<?xml version="1.0" encoding="ISO-8859-1"?><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance">
<front>
<journal-meta>
<journal-id>0798-7269</journal-id>
<journal-title><![CDATA[Zootecnia Tropical]]></journal-title>
<abbrev-journal-title><![CDATA[Zootecnia Trop.]]></abbrev-journal-title>
<issn>0798-7269</issn>
<publisher>
<publisher-name><![CDATA[Instituto Nacional de Investigaciones Agricolas INIA, Maracay, Venezuela. ]]></publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id>S0798-72692014000100002</article-id>
<title-group>
<article-title xml:lang="en"><![CDATA[Impact of grazing on soil organic storage carbon in high lands of Anaime, Tolima, Colombia]]></article-title>
<article-title xml:lang="es"><![CDATA[Impacto del pastoreo en el almacenamiento de carbono orgánico de suelos en el Páramo de Anaime, Tolima, Colombia]]></article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Andrade Castañeda]]></surname>
<given-names><![CDATA[Hernán Jair]]></given-names>
</name>
<xref ref-type="aff" rid="A01"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Espinosa Gómez]]></surname>
<given-names><![CDATA[Edna Lucia]]></given-names>
</name>
<xref ref-type="aff" rid="A02"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Moreno Baltán]]></surname>
<given-names><![CDATA[Herney Armando]]></given-names>
</name>
<xref ref-type="aff" rid="A02"/>
</contrib>
</contrib-group>
<aff id="A01">
<institution><![CDATA[,Universidad del Tolima Facultad de Ingeniería Agronómica ]]></institution>
<addr-line><![CDATA[Ibagué ]]></addr-line>
<country>Colombia</country>
</aff>
<aff id="A02">
<institution><![CDATA[,Universidad del Tolima Facultad de Ingeniería Agronómica Grupo de Investigación PROECUT]]></institution>
<addr-line><![CDATA[ ]]></addr-line>
<country>Colombia</country>
</aff>
<pub-date pub-type="pub">
<day>00</day>
<month>03</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="epub">
<day>00</day>
<month>03</month>
<year>2014</year>
</pub-date>
<volume>32</volume>
<numero>1</numero>
<fpage>7</fpage>
<lpage>21</lpage>
<copyright-statement/>
<copyright-year/>
<self-uri xlink:href="http://ve.scielo.org/scielo.php?script=sci_arttext&amp;pid=S0798-72692014000100002&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://ve.scielo.org/scielo.php?script=sci_abstract&amp;pid=S0798-72692014000100002&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://ve.scielo.org/scielo.php?script=sci_pdf&amp;pid=S0798-72692014000100002&amp;lng=en&amp;nrm=iso"></self-uri><abstract abstract-type="short" xml:lang="en"><p><![CDATA[The mineralization of the soil organic matter may be accelerated by productive activities as grazing and pasture management. The objective of this study was to evaluate the impact of grazing on soil organic carbon (COS) in the high plateau of Anaime, Tolima, Colombia. An unbalanced completely randomized design was used, selecting three treatments: 1) pastures with 20 years in conservation; 2) pastures currently in use; and 3) Andean high-land native forests. The COS storage was estimated using organic carbon concentration and bulk density at a depth of 0-30 cm. Aboveground biomass (AB) of forage plants was also estimated in the case of pastures. Correlation and regression analysis were carried out between COS and time of use, occupation period and stocking rate. The pastures in use stored more COS than those conserved and native forests (P<0.05; 34.4 vs 22.0 vs 21.6 tC/ha, respectively). In contrast, conserved pastures stored more carbon in AB that those in use (P<0.05; 8.3 vs 2.1 tC/ha). It was found that extensive livestock management had a positive impact in the COS in the Páramo of Anaime. However, a change from forests to pastures would cause a CO2 emission by effect of burns and/or decomposition of the organic matter and the loss of other environmental services as water regulation and those derived from conservation of biodiversity.]]></p></abstract>
<abstract abstract-type="short" xml:lang="es"><p><![CDATA[La mineralización de la materia orgánica puede ser acelerada por actividades productivas como el pastoreo y el manejo de pasturas. El objetivo de este estudio fue evaluar el impacto del pastoreo en el almacenamiento de carbono orgánico en suelos (COS) del Páramo de Anaime, Tolima, Colombia. Se realizó un diseño completamente al azar desbalanceado, seleccionando tres tratamientos: 1) pasturas que llevan 20 años en conservación; 2) pasturas actualmente en uso y 3) bosques nativos altoandinos. El almacenamiento de COS se estimó por medio de la concentración de carbono orgánico y la densidad aparente a una profundidad de 0-30 cm. La biomasa arriba del suelo (BAS) de forrajeras también fue estimada en el caso de las pasturas. Se realizaron análisis de correlación y de regresión entre el COS y el tiempo de uso, periodos de ocupación y carga animal. Las pasturas en uso almacenaron más COS que aquellas conservadas y los bosques nativos (P<0,05; 34,4 vs 22,0 vs 21,6 tC/ha, respectivamente). En contraste, las pasturas conservadas almacenaron más carbono en BAS que las pastoreadas (P<0,05; 8,3 vs 2,1 tC/ha). El manejo de ganadería extensiva tiene un impacto positivo en la acumulación de carbono orgánico en el suelo en el páramo de Anaime. No obstante, un cambio de bosques a pasturas causaría emisión de CO2 por efecto de la quema y/o descomposición de la materia orgánica y la pérdida de otros servicios ambientales como la regulación hídrica y aquellos derivados de la conservación de la biodiversidad.]]></p></abstract>
