Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

Aboveground biomass in a post-mining forest succession

in the Colombian Pacific

Jhon Jerley Torres-Torres*1; https://orcid.org/0000-0002-0503-837X

Harley Quinto-Mosquera2; https://orcid.org/0000-0001-5989-4334

Mayira Guerrero-Machado3; https://orcid.org/0009-0003-0411-0493

1. Maestría en Bosques y Conservación Ambiental, Universidad Nacional de Colombia, Colombia, Cra. 65 #59a-110;

jhtorrest@unal.edu.co (*Correspondence)

2. Universidad Tecnológica del Chocó, Programa de Biología. Facultad de Ciencias Naturales, Quibdó, Colombia, 22

N°18B-10; hquintom@gmail.com

3. Fundación Biodiversidad, Cambio Climático y Bienestar Social, Quibdó, Colombia; mayiraguerrero@hotmail.com

Received 17-I-2024. Corrected 07-V-2024. Accepted 12-VIII-2024.

ABSTRACT

Introduction: Mining is one of the main drivers of deforestation of tropical forests. This activity affects the stor-

age of aboveground biomass of these ecosystems; therefore, their ability to contribute to the mitigation of global

climate change.

Objective: To assess the influence of soil properties on the aboveground biomass storage of post-mining forests

in the Colombian Pacific.

Methods: Plots were established in areas post-mining and with different successional ages (12-15 years, 30-35

years, and mature forest). The aboveground biomass and physicochemical parameters of the soil were measured.

Results: An aboveground biomass of 15.58 t ha-1, 35.17 t ha-1, and 178.32 t ha-1 was recorded at 12-15 years,

30-35 years, and mature forests, respectively. The species with the highest biomass content in post-mining for-

ests were Cespedesia spathulata and Clidemia septuplinervia. The aboveground biomass was positively correlated

with organic matter (OM), calcium (Ca), magnesium (Mg), CICE, total nitrogen (N), and silt. In contrast, the

relationship was negative with sand, aluminum (Al), and potassium (K) content. It was evidenced that the rela-

tionship between aboveground biomass and soils differed in each successional age. When evaluating the changes

of aboveground biomass and soils in the succession, it was observed that the aboveground biomass and total N

increased with the recovery time. At the same time, the P and K decreased with succession. On the other hand,

the contents of OM, Mg, Al, Ca, and CICE showed curvilinear tendencies since they increased in the first stages

and then decreased in the advanced successional stages.

Conclusions: Aboveground biomass increases with forest recovery time in the study area. This increase is influ-

enced by the presence of two dominant species shared among the investigated ecosystems and by the soil’s N, P,

and K content.

Key words: aluminum toxicity; carbon; Chocó biogeographical; nutrient limitation; restoration; plant succession.

RESUMEN

Biomasa aérea en una sucesión de bosques post-minería del Pacífico colombiano

Introducción: La minería es una de las principales causas de deforestación de los bosques tropicales. Esta activi-

dad afecta el almacenamiento de biomasa aérea de estos ecosistemas; y, por tanto, su capacidad para contribuir a

la mitigación del cambio climático global.

https://doi.org/10.15517/rev.biol.trop..v72i1.55276

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

INTRODUCTION

Tropical forests are crucial in mitigating

global climate change (Pachauri et al., 2014), as

they store approximately 59 % of the total car-

bon in terrestrial ecosystems. Additionally, due

to their high net primary productivity, these

forests capture an estimated 30 % of the atmo-

spheric carbon dioxide annually, significantly

contributing to carbon sequestration world-

wide (Yguel et al., 2019). These ecosystems are

estimated to store 470 billion tons of CO2 in

aboveground and belowground biomass (Pan

et al., 2011; Pugh et al., 2011). Despite their

importance, tropical forests have faced sig-

nificant deforestation and degradation in recent

decades, resulting in net losses of 3.9 million

hectares in Africa and 2.6 million hectares in

South America over the last decade (FAO &

PNUMA, 2020). Anthropogenic activities, par-

ticularly mining, have been the primary drivers

of these alarming deforestation rates (Primack

et al., 2019; FAO & PNUMA, 2020).

