An overburden is known as the dumping of coal mine tailings and other reject materials. It is nutrient-poor, contains elevated concentrations of trace metals and loosely adhered particles of shale, stones, and boulders, and is devoid of true soil characteristics (Dowarah et al., 2009; Novianti et al., 2018). Therefore, the presence of plants on the overburden (OB) is crucial, because not all species can colonize the sites under these conditions, and their presence may facilitate the presence of subsequent species. Some plants do grow on the OB (Novianti et al., 2017). However, a few of these have been identified as exotic species.

Invasion by exotic plant species in an ecosystem is considered a threat because of the deleterious effects they have for people and nature (Kawaletz et al., 2013; Setyawati et al., 2015), and exotic species should be removed or killed whenever possible. However, the potential consequences of invasive species vary widely across ecosystems (Pejchar & Mooney, 2009), and for both people and nature (Koutika & Richardson, 2019). In Indonesia, there are more than 2000 exotic species, 300 of which are classified as invasive (Setyawati et al., 2015), indicating that possibly not all exotic species are invasive. According to Woods and Mariarty (2001), the role of exotic species is unclear.

In previous studies, 123 plant species have been identified from OB (Novianti et al., 2017). The aim of this study was to classify the species in the OB as native and exotic, and to determine the role of the exotic species. Relative coverage was used to determine the dominance of the species. In primary succession, coverage of species may be more representative in describing the dominance of space than the density of species. OB is presented as a model system to study primary succession (Prach et al., 2013) because it is one of the most important human-mediated disturbances that create this condition.

Information about plant species and their roles is essential to understand succession and restoration. An understanding of succession can used as a tool for restoration efforts (Kangas, 2004) because it can be used to accelerate natural succession (Bradshaw, 1987). Therefore, it is necessary to investigate changes in species composition and associated substrates over time to manipulate succession (Dowarah et al., 2009; Novianti, 2020; Walker et al., 2007).

Materials and Methods

Description of study area

The study was conducted in a coal OB dumping area located in Satui District, South Kalimantan, Indonesia. The sampling was carried out on an out-pit dump (i.e., on an OB dumped at certain disposal sites outside of the mine pit), and without leveling on its surface (known as free dump). The determination of OB heaps was based on the following conditions: (1) no disposal process (final dump), (2) known age, (3) identified origin depth, and (4) identified geological formation. According to this, six OBs were used as the study area of primary succession using a chronosequence approach (Table 1).

Vegetation analysis

OB heaps of different ages were selected, that is, 7, 10, 11, 42, 59, and 64 months old. A vegetation study was conducted using the line-transect method. For each transect, the line-intercept method was used (Mueller-Dombois & Ellenberg, 1974); each plant species that was covered by the transect line was recorded, and plant coverage was estimated by measuring the width of each individual from the transect line. This coverage measurement can produce >100% coverage because of overlapping plants. The number of transect lines was adjusted according to the length of the area with a distance between lines of 5 m, while the length of the transect line followed the width of each area (Novianti et al., 2017; 2018).

Species identification

All vegetation was identified using information from the local communities, plant identification books, and environmental impact assessment reports of PT. AI Satui Mine Project, and websites. Exotic and native species and the origin region were identified based on the identification books of Flora van Java (Van Steenis, 1992), a guidebook for invasive alien plant species in Indonesia (Setyawati et al., 2015), and websites (

Data analysis

Data for the native and exotic species were analyzed descriptively. The relationship between the number of species and time was analyzed using linear correlation at a significance level of 5% (α=5%). The range of vegetation was calculated for each species at each site, including coverage and relative coverage, to determine dominance.


The numbers of native and exotic plant species were 57 and 50, respectively. Sixteen species were not identified (Fig. 1). The native species present at all ages of the OB embankment were Cyperus sulcinux, Fimbristylis dichotoma, Paspalum scrobiculatum, Rhyncospora corymbosa, Scleria sumatrensis (herb), and Trema orientalis (shrub). The exotic species consisted of Echinochloa colona, Leersia hexandra, Paspalum conjugatum, Paspalum dilatatum (herbs), Chromolaena odorata, Clibadium surinamense, and Trema micrantha (shrub) (Table 2).