<kwd-group>
<kwd lng="en"><![CDATA[Aboveground biomass]]></kwd>
<kwd lng="en"><![CDATA[bulk density]]></kwd>
<kwd lng="en"><![CDATA[concentration of soil organic carbon]]></kwd>
<kwd lng="en"><![CDATA[stocking rate]]></kwd>
<kwd lng="es"><![CDATA[Biomasa arriba del suelo]]></kwd>
<kwd lng="es"><![CDATA[carga animal]]></kwd>
<kwd lng="es"><![CDATA[concentración de carbono orgánico del suelo]]></kwd>
<kwd lng="es"><![CDATA[densidad aparente]]></kwd>
</kwd-group>
</article-meta>
</front><body><![CDATA[   <font FACE="Verdana">     <p ALIGN="center"><b>Impact of grazing on soil organic storage carbon in high  lands of Anaime, Tolima, Colombia</b></p>     <p ALIGN="center"><b>Impacto del pastoreo en el almacenamiento de carbono  orgánico de suelos en el Páramo de Anaime, Tolima, Colombia</b></p> </font><font FACE="Verdana" size="2">     <p ALIGN="center"><b>Hernán Jair Andrade Castañeda<sup>1</sup>, Edna Lucia  Espinosa Gómez<sup>2</sup> y Herney Armando Moreno Baltán<sup>2</sup></b></p>     <p ALIGN="justify">1 Universidad del Tolima. Facultad de Ingeniería Agronómica.  Grupo de Investigación “Producción Ecoamigable de Cultivos Tropicales”. Ibagué,  Colombia. Correo electrónico: <a href="mailto:hjandrade@ut.edu.co"> hjandrade@ut.edu.co</a></p>     <p ALIGN="justify">2 Universidad del Tolima. Facultad de Ingeniería Agronómica.  Ingenieros Agrónomos, Grupo de Investigación PROECUT. Colombia.</p>     <p ALIGN="justify"><b>ABSTRACT</b></p>     <p ALIGN="justify">The mineralization of the soil organic matter may be  accelerated by productive activities as grazing and pasture management. The  objective of this study was to evaluate the impact of grazing on soil organic  carbon (COS) in the high plateau of Anaime, Tolima, Colombia. An unbalanced  completely randomized design was used, selecting three treatments: 1) pastures  with 20 years in conservation; 2) pastures currently in use; and 3) Andean high-land  native forests. The COS storage was estimated using organic carbon concentration  and bulk density at a depth of 0-30 cm. Aboveground biomass (AB) of forage  plants was also estimated in the case of pastures. Correlation and regression  analysis were carried out between COS and time of use, occupation period and  stocking rate. The pastures in use stored more COS than those conserved and  native forests (P&lt;0.05; 34.4 vs 22.0 vs 21.6 tC/ha, respectively). In contrast,  conserved pastures stored more carbon in AB that those in use (P&lt;0.05; 8.3 vs  2.1 tC/ha). It was found that extensive livestock management had a positive  impact in the COS in the Páramo of Anaime. However, a change from forests to  pastures would cause a CO2 emission by effect of burns and/or decomposition of  the organic matter and the loss of other environmental services as water  regulation and those derived from conservation of biodiversity.</p>     <p ALIGN="justify"><b>Key words</b>: Aboveground biomass, bulk density,  concentration of soil organic carbon, stocking rate.</p>     <p ALIGN="justify"><b>RESUMEN</b></p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">La mineralización de la materia orgánica puede ser acelerada  por actividades productivas como el pastoreo y el manejo de pasturas. El  objetivo de este estudio fue evaluar el impacto del pastoreo en el  almacenamiento de carbono orgánico en suelos (COS) del Páramo de Anaime, Tolima,  Colombia. Se realizó un diseño completamente al azar desbalanceado,  seleccionando tres tratamientos: 1) pasturas que llevan 20 años en conservación;  2) pasturas actualmente en uso y 3) bosques nativos altoandinos. El  almacenamiento de COS se estimó por medio de la concentración de carbono  orgánico y la densidad aparente a una profundidad de 0-30 cm. La biomasa arriba  del suelo (BAS) de forrajeras también fue estimada en el caso de las pasturas.  Se realizaron análisis de correlación y de regresión entre el COS y el tiempo de  uso, periodos de ocupación y carga animal. Las pasturas en uso almacenaron más  COS que aquellas conservadas y los bosques nativos (P&lt;0,05; 34,4 vs 22,0 vs 21,6  tC/ha, respectivamente). En contraste, las pasturas conservadas almacenaron más  carbono en BAS que las pastoreadas (P&lt;0,05; 8,3 vs 2,1 tC/ha). El manejo de  ganadería extensiva tiene un impacto positivo en la acumulación de carbono  orgánico en el suelo en el páramo de Anaime. No obstante, un cambio de bosques a  pasturas causaría emisión de CO2 por efecto de la quema y/o descomposición de la  materia orgánica y la pérdida de otros servicios ambientales como la regulación  hídrica y aquellos derivados de la conservación de la biodiversidad.