In particular, open-pit mining of metals,

such as gold, has become one of the main driv-

ers of deforestation and degradation of tropical

forests (Kalamandeen et al., 2020; Primack et

al., 2019). It is estimated that between 2001

and 2013, about 1 680 km2 of tropical forests

in South America were lost (Primack et al.,

2019), of which 41 % corresponds to mining

carried out in Guyana, 28 % in the Southwest

of the Amazon, 11 % in the Tapajos Xingú for-

ests in Brazil, and 9 % in the Magdalena basin

in Colombia (Alvarez-Berríos & Aide, 2015).

Between 2010 and 2017, it is estimated that

between 57 000 and 60 000 ha of natural forests

were lost to gold mining in Guyana and Peru,

respectively (Kalamandeen et al., 2020), which

not only affects the biodiversity of the region

threatened but also affects the carbon content

stored and the capacity of ecosystems to miti-

gate global climate change (Quinto et al., 2013).

Open pit mining affects the functioning

of the ecosystem because it causes deforesta-

tion of forests, edaphic erosion, changes in the

physicochemical conditions of the soil, chemi-

cal contamination by substances such as mer-

cury, excavation of the subsoil, sedimentation

of rivers and streams, loss of OM, changes in

nutrient content, alteration of biogeochemical

cycles, decrease in forest biomass and carbon,

changes in species composition, and biodiver-

sity loss (Kalamandeen et al., 2020; Quinto et

al., 2013; Ramírez et al., 2019). However, due to

the enormous carbon emissions into the atmo-

sphere generated by this economic activity, it is

Objetivo: Evaluar la influencia de las propiedades del suelo en el almacenamiento de la biomasa aérea de bosques

post-minería del Pacífico colombiano.

Métodos: Se establecieron parcelas en áreas post-minería con diferentes edades de sucesión (12-15 años, 30-35

años y bosque maduro). Se midió la biomasa aérea y parámetros fisicoquímicos del suelo.

Resultados: Se registró una biomasa aérea de 15.58 t ha-1, 35.17 t ha-1 y 178.32 t ha-1 en 12-15 años, 30-35 años

y bosque maduro, respectivamente. Las especies con mayor contenido de biomasa en los bosques post-minería

fueron Cespedesia spathulata y Clidemia septuplinervia. La biomasa aérea se correlacionó positivamente con

la materia orgánica (MO), calcio (Ca), magnesio (Mg), CICE, nitrógeno total (N) y limo. Por el contrario, la

relación fue negativa con el contenido de arena, aluminio (Al) y potasio (K). Se evidenció que la relación entre

la biomasa aérea y los suelos difería en cada edad sucesional. Al evaluar los cambios de la biomasa aérea y los

suelos en la sucesión, se observó que la biomasa aérea y el N total aumentaron con el tiempo de recuperación.

Al mismo tiempo, el P y el K disminuyeron con la sucesión. Por otro lado, los contenidos de OM, Mg, Al, Ca, y

CICE mostraron tendencias curvilíneas ya que aumentaron en los primeros estadios y luego disminuyeron en los

estadios sucesionales avanzados.

Conclusiones: la biomasa aérea aumenta con el tiempo de recuperación del bosque en el área de estudio. Este

incremento está influenciado por la presencia de dos especies dominantes compartidas entre los ecosistemas

investigados y por el contenido de N, P y K del suelo.

Palabras clave: toxicidad de aluminio; carbono; Chocó biogeográfico; limitación de nutrientes; restauración;

sucesión vegetal.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

considered to reduce the climate change mitiga-

tion potential of tropical forests substantially

(Kalamandeen et al., 2020; Poorter et al., 2016).

Therefore, to recover the functionality and role

of the post-mining ecosystems in the carbon

balance and the mitigation of global warming,

it is necessary to evaluate the aboveground

biomass storage and the environmental factors

that favor it.

The biomass recovery in areas degraded

by anthropogenic activities (mining, logging,

livestock, and agriculture) is conditioned by

factors such as climate, water availability (pre-

cipitation), recovery time, type and disturbance

intensity, surrounding forest cover, soil condi-

tions and fertility, among others (Guariguata

& Ostertag, 2001; Kalamandeen et al., 2020;

Oberleitne et al., 2021; Poorter et al., 2016).