Neither the number of exotic species (r=0.7093, N=6, P<0.05) nor the number of native species (r=0.7051, N=6, P<0.05) showed a significant relationship with time. However, the number of native species tended to be higher than that of exotic species over time (Fig. 2). Based on the species dominance, Paspalum conjugantum, an exotic plant dominated in the first five heap ages and was replaced at 64 months of stockpiling by Neyraudia reynaudiana, a native species. Meanwhile, other species dominated only at a certain age of the heap spoil (Table 3). The relative dominance of native (r=0.0954, N=6, P<0.05) and exotic (r=0.2512, N=6, P<0.05) species did not significantly correlate with time (Fig. 3). In addition, the dominance of exotic species was higher than that of native species at all six stockpile ages. A community dynamic relationship was observed between exotic and native species, that is, a decrease in the percentage coverage area by exotic species was followed by an increase in the percentage coverage area by native species. The decrease in spatial control by exotic species and increase in dominance by native species began to occur at 59 months of stockpiling age.


This study showed that both native and exotic species are present at an early successional stage in OB stockpiles. A higher percentage of native species indicated that seed sources were still available. They can be a source for pioneer vegetation during restoration, considering that the plant species used are generally brought from outside Indonesia, such as Centrosema pubescens (Hindersah et al., 2021). Moreover, the surroundings of the mine have been dominated by fast-growing trees, such as Acacia mangium, as the main tree used in reclamation (Lewis et al., 2022). The native species present throughout the heap were herbs, such as sedges, grasses, and trees, whereas exotic plants were grasses, shrubs, and trees. The presence of plants in the early stages of primary succession is influential because they must cope with unfavorable conditions for establishment. Their growth and distribution are often restricted by nutrient availability because of bare substrates (Dalling, 2008; Glenn-Lewin et al., 1992; Novianti et al., 2018). Furthermore, when these plants die, they become an organic source that will improve substrate conditions, increase safe microsites over time, and consequently increase the number of species (Marrs & Bradshaw, 1993; Novianti, 2013; Walker & del Moral, 2011).

Initially, exotic species dominated both in terms of the number of species and their coverage, perhaps because of their higher response (per capita growth) to the opportunities of niche and resources than that of the resident species (Shea & Chesson, 2002). However, the number of species and the coverage of native species increased with time. Once a surface is colonized, future generations of colonists are likely to be controlled by local seed production or vegetative expansion, and the disturbance may be colonized in successively expanding plant nuclei (Walker & dan del Moral, 2011).

Some native and exotic have roles as pioneers and some as “followers” in the chronosequence of the OB spoil. Pioneers are those that arrive and grow quickly on very poor substrata and appear to facilitate the subsequent vegetation; “followers” are the ones that appear subsequently (Novianti, 2020; Novianti et al., 2018). Temporary dominance by native and exotic plants shows a temporal pattern during succession in post-mining landscapes (Baasch, 2010). These results indicate that both native and exotic species assist in primary succession. These processes may take longer without their presence during the early primary succession. Herbs are present and dominate at the beginning of succession (Dalling, 2008). In addition, shrubs and trees colonize very quickly and dominate after only 64 months, probably because of their close proximity to a diverse species pool and the constant high spoil moisture content (Maharana & Patel, 2013; Novianti et al., 2018).

In conclusion, the species that predominate at the beginning of primary succession in the OB of coal mines are exotic species. The temporary presence of exotic species assists the primary succession process in OB areas by improving the condition of the OB substrate, without which the succession process may possibly take longer. These results indicate that not all exotic species are invasive. Information about the role of exotic species in primary succession is needed as part of the integration between scientific and management concerns for practical knowledge in doing restoration.


This work would not have been possible without financial support and all the facilities from PT. Arutmin Indonesia, Satui Mine Project, South Kalimantan. Author is also thankful to Dr. Achmad Sjarmidi for his critical suggestions and comments that improved the manuscript.

Conflict of Interest

The author declares that (s)he has no competing interests.



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Figures and Tables
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Fig. 1

Number of native and exotic plant species on overburden heaps.