</p>     <p ALIGN="justify"><b>Palabras clave</b>: Biomasa arriba del suelo, carga  animal, concentración de carbono orgánico del suelo, densidad aparente.</p>     <p ALIGN="justify">Recibido: 21/08/13 Aprobado: 21/11/14</p>     <p ALIGN="justify"><b>INTRODUCTION</b></p>     <p ALIGN="justify">The páramo is an ecosystem with great environmental and  socioeconomical importance, which is threatened by man activities, reducing its  capacity to offer environmental services to the region and World. The mountain  regions in the Andean region have been identified as vulnerable to climate  change (Büchler et al., 2004). Otero et al. (2011) argued that Andean páramo and  cloud forest ecosystems in South America are changing to agricultural activities  such as potato farming and cattle grazing.</p>     <p ALIGN="justify">The páramo provides invaluable ecosystem services as constant  supply of water and storage of soil organic carbon (SOC), that are dependent of  soil behavior (Hofstede and Mena, 2011).The soils in páramo ecosystem have a  high content of organic matter and can store, by a long period, organic carbon  in this component, due to low temperature decreases mineralization rates (Hofstede  and Mena, 2011). However, soil land activities, such as grazing and fires, can  accelerate organic matter decomposition and decrease storage of SOC (Maia et  al., 2009; Román-Cuesta et al., 2011). On the other hand, several studies in  high Andean mountains of Colombia and Venezuela have not shown an increase in  the total soil carbon during the fallow period (Sarmiento and Bottner, 2002).  Some authors have found a negative impact of livestock activities in SOC storage  (Han et al., 2008; Luan et al., 2014; Silva et al., 2014); while others have  argued that a good management can contribute to increase carbon sinks (De  Camargo et al., 1999; Fisher et al., 2004; Andrade et al., 2008; Maia et al.,  2009; Wang et al., 2014).</p>     <p ALIGN="justify">The government of Colombia has supported the research in high  land ecosystems to understand the impacts of climate change and land use on  cycles of water and carbon (Brown et al., 2007). The relevance of these  terrestrial ecosystems is obvious, given the importance of these carbon  reservoirs in páramo soils, thus the objective of this study was to estimate the  impact of grazing on storage of SOC and its potential emission of carbon dioxide  towards the atmosphere in this region.</p>     <p ALIGN="justify"><b>MATERIALS AND METHODS</b></p>     <p ALIGN="justify"><b>Study area</b></p>     <p ALIGN="justify">This study was conducted in the Reserva Natural Semillas de  Agua and adjacent grasslands, located on east face of Central Mountain Range of  Colombia, in the municipalities of Cajamarca, Rovira and Roncesvalles, Tolima,  Colombia. This reserve, which covers 979 ha, is composed by páramo ecosystems,  páramo native forests, Andean high land forests and grasslands and is dedicated  to conservation and research (Birdlife, 2011).</p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">The mean annual temperature is 7.3oC, with a minimal and  maximal temperature of -3 and 25oC, respectively. The mean annual  evapotranspiration is 680 mm and a vertical rainfall of 2141 mm/year. The  rainfall shows a bimodal behavior with a dry season in January, February, March,  September and December; whilst rainy season covers April, May, July, October and  November. Strong winds are presented in August, September and January, with  predominant direction southeast to northeast (Asociación Red Colombiana de  Reservas Naturales de la Sociedad Civil, 2011).</p>     <p ALIGN="justify">The soils in the study area are derived from volcanic ashes,  slags and vitreous pyroclastics with a thick layer of organic matter little  weathered, which is typical of páramos. Experimental plots were located in two  cartographic units: MDBf and MGCe (IGAC, 2004). The unit MDBf is dominated by  pyroclastics, andesite and the metamorphic rocks (shale). This soil association  is composed by soils Lithic Cryorthentsen in a 60% and Rocky outcrops in 40% of  area. These soils have the following characteristics: a low evolution shallow  due to hard rock, high content of organic matter, highly acid, high cationic  exchangeable capacity, poor in changeable bases, low in available phosphorus and  very low fertility. Steep slopes (50-75%), low temperatures and low soil  effective depth are the main limiting characteristics for use.</p>     <p ALIGN="justify">The unit MGCe has volcanic ashes as parental material, which  is deposited on andesite. Soils of this consociation are represented by Typic  Haplundands in a 90% and inclusions of Humic Udivitrands in a 10% of area. These  soils are characterized by high depth, with black color due to accumulation of  organic matter, medium tomoderately gross textures, well-structured and well-drained.  These soils have a high cationic exchangeable capacity, high phosphorus content  in top layers and moderate fertility. Slopes range between 25 and 50% in the  area.