Due to this, secondary forests with 20 years of

regeneration (without significant soil modifi-

cation) present aboveground biomass ranges

between 20 and 225 tons per hectare (t ha-1),

with 122 t ha-1 on average (Poorter et al., 2016).

Likewise, these areas can reach up to 90 % of

the aboveground biomass of a primary forest

in an average time of 66 years (Poorter et al.,

2016). In areas affected by mining, the recovery

of aboveground biomass is limited by soil nutri-

ents, especially N (Kalamandeen et al., 2020).

Therefore, it is evident that these factors are

essential in forest recovery.

Although few studies directly link aboveg-

round biomass with soil nutrients in post-min-

ing areas, some research results in ecosystems

under ecological succession (other than post-

mining) indicate that as aerial biomass and

succession increase, soil nitrogen (N) tends to

increase. In contrast, phosphorus (P) tends to

decrease (Feldpausch et al., 2004). This obser-

vation aligns with the hypothesis of nutritional

limitation of available P and total N with suc-

cession (Walker & Syers, 1976). According

to this hypothesis, tropical soils in early suc-

cessional stages have limited availability of N

(Davidson et al., 2004). However, as ecosystem

biomass and succession progress, the availabil-

ity of N increases (Davidson et al., 2004; Reed

et al., 2011). On the other hand, P levels tend

to be high in the initial successional stages,

but over time, its availability diminishes and

becomes limiting in the ecosystem (Reed et al.,

2011; Vitousek et al., 1993; Vitousek et al., 2010;

Walker & Syers, 1976). Nevertheless, these pat-

terns and hypotheses have yet to be tested in

post-mining areas.

The Colombian Pacific is one of the raini-

est regions in the world, with places that have

rainfall levels higher than 10 000 mm per year

(Poveda et al., 2004). In this region, open pit

gold mining generates the deforestation and

degradation of more than 360 ha of forest annu-

ally (Ramírez & Ledezma, 2007). Due to this,

these ecosystems offer us an opportunity to

evaluate the hypotheses above about nutritional

limitations. Therefore, our objective was to

evaluate the influence of soil properties on the

aboveground biomass storage of post-mining

forests in the Colombian Pacific. The follow-

ing questions guided this objective: How does

aboveground biomass vary as a function of suc-

cession time in areas post-mining? Which tree

species and botanical families have the highest

aboveground biomass in areas post-mining? To

what extent do edaphic conditions, especially

soil nutrients, determine the aboveground bio-

mass of areas post-mining in these high-rainfall

tropical ecosystems? This inquiry is especially

pertinent considering that, as noted by Austin

& Vitousek (1998), high precipitation levels can

lead to decreased nutrient content due to runoff

and leaching.

MATERIALS AND METHODS

Study Area: The present study was carried

out in forested areas previously degraded by

open-pit gold mining in the town of Jigualito

(5º06’01” N & 76º32’44” W), municipality of

Condoto, in the Colombian Pacific, which has

an average rainfall of 8 000 mm per year, an

altitude of 70 m and flat topography. This local-

ity is part of the Chocó biogeographical Chocó

subregion, which includes the upper basins of

the Atrato and San Juan rivers, in Piedemonte

and Colinas low landscape units with humid

terraced soils and with a type of transitional

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

sedimentary rock (Poveda et al., 2004). The

forests are primarily secondary, with different

recovery ages, because mining has been carried

out in the area at different times.

The soils are ultisols, but due to mining,

they were characterized by a lot of rocky mate-

rial and sand. In addition, they are acidic and

have high contents of OM, total N, available P,

Al, and clay. At the same time, the concentra-

tions of Ca, K, Mg, CICE, and silt are deficient

in areas of recent mining activity, but their

content is higher in areas with more recovery

time (Ramírez et al., 2019). On the other hand,

the soils of the forested areas surrounding the

mines present extreme acidity, with high con-

tents of Al, MO, and total N and low amounts

of P, Mg, and Ca. Likewise, the K contents are

intermediate, and the CICE is low (Quinto &

Moreno, 2016; Quinto et al., 2022).