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Fig. 2

Correlation between time (months) and number of native and exotic species on the chrono sequence on coal overburden spoils

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Fig. 3

Correlation between time (months) and total relative dominance of native and exotic species on the chronosequence on coal overburden spoils.

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Table 1

Characteristics of OB dumping sites that were selected as the study area

No. Age of mine OB (mo) Height of OB dump (m) Width of OB dump (ha) Origin depth (m asl)
1 7 38.18 2.68 30 to −80
2 10 20.07 2.06 30 to −80
3 11 16.18 3.66 30 to −80
4 42 19.90 7.09 30 to −80
5 59 24.11 2.14 30 to −80
6 64 29.85 11.87 30 to −80

OB, overburden.


Data from the article of Novianti et al. J Degrade Min Land Manage 2017;4:927-936.

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Table 2

Native and exotic plant species and their relative dominances on the chronosequence on coal overburden spoils

No. Species Life form Time (mo) N E Origin region

7 10 11 42 59 64
1 Acacia mangium Wild. Pt 0.506 0.036 1.080 v Maluku to N. Queensland
2 Ageratum conyzoides Linn. Ah 0.021 0.001 0.003 v Central America and the Caribbean, is now found throughout the world
3 Alternanthera pungens Kunth. Ph 0.288 0.194 v South America; naturalized in Bhutan, Myanmar, Thailand, other parts of Indo-China, Australia, and United States
4 Alternanthera sessilis Linn. Ph 1.445 3.687 6.442 3.947 0.041 v South America
5 Andropogon aciculatus Retz. Ph 0.036 v Indian Ocean, Tropical & Subtropical Asia to Pacific
6 Andropogon chinensis (Ness) Merr. Ah 0.021 v Tropical & S. Africa, SW. Arabian Peninsula, India to S. China and Indo-China
7 Anthocephalus macrophyllus Havil Pt 0.023 v South Asia and Southeast Asia, including Indonesia
8 Araujia hortorum E. Four Pl 0.484 0.127 1.626 v South America
9 Axonopus compressus (Sw.) P.Beauv. Ah 0.125 v Tropical America
10 Benincasa hispida (Thunb.) Cogn Pl 0.190 v Probably native in Japan and Java
11 Blechnum orientale Linn. Pf 0.033 0.138 v Tropical & Subtropical Asia to Pacific
12 Blumea balsaminifera (Linn.) DC Ps 0.013 0.017 0.051 v India to Burma (Myanmar), Indo-China, southern China, Taiwan, Malaysia, Indonesia and the Philippines
13 Blyxa japonica (Mix.) Maxim. Ah 0.185 v Bangladesh, China, Hong Kong, India, Irian Jaya, Kalimantan, Japan, Korea, Laos, Malaysia, Myanmar, Nepal, Papua New Guinea, Taiwan, Thailand and Vietnam
14 Bryophyta 0.497 1.082 0.055 1.615 4.773
15 Celosia argentea Linn. Ah 1.878 0.014 0.001 v The tropical Americas
16 Centotheca lappacea (Linn.) Desv. Ph 0.026 0.055 v W. & W. Central Tropical Africa, Madagascar, Tropical & Subtropical Asia to Pacific
17 Centrosema molle Benth. Pl 0.052 0.048 v The native range of this species is S. Mexico to Tropical America
18 Centrosema pubescens Benth. Pl 0.488 0.056 v Mexico to Tropical America
19 Chloris barbata Swartz. Ph 0.009 v Tropical & Subtropical Old World