</p>     <p ALIGN="justify"><b>Selection of systems to evaluate</b></p>     <p ALIGN="justify">An unbalanced completely randomized design was used,  selecting three treatments: 1) pastures with 20 years in conservation (PC); 2)  pastures currently in use (PU); and 3) Andean high-land native forests (F).  Selection of pastures was carried out with the advisory of local personal with  knowledge of this zone. Producers were asked about time of use of pastures, mean  stocking rate and periods of resting and occupation.</p>     <p ALIGN="justify">A total of 18 experimental units were evaluated: four PC in  the conservation area of this natural reserve, 11 PU in adjacent farms and three  F. This last system was considered as control to estimate the impact of land use  change in storage of SOC. Each experimental unit was geographically located to  identify the type of soil. Statistical analyses were carried out to define a  significant effect of soil unit in results.</p>     <p ALIGN="justify"><b>Estimation of storage of soil organic carbon</b></p>     <p ALIGN="justify">Estimation of SOC was done to a depth of 0-30 cm, employing  bulk density (BD), concentration of SOC and portion of gross fragments (rocks;  Don et al., 2007; Maia et al., 2009). To avoid overestimations of SOC, it was  applied a correction of the soil weight according to BD following the approach  recommended by Ellert et al. (2002) and Buurman et al. (2004). The treatment  with lowest BD was taken as reference, assuming this variable was similar in all  experimental plots before land use change.</p>     <p ALIGN="justify">One combined soil sample per plot was obtained by mixing ten  sub-samples collected with a manual auger in each experimental plot.  Concentration of SOC was estimated by method of Walkley and Black (1934) in the  Laboratorio Laserex of the Universidad del Tolima. BD was estimated using the  cylinder approach (Andrade and Ibrahim, 2003), taking three samples by  experimental unit with a cylinder of 100.13 cm3. In case of soils with a high  content of rocks, gross-fragment content was calculated doing a soil pit of 50 x  50 x 50 cm and estimating volume of rocks by water displacement.</p>     <p ALIGN="justify"><b>Estimation of herbaceous biomass</b></p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">Aboveground biomass of grasses was estimated using the  methodology proposed by Andrade and Ibrahim (2003), taking 10 quadrats of 50 x  50 cm, which were randomly located in each experimental plot. In these quadrats,  all aboveground biomass of grasses was cut and weighted in fresh, taking a  subsample of around 200 g to estimate dry matter content (65ºC until constant  weight). Biomass was transformed to carbon multiplying by a carbon fraction of  0.5, which is recommended by IPCC (2007).</p>     <p ALIGN="justify"><b>Statistical analyses</b></p>     <p ALIGN="justify">An analysis of variance was carried out to those normal  variables. Variable with no normality, such as concentration of SOC (%), were  transformed. In case of found statistical differences among treatments, a  comparison test of Tukey was executed. Analysis of correlation and lineal  regression, with logarithmic transformation and untransformed variables, between  storage of SOC and BD, and management factors such as annual stocking rate,  occupation period and time of use were carried out. To evaluate the effect of  soils in response variables, it was developed an analyses of variance taking  soil units as classification variables. All statistical analyses were carried  out using the software InfoStat.</p>     <p ALIGN="justify"><b>RESULTS AND DISCUSSION</b></p>     <p ALIGN="justify"><b>Concentration of soil organic carbon</b></p>     <p ALIGN="justify">No significant difference (P&gt;0.05) was found in concentration  of SOC in the top 30 cm among land use systems. However, the lowest  concentration of SOC was registered in native forests (2.9%), which was  statistically similar (P&gt;0.05) to those in conserved pastures, but a 37% lower  (P&lt;0.05) than in pastures in use (<a href="#fig1">Figure 1a</a>).</p>     <p ALIGN="center"><a name="fig1"> <img border="0" src="/img/fbpe/zt/v32n1/art02fig1.gif" width="429" height="737"></a></p>     
<p ALIGN="justify">Bulk density BD was not significantly affected (P&gt;0.05) by  type of soil, ranging between 0.46 and 0.63 g/ cm³. It was found that soils in  native forests had a lower BD (0.46 g/cm³), followed by conserved pastures and  those in use where this variable has increased in 9 and 37%, respectively (<a href="#fig1">Figure  1b</a>).</p>     <p ALIGN="justify">It was found a decreasing lineal relationship between BD and  concentration of SOC in pastures in use, indicating an increase of SOC decreases  of this first variable. In spite this relationship is just explained in a 0.5% (<a href="#fig2">Figure  2a</a>). On the contrary, an increasing lineal relationship between BD and  concentration of SOC, when all land use systems were included, was found with an  explication of 12% (<a href="#fig2">Figure 2b</a>).</p>     <p ALIGN="center"><a name="fig2"> <img border="0" src="/img/fbpe/zt/v32n1/art02fig2.gif" width="431" height="744"></a></p>     