Experimental Design: A design strati-

fied by age of succession was used, with three

strata for sampling. Stratum 1, Initial Age (IA)

included areas post-mining, with a succession

time of 12-15 years. This stratum presented

a small shrubby and woody vegetation with

a smaller average diameter and tree species

richness. In contrast, in stratum 2, recovery

Age (RA) corresponded to areas with a 30-35

year recovery time. In this tree, vegetation

with greater diameter and specific richness was

found. The stratum 3 corresponded to primary

forests present in the region and was taken as

the reference scenario.

Establishment of plots: In stratum 1 (EI),

mine areas with a recovery time between 12-15

years were selected. In these mines, 21 per-

manent plots of 50 x 50 m (2 500 m2) were

installed; these plots were the sampling units.

Similarly, in stratum 2 (ER), which corre-

sponded to another forested area with more

than 30-35 years of regeneration, 11 permanent

plots of 50 x 50 m (2 500 m2) were installed;

these plots were the sampling units. In stratum

3, seven plots of one hectare (100 x 100 m)

located in forests of the localities of Opogodó,

Pacurita, and Salero, in the Colombian Pacific,

were used, which were divided into 25 quad-

rants of 20 x 20 m (400 m2).

Measurement of tree diameters: The cir-

cumference at breast height in cm (1.30 m

above ground level) was measured with a tape

measure for all trees with DBH ≥ 10 cm in each

quadrant; later, the circumference values were

transformed to DBH. The perimeter of the tree

trunk where DBH was measured was marked

with yellow spray to guarantee that subsequent

measurements were made in the same strip as

the first. All measured trees were marked with

aluminum plates. Additionally, growth habits

were identified, and the characteristics of each

individual were recorded.

Botanical identification: Trees were iden-

tified to the highest possible taxonomic level

(NN, species, genus, botanical family) in the

herbarium of the Universidad Tecnológica del

Chocó’s “Herbario Chocó.” Gentry (1993) spe-

cialized key was used to make this identification.

Estimation of wood density: To estimate

this variable, the values published in two inter-

national databases of wood density were taken

and generated in tropical forests (Brown, 1997).

In cases where a species or genus found in the

plots was not reported in these databases, the

average genus or family of the species was used.

Estimation of aboveground biomass: To

determine the aboveground biomass of the

trees, the model of Álvarez et al. (2012) includes

DBH and wood density as variables. The model

was:

AB (kg) = EXP(1.59 − 1.22*ln(DBH) +

1.23*(ln(DBH 2) − 0.12*(ln(DBH 3) +

0.69*(ln(pi)))

Where AB is aboveground biomass, DBH

is the diameter, Ln is the Neperian logarithm,

and pi is the wood density. The aboveground

biomass was determined at the ecosystem level,

successional age, plant species, and botanical

family. The biomass growth rate was estimated

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

based on the forest age and the net aboveg-

round biomass values.

Measurement of edaphic characteristics:

In each sampling unit, five composite soil

samples were taken at a depth of 20 cm in the

corners and the center of each 10 x 10 m and

20 x 20 m quadrant. In total, 300 composite soil

samples were taken, 125 in the EI and ER suc-

cessional strata of areas post-mining and 175 in

primary natural forests. The collected samples

were sent to the Biogeochemistry Laboratory

of the National University of Colombia, Medel-

lin. In said laboratory, the samples were ana-

lyzed for texture (sand, silt, and clay), pH, OM

content, Al, effective cation exchange capacity

(ECEC), and nutrient content (N, P, K, Ca and

Mg); using the techniques that are referenced

below: the texture with the Bouyoucos tech-

nique, the pH with the Potentiometer of soils:

water 1:2, the OM with the Walkley and Black

technique and with volumetry, the available P

with Acid L ascorbic acid, UV-VIS spectropho-

tometer, total N with the Micro-Kjeldahl ana-

lytical technique, Al with 1M KCl/Volumetry,

NTC 5 263, nutrients (Ca, Mg, K) with the 1N

ammonium acetate method, neutral, atomic

absorption (Osorio, 2014).

Statistical analysis: To evaluate the varia-

tion of the aboveground biomass as a func-

tion of the recovery time, the non-parametric

Kruskal-Wallis test was used since the assump-

tions of normality and homogeneity of vari-

ances of the data and their residuals were not

met, evaluated with the Bartlett, Hartley, and

Kurtosis statistics (between +2.0 and −2.0).