20 Christella dentata (Forsk.) Browney & Jerm Ph 0.005 v Tropical & Subtropical Old World to Pacific
21 Chromolaena odorata (Linn.) King & Robinson Ps 2.855 0.564 0.617 2.436 5.281 2.676 v Central & South America
22 Citrullus lanatus (Thunb.) Al 0.047 v Egypt, Ethiopia, Libya, Sudan
23 Clibadium surinamense Linn. Ps 2.053 0.270 10.64 6.946 0.602 6.811 v Tropical America
24 Cynodon dactylon (Linn.) Pers. Ph 0.234 1.261 0.225 4.633 0.236 v Temp. & Subtropical Old World to Australia
25 Cyperus babakans Steud. Ah 0.041 0.023 v South East
26 Cyperus brevifolius (Rottb.) Hass Ah 1.239 v Tropics & Subtropics
27 Cyperus compactus Retz. Ah 0.141 0.094 0.297 0.005 v Madagascar, Tropical & Subtropical Asia to N. Australia
28 Cyperus compressus Linn. Ah 0.005 0.096 v Tropics & Subtropics
29 Cyperus difformis Linn. Ah 0.017 0.303 v Tropical & Subtropical Old World
30 Cyperus entrerianus Boeckl. Ah 0.211 0.610 v Mexico to N. Argentina, Caribbean
31 Cyperus halpan Linn. Ah 0.072 0.108 v Tropics & Subtropics
32 Cyperus iria Linn. Ah 0.208 0.007 v Tropical & Subtropical Old World to Central Asia
33 Cyperus javanicus Houtt Ah 0.894 1.593 0.398 0.040 0.059 v Indian Ocean, Tropical & Subtropical Asia to Pacific
34 Cyperus kyllinga Endl. Ah 0.043 v SE. U.S.A. to N. South America
35 Cyperus polystachyos Rottb. Ah 2.220 1.363 v Tropics & Subtropics
36 Cyperus pulcherrimus Will. Ex. Kunth. Ah 0.071 0.002 v Tropical Asia
37 Cyperus pygmaeus Rottb. Ah 0.042 v Tropical & Subtropical Old World to Russian Far East
38 Cyperus sp.1 Ah 0.012
39 Cyperus sp.2 Ah 0.036
40 Cyperus sp.3 Ah 0.006
41 Cyperus sp.4 Ah 0.007 0.063 0.015
42 Cyperus sp.5 Ah 0.065
43 Cyperus sp.6 Ah 0.039
44 Cyperus sulcinux C. B. Clarke. Ah 0.037 0.310 1.280 0.117 0.233 1.641 v Tropical & Subtropical Asia to Queensland
45 Dactyloctenium aegyptium (Linn.) Willd. Ph 0.034 v Tropical & Subtropical Old World
46 Desmodium heterophyllum (Willd.) DC. Ah 0.121 v Madagascar, Tropical & Subtropical Asia
47 Digitaria ciliaris (Retz.) Koeler Ph 0.064 v Tropical & Subtropical Old World
48 Echinochloa colona (Linn.) Link Ah 1.134 3.939 4.291 0.007 0.001 0.370 v India (Gujarat)
49 Eclipta prostrata Linn. Ah 3.859 v Tropical America
50 Elaphoglossum blumeanum (Fée) J. Sm Ph 0.003 0.003 v Malesia to Solomon Islands
51 Eleocharis dulcis (Burm. f.) Trin. ex. Henschel. Ah 1.470 0.097 0.671 v Tropical & Subtropical Old World
52 Eleusine indica (L.) Gaertn. Ah 0.115 0.022 2.036 0.041 0.018 v India
53 Emilia sonchifolia (Lin.) DC Ah 0.016 0.004 0.194 0.014 v Tropical Africa
54 Eragrostis japonica (Thunb.) Trin. Ah 4.505 7.391 5.387 0.356 1.571 v Tropical & Subtropical Old World
55 Eragrostis leptostachya (R. Br.) Steud Ah 0.369 v E. & SE. Australia
56 Eragrostis tenella (Linn.) P Beau Ah 1.530 1.650 1.165 0.056 0.421 v Tropical & Subtropical Old World.
57 Eragrostis unioloides (Retz.) Nees ex Steud Ah 0.043 0.015 4.785 v S. E. Asia
58 Erechtites valerianifolia (Wollf) DC. Ah 0.028 v Tropical & Subtropical America
59 Erigeron sumatrensis Retz. Ah 0.012 0.011 v Mexico to S. Tropical America