]]></body>
<body><![CDATA[<p ALIGN="justify"><b>Impact of grazing on soil bulk density</b></p>     <p ALIGN="justify">Stocking rate had a positive impact on BD (<a href="#fig3">Figure  3a</a>), in the studied range (0 – 0.19 animals/ha/ year), which are so low due  to own characteristics in the zone. In the same way, an increase in stocking  rate of 1 animal/ha/year causes an increase of BD in an average of 1.1 g/cm<sup>3</sup>  (<a href="#fig3">Figure 3a</a>). The period of occupation affected in a very  similar way than stocking rate, to BD (<a href="#fig3">Figure 3b</a>),  increasing it a mean of 0.1 g/cm<sup>3</sup> every 100 days of grazing.  Similarly, the time of use of pastures increases BD in 0.03 g/cm<sup>3</sup>  every 10 years of use (<a href="#fig3">Figure 3c</a>). These predictive models  explain a total of 21-23% the variation of BD.</p>     <p ALIGN="center"><a name="fig3"> <img border="0" src="/img/fbpe/zt/v32n1/art02fig3.gif" width="391" height="958"></a></p>     
<p ALIGN="justify"><b>Storage of soil organic carbon</b></p>     <p ALIGN="justify">Pastures in use had a significantly highest (P&lt;0.05) SOC in  top 30 cm than that in conserved grasslands and high land native forests (34.4 ±  3.4 vs 22.0 ± 2.1 vs 21.6 ± 3.8 tC/ha, respectively; <a href="#fig4">Figure 4</a>).</p>     <p ALIGN="center"><a name="fig4"> <img border="0" src="/img/fbpe/zt/v32n1/art02fig4.gif" width="541" height="569"></a></p>     
<p ALIGN="justify"><b>Impact of grazing on storage of soil organic carbon</b></p>     <p ALIGN="justify">A quadratic relationship between storage of SOC and animal  stocking rate and period of occupation (<a href="#fig5">Figures 5a and 5b</a>)  was detected. Accumulation of SOC is maximized when annual stocking rate is 0.1  animals/ha and 115 days of occupation/ year (<a href="#fig5">Figures 5a and 5b</a>).  These models have a high explanation of storage of SOC in these ecosystems  (77-78%). In contrast, time of use of pastures always increased SOC storage at a  mean rate of 0.18 tC/ha/year (<a href="#fig5">Figure 5c</a>).</p>     <p ALIGN="center"><a name="fig5"> <img border="0" src="/img/fbpe/zt/v32n1/art02fig5.gif" width="399" height="960"></a></p>     
<p ALIGN="justify">Carbon storage in aboveground biomass Statistical differences  (P&lt;0.05) in accumulation of aboveground biomass of herbaceous species between  conserved and in-use pastures (8.3 ± 0.3 vs 2.1 ± 0.9 tC/ha, respectively). This  can be seen in <a href="#fig4">Figure 4</a>, with an estimation of aboveground  biomass of high-land native forests reported by other authors.</p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">Total carbon storage in pastures and native high-land forests  Taking information about carbon storage in aboveground biomass in high-land  forests (Gálmez and Kómetter, 2009), it is possible to show a tendency in  finding more carbon accumulation, as expected, in native forests than in in-use  and conserved pastures (65.8, 36.6 and 30.3 tC/ha, respectively; <a href="#fig4"> Figure 4</a>). In native forests, most of this carbon is located in aboveground  biomass; whereas in pastures this is stored in soils (<a href="#fig4">Figure 4</a>).</p>     <p ALIGN="justify">Concentration of soil organic carbon The SOC concentration in  pastures in conservation (2.9%) was lower than that reported by Farley et al.  (2004), who found a value of 4.2% in páramo soils in the equadorian Andean  region at a depth of 0-10 cm. The results of this study and quoted one are  coinciding when inuse pastures are considered, despite of slight differences in  the studied soil depth. However, Han et al. (2008) found that SOC decreased with  grazing intensity at the same depth of this study.</p>     <p ALIGN="justify">The positive impact of extensive livestock in the páramo on  concentration of SOC may be explained by an increase of biomass of fine roots  which at death and decomposition supply carbon at soil (FAO, 2007; Andrade et  al., 2008; Maia et al., 2009). It is considered that in forests and conserved  pastures (with no grazing for more than 20 years) there is not an important  contribution of carbon by fine root senescence. Similarly, necromass due to fine  roots can be maintained by a relative long time due to low temperatures in the  zone, which decline decomposition rate and mineralization of organic matter.  According to Pansu et al. (2009), rhizodeposition provides a force for microbial-mediated  process, including soil carbon sequestration. This analysis is important to  understand the dynamics of SOC in Latin America, due to the change from forests  to pastures is one of the most important tendencies in the region (Harvey et  al., 2005). </p>     <p ALIGN="justify"><b>Bulk density</b></p>     <p ALIGN="justify">The BD found in this study (0.46 – 0.63 g/ cm3) were much  lower than those reported by Farley et al. (2004) in top 10 cm in pastures in  ecuadorian Andean region (1.17 g/cm3). This clearly indicates an effect of soil  genesis in BD and show the importance of considering the soil type when studying  the impact of land use and management practices in this soil physical variable.