Then, to determine the linear correlations and

associations between edaphic variables of each

successional stage and aboveground biomass,

multiple regressions were used with the method

of selection of significant variables “backward,”

for which the aboveground biomass and soil

data were transformed with the natural loga-

rithm, due to their lack of statistical normal-

ity. In addition, Spearman rank correlations

and Pearson, as well as multiplicative, recipro-

cal, logarithmic, and linear regressions, were

used to determine correlations and associations

between the physical-chemical variables of the

soil and aboveground biomass. Finally, regres-

sion models were used to evaluate the changes

in the variables through successional time.

These analyses were carried out for the strata of

12-15 and 30-35 years of recovery together and

later separately. The analyses were performed

in the R programming environment (R Core

Team, 2013).

RESULTS

In areas post-mining, aboveground bio-

mass increased significantly with recovery time

(Kruskal-Wallis = 176.9; P = 0.00004). Specifi-

cally, in strata with 12-15 years of succession, it

was 35.17 t ha-1; in those with 30-35 years old

it was 56.30 t ha-1, and in the reference primary

forests it was 178.32 t ha-1 (Fig. 1). Based on the

aboveground biomass values and succession

time, an aboveground biomass accumulation

rate of 2.34 t ha-1year-1 for areas of 12-15 years

of succession, and 1.87 t ha-1year-1 for areas

with 30-35 years of recovery.

It was observed that aboveground bio-

mass at the level of species, the stage from

12-15 years, presented a higher proportion in

Cespedesia spathulata, Cyathea sp., Hampea

romeroi, Vismia sp., and Tovomita weddeliana

(Fig. 2A), while the stage from 30-35 years

presented a higher proportion in Tovomita

weddeliana, Cespedesia spathulata, Clidemia

septuplinervia, Inga lopadadenia, and Cecro-

pia peltate (Fig. 2B). On the other hand, at

the level of botanical families, those with the

highest aboveground biomass were Ochna-

ceae, Cyatheaceae, Malvaceae, Hypericaceae,

Asteraceae, and Clusiaceae in stages of 12-15

years (Fig. 2C), while the families Clusiaceae,

Ochnaceae, Fabaceae, Melastomataceae, and

Urticaceae, were the most representative in the

30–35-year-old stage (Fig. 2D).

The aboveground biomass was positively

correlated with OM, Ca, Mg, CEC, and total N

(Table 1). In the areas with 12-15 years, aboveg-

round biomass was positively related to CICE,

but the association was negative with sand and

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

silt (Table 2). On the other hand, in the areas

with 30-35 years, aboveground biomass was

positively related to P and silt, but the relation-

ship was negative with Ca (Table 3).

When evaluating the changes in aboveg-

round biomass and physicochemical param-

eters of soil through the successional ages, it

was observed that aboveground biomass and

total N increased with the recovery time. At the

same time, P and K decreased with succession

(Fig. 3). On the other hand, OM, Mg, Al, Ca,

and CICE showed curvilinear trends since they

increased in early stages and then decreased in

advanced stages (Fig. 3).

Fig. 1. Aboveground biomass in areas post-mining with different successional ages in Jigualito, Colombian Pacific.

Tabl e 1

Correlations of aboveground biomass and soil parameters

in areas post-mining, with 12-15 years and 30-35 years of

recovery in Jigualito, Colombian Pacific.

Soil parameters

Aboveground biomass

Correlation of

Spearman

Correlation of

Pearson

Organic matter 0.34 (0.016)

Calcium 0.44 (0.0018)0.29 (0.036)

Magnesium 0.32 (0.025)

ECEC 0.40 (0.005)0.32 (0.024)

Nitrogen 0.29 (0.039)

The values in brackets correspond to the p-value. Significant

correlations are included in the table.

Tabl e 2

Multiple regression of aboveground biomass based on edaphic variables in abandoned mines from 12-15 years in Jigualito,

Colombian Pacific.