60 Eulophia graminea Lindl. Ah 0.007 v Tropical & Subtropical Asia to Marianas (Guam)
61 Fern/Paku sp.1 0.001
62 Fimbristylis dichotoma (Linn.) Vahl. Ah 13.810 15.680 2.611 1.897 7.986 0.013 v Tropics & Subtropics
63 Fimbristylis litoralis Gaudich Ah 0.672 2.282 4.841 v Tropics & Subtropics
64 Fimbristylis miliaceae (Linn.) Vahl Ah 0.448 0.833 0.025 v Tropical America
65 Fimbristylis schoenoides (Retz.) Vahl Ah 0.056 0.912 v S. E. Asia
66 Fimbristylis sp.1 Ah 0.152
67 Fimbristylis sp.2 Ah 0.013
68 Hodgsonia heteroclita (Roxb.) Hook f. & Thomson Pl 1.037 v E. Himalaya to China (S. Yunnan, Guangxi) and Indo-China
69 Homalanthus populifolius Graham Pt 0.241 0.123 0.328 0.109 v Papua New Guinea to Solomon Islands, E. Australia, Norfolk Island, Lord Howe Island
70 Hyptis capitata Jacq. Ah 0.012 0.075 v Tropical America
71 Imperata cylindrica (L.) Raeuschel Ph 5.909 0.151 2.134 4.147 3.752 v Medit. to Africa and Afghanistan
72 Ipomoea aquatica Forsk. Ah 0.111 v Tropical & Subtropical Old World
73 Lantana camara Linn. Ps 0.064 v Tropical America
74 Leea indica (Burm.f.) Merr. Ps 0.014 v Tropical & Subtropical Asia to W. Pacific
75 Leersia hexandra Swartz Ph 0.437 0.166 0.022 0.047 1.790 0.538 v Tropical America
76 Lindernia crustacea (Linn.) F.Muell. Ah 0.588 0.023 v Tropics & Subtropics
77 Ludwigia hyssopifolia (G. Don) Exell. Ph 0.450 0.004 0.031 0.012 v S. Mexico to Tropical America, N. Australia
78 Lycopodium cernuum Linn. Af 0.004 0.013 v Tropics & Subtropics
79 Lygodium microphyllum (Cav.) R Br. Af 0.149 0.028 v Tropical & Subtropical Old World
80 Macaranga gigantea (Reichb.f.& Zoll.) Műll.Arg. Pt 0.040 v S. Myanmar to W. & Central Malesia
81 Macaranga tanarius (L.) Muell.Arg. Pt 0.018 0.002 v Tropical & Subtropical Asia to W. Pacific
82 Melastoma malabathricum Linn. Ps 0.108 0.004 1.089 0.077 v Asia
83 Melochia corchorifolia Linn. Ah 0.114 0.020 0.000 v Tropical & Subtropical Old World
84 Merremia peltata (L.) Merr. Pl 6.485 v Tanzania, W. Indian Ocean, Tropical Asia to Pacific
85 Mikania micrantha Kunth. Pl 0.336 0.288 v Central and South America
86 Mimosa pudica Linn. Ps 1.736 4.349 6.627 5.488 v Tropical America/S. America
87 Mitracarpus hirtus (Linn.) DC Ah 0.004 v Mexico to Tropical America
88 Mollatus paniculatus (Lam.) Mull.Arg. Pt 0.326 0.579 v Tropical & Subtropical Asia to N. & NE. Queensland
89 Morinda citrifolia Linn. Pt 0.006 v Tropical & Subtropical Asia to N. Australia
90 Nephrolepis sp. Af 0.000
91 Neyraudia reynaudiana (Kunth) Keng ex Hitchc Ph 0.325 0.487 0.533 2.059 15.590 v Himalaya to Central China and Malesia
92 Ochroma pyramidale (Cav. Ex Lam.) Urb. Pt 0.241 v S. Mexico to Tropical America
93 Palaquium oblongifolium (Burck) Burck Pt 0.145 v W. Malesia
94 Panicum repens Linn. Ph 0.778 v Asia or Africa
95 Paspalum conjugatum Berg. Ph 20.620 32.610 13.56 55.030 22.310 6.439 v Tropical America
96 Paspalum dilatatum Poir. Ph 7.501 8.441 3.756 1.131 7.594 14.670 v SE. & S. Brazil to S. South America
97 Paspalum scrobiculatum Linn. Ph 1.104 1.163 0.882 0.140 0.547 7.468 v Tropical & Subtropical Old World to N. & E. Australia