</p>     <p ALIGN="justify">The increment of BD in soils is a clear and widely studied  process. Pressure of cattle hooves causes compaction, mainly when previous land  use was native forests. These changes were not significant in this study, due to  the nature of soil, mainly by its origin from volcanic ashes. In a research  developed by Sadeghian et al. (1999), it is reported that in spite of finding  significant changes in soil compaction by animal trampling, the values of BD  were not so great due to its parental material of volcanic nature (andisols with  low bulk and real densities), its textural classes, dominated by sand and high  content of organic matter. Similar results were found by Medina- Roldán et al.,  2012, who reported an increase of soil BD by grazing effect in comparison to  grazing exclusion (0.18 vs 0.14 g/cm3, respectively). In contrast, Larreguy et  al. (2014) found the lowest BD in moderate grazing disturbance in comparison to  low and high grazing disturbance (1,08 vs 1,15 vs 1,23 g/cm3, respectively) in  arid soils of Argentinian Patagonia.</p>     <p ALIGN="justify">BD in currently in-use pastures is being affected by two  simultaneous, but opposite factors: grazing and addition of organic matter. The  last one is derived from the senescence of fine roots caused by grass  defoliation (FAO, 2007; Andrade et al., 2008; Maia et al., 2009). Pressure of  hooves of animals has a negative impact on soil due to an increase of BD as a  consequence of reduction of porosity. This is confirmed by the study of Alegre  and Lara (1991), who found a rise in soil BD by the effect of animals grazing.  In the same way, in the Páramo of Chontales (Boyacá), Avellaneda (2006) claimed  that grazing caused a hardening and impermeability of soils, affecting water  movement and, therefore, altering the hydrologic cycle.</p>     <p ALIGN="justify">Grazing is contributing to improve the storage of organic  matter (OM) in soil, which would improve the soil structure. According to  Fassbender and Bornemisza (1987), OM favor the formation of individual  aggregates, reduce soil global aggregation, improve water infiltration,  intensify soil aeration, contributes to better water distribution in soil  profile and augment water retention capacity. Civeira (2010) also found that  soil physical properties improve with addition of compost: an increase of OM  which reduces BD and rises infiltration. Sadeghian et al. (1999) found, in the  Department of Quindío (Colombia), than organic matter has significantly  contributed to reduce BD and helped to improve porosity and water retention,  additionally to increase structural stability.</p>     <p ALIGN="justify">The low explication (0.5%) of SOC concentration by BD  indicates that other factors, such as grazing are affecting carbon sequestration  in soils. It was observed a non-expected relationship; in other words, when  concentration of SOC is increased BD is also raised, with an explanation of 12%.  This confirm that addition of organic matter is not having a positive effect on  reducing BD, on the contrary, it was observed a tendency in rising BD by  livestock activity. It must be clarify that stocking rates were low (0.01 – 0.19  animals/ha/ year), due to extensive management of livestock in the study area.  That is to say that the evaluated stocking rate is not enough to promote a  change in BD. Growth rate of grasslands is so slow, as a consequence of low  temperatures; consequently, resting periods are so long in order to allow the  recovery of grasses (between 183 and 329 days/ year). As a result of this, the  occupation periods are relatively short, ranging between 36 and 205 days/year.  This extensive use does not cause a significant impact on BD.</p>     <p ALIGN="justify">No variation in BD can also be explained by the addition of  organic matter from grazing, which increases the capacity of soil to hold high  stocking rates (Martínez et al., 2008). In other words, functional resilience of  soil pores is closely related to presence of organic components (Dörner et al.,  2009), that indicates that under the same management conditions a soil is  enriched with organic matter this should increase its capacity to maintain  stocking rates in comparison to soils with lower organic matter (Martínez et  al., 2008). Results of this study coincides with findings of Alarcón et al.  (2010), where it was found that BD and total porosity of soil were not  significantly affected by increase of grazing intensity, demonstrating the soil  capacity to endure animal transit and trampling with no a significant  deformation of soil volume.</p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">A significant change in BD would have to be the result of  intensification of livestock activity, such as reports by Sadeghian et al.  (1999), who found that magnitude of diminishing of pore spaces found were  greater with an increase of the number of animals by area. In the same way, it  was reported that trampling of animals in 2 to 3 years in intensive systems of  meat and milk production caused a similar compaction to that extensive systems  during more than 15 years.</p>     <p ALIGN="justify"><b>Soil organic carbon</b></p>     <p ALIGN="justify">The extensive livestock has a positive impact in storage of  soil organic carbon in this zone, allowing the capture of 12.5 tC/ha in  comparison with pastures in conservation by more than 20 years. In other words,  the use of extensive cattle could be causing a SOC fixation of around 0.6 tC/ha/year.  This response can be explained by a potential higher biomass and extension of  grasses fine roots, mainly in top soil, than trees as found by Harden et al.  (2013) in an Ecuadorian páramo. Stockmann et al. (2013) argued that improved  grazing with best management practices could fix carbon at a rate of 0.4-0.8 tC/ha/year,  mainly in the top 30 cm of soil, the zone with the highest root and microbial  activity and plant inputs. The effect of grazing intensities is contrasting  among different studies. For example, Silva et al. (2014) found that low and  moderate grazing intensities had lower soil organic carbon losses than high  grazing intensity in Southern Brazil. Luan et al. (2014) reported that grazing  exclosures of five years significantly increased SOC stock in alpine ecosystems  in east Qinghai–Tibet Plateau; which contrasts with Wang et al. (2014) who found  a reduction in carbon content in the topsoil in Loess Plateau (China) after  grazing exclusion.</p>     <p ALIGN="justify">These results agree with those reported by Fisher et al.  (2004) in the Colombian Orinoquia and by Ibrahim et al. (2007) in Colombia and  Costa Rica, who found that forests store lower carbon in soils than improved  grasslands (Brachiaria humidicola). De Camargo et al. (1999) found similar  results on Brazilian Amazon, where land uses of improved pastures (B. humidicola)  incorporate more carbon than forests of forestry plantations. Maia et al. (2009)  found in oxisols in Rondônia and Mato Grosso, Brazil than degraded pastures  caused a reduction of SOC in 0.27-0.28 tC/ha/year, contrasting with wellmanaged  and improved pastures (0.61 – 0.72 tC/ ha/year). In contrast, Zimmermann et al.  (2010) did not find statistical differences in SOC stocks among forests,  shrublands and grasslands in the Peruvian Andes (11.8 vs 14.7 vs 11.9 kgC/m<sup>2</sup>,  respectively).</p>     <p ALIGN="justify">The previous statement does not mean that the ideal situation  is land use change, from native forests to grasslands, due to it is only taking  into account the SOC storage. The global cycle of carbon must consider the  carbon accumulation in biomass in the forest ecosystems. In the same way, any  possible land use change must take into account the functions of protection and  offer of environmental services, such as water regulation and soil conservation  at protect it from erosive effects of water and air (FAO, 2010).</p>     <p ALIGN="justify">Similarly, it must be highlighted the negative and opposite  impact of livestock in other nonevaluated issues, such as the loss of water  sources and water quality, decrease in water regulation, loss of natural  ecosystems, decrease of biological diversity, erosion and mass removals (Murgueitio,  2003).</p>     <p ALIGN="justify">The results suggest that extensive livestock in the zone is  contributing to increase carbon reservoirs in soils; however, changes in  management practices, such as an increase in livestock intensification, could  cause a negative impact in carbon storage in the future, generating an CO<sub>2</sub>  emission to atmosphere. Grazing intensity not affect COS, according to  Medina-Roldán et al. (2012) in an upland grassland in England and Silveira et  al. (2013) in perennial grass pastures in the southeastern USA. It means that if  the resting periods, where grasses can recovery, are not taken and unsustainable  practices are used in intensive livestock a negative impact could be caused,  transforming these ecosystems from sinks to sources of carbon.</p>     <p ALIGN="justify">This study suggests that the management of these pastures be  extensive, in a natural way without the use of chemical-synthesis fertilization;  avoid seasonal fires and elimination of vegetal succession by chemical methods.  Unsustainable practices, such as the use of concentrates, conduce to an  intensification of cattle production which causes a negative environmental  impact much higher than current systems. Murgueitio (2003) found that in  intensive production systems of milk and fattening steers with high stocking  rates, a reduction of diversity of plant species and soil fauna was produced.</p>     <p ALIGN="justify">The importance of páramo zones for water conservation and  regulation must not be forgotten. Herrera (2011) argued that the “capacity of  páramo in the storing and distribution of water from rains and cloud  condensation is based on their special soil structure, safeguarded by vegetation”.</p>     <p ALIGN="justify"><b>Carbon storage in aboveground biomass</b></p>     ]]></body>