Source Sum of Squares Df Mean Square Test F P-value

Model 6.9414 3 2.3138 4.62 0.0145

Residue 9.02074 18 0.501152

Total (Corr.) 15.9621 21

Parameter Estimate Standard Error T Statistic P-value

Constant 10.1892 3.49669 2.91397 0.0093

Ln(ECEC) 0.768866 0.30179 2.54768 0.0202

Ln(Sand) -1.31407 0.606447 -2.16683 0.0439

Ln(Silt) -1.00182 0.365238 -2.74292 0.0134

Where R2 = 43.48 %, R2(adjusted) = 34.06 %. Backward variable selection method.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

Fig. 2. Percentage of aboveground biomass of the dominant tree species and botanical families in areas post-mining in

Jigualito, Colombian Pacific.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

Tabl e 3

Multiple regression of aboveground biomass based on edaphic variables in abandoned mines aged 30-35 years in Jigualito,

Colombian Pacific.

Source Sum of Squares Df Mean Square Test F P-value

Model 1 679.22 3 559.739 4.60 0.0375

Residue 973.913 8 121.739

Total (Corr.) 2 653.13 11

Parameter Estimate Standard Error T Statistic P-value

Constant 26.9221 11.0445 2.43759 0.0407

Phosphorus 0.903366 0.32787 2.75526 0.0249

Calcium -9.31647 3.30217 -2.82132 0.0224

Silt 1.56158 0.425212 3.67248 0.0063

Where R2 = 63.29 %, R2(adjusted) = 49.5 %. Backward variable selection method.

Fig. 3. Changes in aboveground biomass and chemical parameters of the soil in areas post-mining at different successional

stages in Jigualito, Colombian Pacific.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

DISCUSSION

After thirty years of succession, areas post-

mining can recover up to 31.57 % of the

aboveground biomass of a primary forest in

the region. Contrary to this, Kalamandeen et

al. (2020) reported an average aboveground

biomass between 1.06 and 6.63 t ha-1 in areas

post-mining for 3 and 4 years in the Guyanese

Amazon. Consequently, in these ecosystems,

after four years, only 1.62 % of the aboveground

biomass of the mature reference forest was

recovered (Kalamandeen et al., 2020). These

results show that, both in the Amazon and in

the Colombian Pacific, after mining, the forests

take many years to recover the aboveground

biomass characteristic of the primary forest of

each region. Consequently, open-pit mining

would affect not only the amounts of carbon in

the ecosystem but also their ability to recover in

times when they would do so with another type

of intervention or disturbance, such as crop

abandonment and forest use.

In the forests of the Colombian Pacific,

Torres-Torres et al. (2017) measured aboveg-

round biomass ranging from 62.6 to 136.1 t ha-1

in secondary forests aged 12 to 40 years, which

had previously been impacted by logging and

abandoned agriculture. This illustrates that,

even within the same biogeographic region,

there are variations in the recovery of aboveg-

round biomass in tropical forests affected by

different human-induced disturbances. Addi-

tionally, Alves et al. (1997) reported that sec-

ondary forests of varying ages and subjected

to different types of human intervention in the

Amazon region could achieve 40-60 % of the

aboveground biomass of a primary forest after

just 18 years of regeneration. Moreover, Ober-

leitner et al. (2021) documented that within a

mere 20 years of regeneration, secondary for-

ests in Costa Rica can recover up to 52 % of the

area’s aboveground biomass of primary forests.

In an analysis carried out a few years ago, at the

level of Neotropical forests, Poorter et al. (2016)

reported that after 20 years’ secondary forests

can reach up to 122 t ha-1 of aboveground bio-

mass in areas previously affected by livestock

and agriculture. Additionally, this study con-

cluded that, after about 66 years of recovery,

they can reach up to 90 % of the aboveground

biomass of a primary forest (Poorter et al.,

2016), which is a situation contrary to what is

recorded in areas post-mining, as mentioned

before. Possibly, this difference is due to the

influence of different factors that affect the suc-

cession in abandoned mines, among which we

can mention the degree of damage caused to

the soil and subsoil, which restricts the recovery

of the ecosystem (Torres-Torres et al., 2023).