98 Passiflora foetida Linn. Pl 3.306 8.001 0.321 1.926 1.505 v Tropical America
99 Phyllanthus urinaria Linn. Ah 0.184 0.571 v Tropical Asia
100 Piper aduncum Linn. Ps 0.002 v Mexico to Tropical America
101 Pityrogramma calomelanos (Linn.) Link Pf 0.017 0.193 0.289 0.200 0.265 v Mexico to Tropical America
102 Polygala paniculata Linn. Ah 0.430 v Tropical America from Mexico and the Antillies to Brazil
103 Porophyllum ruderale (Jacq.) Cass. Ah 0.025 0.035 0.043 1.318 1.187 v C. & S. America
104 Psidium guineense Swartz Pt 0.004 v Mexico to Tropical America
105 Pteridium esculentum (G. Forst.) Cockayne Pf 0.002 v Tropical & Subtropical Asia to SW. Pacific
106 Pteris vittata Linn. Pf 0.235 0.351 0.686 3.039 v Tropical & Subtropical Old World
107 Rhyncospora corymbosa (Linn.) Britton Ph 10.930 2.064 6.577 3.562 4.055 0.495 v Tropics & Subtropics
108 Saccharum spontaneum Linn. Ph 1.273 0.002 v Sicilia, Africa, Asia to N. & NE. Australia.
109 Sacciolepis indica (Linn.) Chase Ph 0.199 0.152 0.025 v India
110 Scirpus mucronatus (Linn.) Palla Ph 0.069 0.952 0.103 v Europe to Central Siberia and Himalaya, Africa, Brazil to NE. Argentina
111 Scleria bancana Miq. Ah 0.007 v Tropical & Subtropical Asia to Caroline Islands
112 Scleria sumatrensis Retz. Ah 4.734 0.090 2.256 7.885 0.352 0.508 v Seychelles, Hainan to Tropical Asia and N. Australia
113 Solanum torvum Swartz Ps 0.013 0.012 0.057 0.045 0.237 v The Antiles
114 Tree sp. 1 Pt 0.123
115 Tree sp. 2 Pt 0.074
116 Tree sp. 3 Pt 0.008
117 Tree sp. 4 Pt 0.001
118 Trema micrantha (L.) Blume Ps 0.556 0.152 3.098 0.032 0.009 0.002 v Tropical & Subtropical America
119 Trema orientalis (L.) Blum Ps 1.660 0.729 7.853 2.572 0.224 2.701 v Tropical & Subtropical Old World
120 Typha angustifolia Linn. Ph 10.110 17.990 1.523 0.018 5.164 0.269 v Temp. Northern Hemisphere
121 Vernonia cinerea (L.) Less Ah 0.042 0.177 0.277 v Unknown/Old World
122 Vitaceae Pl 2.293 0.100
123 Wedelia trilobata (Linn.) Hitchc. Ph 0.025 2.020 v Tropical America

Ah, annual herb; Ph, perennial herb; Pl, perennial liana; Ps, perennial shrub; Pt, perennial tree; N, native; E, exotic.

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Table 3

Top species with top 10 relative dominances

No. Time (mo)

7 10 11 42 59 64
1 P. conjugatum P. conjugatum P. conjugatum P. conjugatum P. conjugatum N.reynaudiana
2 F. dichotoma T. angustifolia C. surinamense S. sumatrensis F. dichotoma P. dilatatum
3 R. corymbosa F. dichotoma P. foetida C. surinamense P. dilatatum P. scorbiculatum
4 T. angustifolia P. dilatatum T. orientalis E. japonica M. pudica C. surinamense
5 P. dilatatum E. japonica E. japonica M. pudica C. odorata P. conjugatum
6 M. peltata E. colona R. corymbosa I. cylindrica T. angustifolia M. pudica
7 I. cylindrica A. sessilis A. sessilis R. corymbosa C. dactylon F. litoralis
8 S. sumatrensis R. corymbosa E. colona T. orientalis R. corymbosa E. uniloides
9 P. foetida C. argantea P. dilatatum C. odorata A. sessilis Bryophyta
10 C. odorata C. javanicus T. micrantha F. dichotoma E. prostrata P. vittata