<body><![CDATA[<p ALIGN="justify">The reduction of plant cover, due to change from native  forests to grasslands, and the current use of pastures does not have a negative  effect in carbon storage in soil in the Páramo of Anaime. Medina-Roldán et al.  (2012) found similar results in upland grasslands of England, no detecting the  impact of seven years of grazing exclusion on total carbon in soil surface.  Moreover, soil loss from water erosion, with a consequent carbon loss, can be  also caused. The highest carbon accumulation in forests is caused by its  greatest carbon in aboveground biomass (44.2 tC/ha, Gálmez and Kómetter, 2009)  counteracting the low SOC storage. In other words, despite of the use in pasture  causes a higher SOC fixation, when a change in land use from forests to pastures  is presented a loss of carbon which is not compensated by an increase in SOC.  The land use change from forests to grasslands could cause an emission of 118  tCO<sub>2</sub>/ha.</p>     <p ALIGN="justify">Although conserved pastures store more carbon in aboveground  biomass, those in-use pastures contain around 6.3 tC/ha more than first ones due  to a greater carbon accumulation in soils. That is to say that pastures  currently in use have caused a total carbon fixation of 1.2 tCO<sub>2</sub>/ha/  year over than those in conservation.</p>     <p ALIGN="justify">Taking in consideration the importance of páramo ecosystems  and theirs vulnerability to climate change and anthropogenic activities, impacts  of livestock activities in the generation of environmental services and  conservation of these ecosystems must be evaluated. With this information, it  can be evaluated and recommended if it is better to conserve pasture, reducing  SOC but increasing carbon in biomass in order to protect this ecosystem from  degradation.</p>     <p ALIGN="justify"><b>CONCLUSIONS</b></p>     <p ALIGN="justify">Conservation of pastures does not contribute positively to  SOC storage. On the contrary, the extensive cattle activity, which is  characteristic of this zone, has a positive impact in the carbon storage in soil  in the Páramo of Anaime, fixing about 1.2 tCO<sub>2</sub>/ha/year more than  those in conservation. This is possible due to the higher dynamics of fine roots  by defoliation in grazing lands, which conduces to incorporation of organic  carbon to soil. This carbon can be stored by a long time due to low temperatures  reduce considerably mineralization of organic matter. In contrast, conservation  of grasslands increases carbon retention in aboveground biomass, resulting in  similar total carbon in conserved and in-use grasslands. However, more research  is needed to find more details about the SOC dynamics in tropical upland  grasslands.</p>     <p ALIGN="justify">Land use change, from forests to pastures, has a positive  impact in the SOC storage. However, in this change must be considered carbon  capture in biomass and other environmental services, such as those derived from  conservation of biodiversity, soil protection and water regulation. Moreover, it  is widely known that conventional cattle activity has negative effects on offer  of invaluable environmental services that are supplied by páramo zones.</p>     <p ALIGN="justify">Although a rising tendency in BD caused by cattle activity,  this relationship was not significant. This can be caused by a higher effect of  parental material, low stocking rate, long resting periods, short occupation  periods and an increase of incorporation of organic matter due to grazing, which  improves soil structure. An increase in stocking rate and occupation period has  a negative effect in SOC storage; in contrast, a higher time of use increases  SOC. This can be explained by soil parent material, low animal stocking rates,  long resting periods, short occupation time and the organic matter of livestock  activities at low level of intensification. An increase in annual stocking rate  and occupation periods has a negative effect in SOC stocks; in contrast, an  increase in time of use of grasslands causes that SOC stocks rises. According to  this, it is needed to maintain sustainable and ecofriendly practices and avoid  intensification of this activity.</p>     <p ALIGN="justify"><b>ACKNOWLEDGMENTS</b></p>     <p ALIGN="justify">The authors of this paper want to express their most deep  gratitude to Semillas de Agua for the invaluable support assistance in  logistical issues and plot selection from local producers. The authors  acknowledge to the town hall of Cajamarca for financial support. We sincerely  appreciate to all cattle producers for their support in this research. Thanks to  Diana Skarly Canal for her support in the Laboratory of Plant Physiology at the  University of Tolima.</p>     <p ALIGN="justify"><b>LITERATURE REVIEW</b></p>     ]]></body>
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