The impact of mining on the soil influ-

ences the type of ecological succession (pri-

mary or secondary) and the time required for

aboveground biomass (AB) recovery in the

ecosystem (Chazdon, 2003). Specifically, the

differences in AB recovery between mining-

affected areas and those impacted by other

disturbances are determined by the type of suc-

cession that follows the disturbance (Chazdon

et al., 2016; Guariguata & Ostertag, 2001). In

the case of mining, primary succession is typi-

cally observed (Kalamandeen et al., 2020), as

this activity involves the removal of the organic

soil horizon, resulting in altered structure and

texture, often leaving rocks and sand on the soil

surface (Ramírez et al., 2019). This is akin to the

processes observed in newly formed substrates,

landslide areas, soils resulting from volcanic

eruptions, and igneous rocks where primary

successions occur (Walker, 1993). In contrast,

secondary successions take place on soils with

organic matter, high nutrient content, and seed

banks, such as those following activities like

livestock, agriculture, and logging (Guariguata

& Ostertag, 2001), leading to a much faster

recovery of species richness and biomass. The

slow recovery of AB after mining underscores

the necessity for ecosystem restoration.

Another factor that may be determining

the recovery of the ecosystem in abandoned

mines is the composition of tree species and

their respective capacity to grow and store

biomass. The species recorded in abandoned

mines are commonly reported for degraded

ecosystems and secondary forests with dif-

ferent stages of succession (Alves et al., 1997;

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

Guariguata & Ostertag, 2001). The composi-

tion, diversity, and biomass of these species are

influenced by the fertility or toxicity of the soil

in which they grow (Martins et al., 2015; Ober-

leitner et al., 2021; Poorter et al., 2016; Torres-

Torres et al., 2023). Notably, in abandoned

mines, a high concentration of Al was recorded,

which is undoubtedly generating toxicity for

plants (Carreño & Chaparro-Giraldo, 2013;

Jansen et al., 2002; Quinto et al., 2022); and

consequently, it follows that the species with

the highest aboveground biomass are possibly

the ones with the highest tolerance to Al toxic-

ity (Carreño & Chaparro-Giraldo, 2013). Such

is the case of trees from botanical families such

as Melastomataceae, Rubiaceae, Euforbiaceae,

and Lauraceae, among others, that in previ-

ous studies have been determined as tolerant

and bioaccumulative of Al (Jansen et al., 2002¸

Watanabe & Osaki, 2002). This probably occurs

in the post-mining forests under study, where

species belonging to these botanical families

are well represented (see floristic composition

in Torres-Torres et al., 2023). Therefore, it

would be worthwhile to verify this assumption

in future research.

High levels of aluminum in the soil may

restrict the aboveground biomass storage of

some species in areas affected by mining. This

is because the high availability of this min-

eral in many species with low tolerance to

its excessive content leads to a decrease in

root elongation, reduced stem growth, altered

metabolic and physiological processes, and

decreased ecosystem productivity (Shetty et al.,

2021). Additionally, edaphic Al reduces nutrient

absorption, causes nutritional stress, chlorosis,

and necrosis, decreases leaf size, and affects

photosynthesis (Chandra & Keshavkant, 2021).

Consequently, Al toxicity may be responsible

for the limited recovery of aboveground bio-

mass observed in post-mining forests over a

30-year succession period.

The edaphic conditions are fundamental

for recovering aboveground biomass in aban-

doned mines (León & Osorio, 2014). In this

sense, this study evidenced a positive relation-

ship between aboveground biomass and soil

nutrients, showing that small soil patches with

higher fertility facilitate carbon accumulation

in the ecosystem. This higher aboveground bio-

mass recorded in more fertile abandoned mines

is similar to that reported by Kalamandeen et

al. (2020), who reported a higher aboveground

biomass in abandoned mines with higher N

content. Likewise, Poorter et al. (2016) deter-

mined that soil fertility in Neotropical sec-

ondary forests determines the percentage of

aboveground biomass accumulation. Similarly,

Tucker et al. (1998), in forests with more than

15 years of succession, compared the recovery

of basal area (an indirect measure of biomass)

in nutrient-rich soils with that of infertile oxi-

sols and found that in fertile soils, the basal

area was much more significant with the pas-

sage of successional time, and reached the

aboveground biomass of a primary forest more

quickly. Thus, the positive influence of soil fer-

tility on the recovery of aboveground biomass

in tropical forests (Guariguata & Ostertag,

2001; Moran et al., 2000; Lu et al., 2002) at local

and regional scales, as occurs in areas post-

mining, is evidenced. Likewise, these results

show the influence of different nutrients on

the restoration of aboveground biomass, which

denotes a limitation due to multiple nutrients

(Kaspari et al., 2007; Sullivan et al., 2014), as

previously evidenced by Paoli et al. (2005) and

has been shown experimentally by Davidson

et al., (2004) on secondary successions, and

Vitousek & Farrington (1997), and Harrington

et al. (2001) in primary successions. These

multiple limitations show the need to develop

restorations, applying various nutrients in the

mines. In addition to soil nutrients, sandy soil

texture negatively influenced aboveground bio-

mass. This result is similar to that reported by

Johnson et al. (2000), who showed that aboveg-

round biomass accumulation on nutrient-poor

sandy soils is lower than on non-sandy soils in

secondary forests, which evidences the joint

influence of soil fertility, texture, and moisture

retention on succession (Chazdon, 2003; John-

son et al., 2000). Possibly, in abandoned mines,

the greater macroporosity of the sand facili-

tates the leaching of nutrients, which favors a

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e55276, enero-diciembre 2024 (Publicado Ago. 21, 2024)

lower accumulation of aboveground biomass,

as denoted in this study.

The hypothesis of aboveground biomass

limitation by nutrients (K, P, and N) in plant

succession in abandoned mines. Nutritional

limitation of P and N has been hypothesized

with succession (Walker & Syers, 1976); accord-

ing to which, in tropical soils with initial suc-

cessional ages, there is little availability of N due

to its reduced biological fixation and scarcity

of leguminous plants (Davidson et al., 2004).

However, to the extent that there is a more

significant colonization of N-fixing plants, the

biomass of the ecosystem increases and succes-

sion advances, its availability increases (Reed

et al., 2011). While the levels of phosphorus

in the soil tend to be high in the first succes-

sional stages, over time, its availability tends

to decrease and be limited in the ecosystem

(Vitousek et al., 2010) due to losses due to

leaching and immobilization in carbon oxides.

Fe and Al, especially in tropical clay soils (Reed

et al., 2011; Vitousek et al., 1993; Walker &

Syers, 1976). This hypothesis was corroborated

in the present investigation since, when evalu-

ating changes in aboveground biomass and

nutrients (available P and total N) of the soil

at different successional ages after mining, it

was observed that aboveground biomass and

total N increase with recovery time. Also, this

result is supported by the recent findings of

Quinto et al. (2024a), who observed significant

increases in leaf nitrogen content after add-

ing nitrogen to the soil. This demonstrates the

limitation of forest productivity components

by nitrogen, as trees generally show better

responses when their scarcest resource is sup-

plied (Quinto et al., 2024b).

However, the K content presented a similar

trend to that of P, with which the hypothesis of

nutritional limitation of edaphic P and K on the

biomass and productivity of low-altitude tropi-

cal rain forests was corroborated. The contents

of OM, Mg, Al, Ca, and CICE showed curvi-

linear trends since they increased in the first

stages. Then, they tend to decrease their edaph-

ic concentration in the advanced successional

stages. This is undoubtedly because plants with

progress in succession tend to store nutrients

(K, Mg, and Ca) in their aboveground biomass

(Feldpausch et al., 2004) as a strategy to reduce

their losses by edaphic leaching and thus a

way to make nutrient recycling more efficient

(Chazdon, 2003; Guariguata & Ostertag, 2001).

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

This research was financed through the

project: Evaluation of the effect of soil fer-

tilization on the net production of the eco-

system in areas degraded by mining, as a

strategy to promote carbon capture and the sale

of environmental services in the Chocó Biogeo-

graphic (CODE 1128-852-72243), presented by

the Technological University of Chocó DLC,

National University of Colombia Medellín,

University of Valladolid (Spain), John Von Neu-

mann Pacific Environmental Research Insti-

tute, and SENA, and approved by the Ministry

of Science, Technology and Innovation. We

thank Yeison Rivas, Darlington, and Jesus Erlin

for their support in the field activities.

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