Nutrient Mineralization and Leaf Litter Preference
by the Earthworm Pontoscolex corethrurus
on Iron Ore Mine Wastes
S. R. Ganihar
Abstract
The effects of the earthworm Pontoscolex corethrurus
(Muller) on the rate of mineralization of cattle dung-
amended iron (Fe2) ore mine wastes and its preference
for partially decomposed leaf litter with contrasting chem-
ical composition were studied in pot trials. The growth and
survival rates of earthworms showed significant positive
correlations with percent of organic matter. During 96
days of exposure, the earthworms significantly increased
exchangeable Ca2, Mg2, PO43 and NH4-N. Iron ore
mine wastes amended with 5–10% organic matter sup-
ported earthworm fauna better than mine wastes amended
with 0–3% organic matter. The leaf litter preference shown
by the earthworm was, in descending order, Phyllanthus
reticulatus, Tamarindus indica, Anacardium occidentale,
Casuarina equisetifolia, Acacia auriculiformis, and Eu-
calyptus camaldulensis. A significant positive correlation
was observed between the survival and growth rates of
Introduction
Among the most severe forms of biological degradation
are those associated with mining. Biologically active soil is
completely removed or significantly reduced by opencast
mining. Abandoned iron ore mines consisting of barren re-
ject dumps occupy nearly 5–10% of the total land area in
Goa, India. The reject dumps are spread over hundreds of
hectares of forest and agricultural land and provide envi-
ronmental conditions that are extremely hostile for living
organisms (Ganihar 1990). Prerequisites for restoring bio-
logical activity in such sites include amelioration of limit-
ing factors, stabilization of the physicochemical environ-
ment, and provision of food sources for decomposer
organisms in the form of organic matter or plant litter (Curry
& Boyle 1995).
A number of studies have revealed that earthworms are
capable of improving the fertility and productivity of re-
claimed soils (Curry 1988). The introduction of selected
earthworms into degraded or newly restored land is
known to promote soil improvements (Butt et al. 1995).
Some studies (e.g., Vimmerstedt & Finney 1973) have
Department of Zoology, Dhempe College of Arts and Science, Miramar,
Panaji, Goa-403 001, India
© 2003 Society for Ecological Restoration International
DECEMBER 2003 Restoration Ecology Vol. 11 No. 4, pp. 475–482
earthworms and the nutrient contents of partially decom-
posed leaf litter. The first three plant species were signifi-
cantly richer in nutrients, mainly organic carbon, calcium,
phosphorus, and nitrogen, than the other two plant spe-
cies. Acacia auriculiformis and E. camaldulensis litter
were preferred less because of their high lignin and
polyphenolic compounds, despite being rich in other
macronutrients like nitrogen and phosphorus. It is con-
cluded that the introduction of P. corethrurus to cattle dung-
amended (5–10%) iron ore mine wastes or revegetation of
the sites with P. reticulatus, T. indica, and A. occidentale
plant species should be attempted before earthworm in-
troduction. The litter from these species acts as a source of
food for earthworms, thereby hastening the process of res-
toration of abandoned iron ore mines of Goa, India.
Key words: amendments, earthworms, iron mine wastes,
organic, restoration.
demonstrated the feasibility of introducing earthworms
into revegetated acid coal mine spoils to increase the rate of
incorporation of organic matter. Emmerling and Paulsch
(2001) studied the impact of various amounts and types of
organic waste materials on the development of earthworm
communities, both in field mesocosms and laboratory mi-
crocosm. Coelho (1990) studied earthworms in relation to
bacterial activity and their possible role in the rehabilita-
tion of iron mine tailings in Goa.
The objectives of the present study were (1) to examine
whether the earthworm Pontoscolex corethrurus (Muller)
could be established on abandoned iron ore mines in Goa,
India in association with suitable organic matter amendments
that may hasten the ecological restoration of mines, and (2)
to determine the earthworm preference for leaf litter of dif-
ferent plant species used for revegetation of mine lands.
Methods
The tropical earthworm P. corethrurus Muller (Family:
Glossoscolecidae) is widely distributed in the Indian pen-
insula and Northwest region of India (Julka 1993). One of
the most dominant species in woodland habitats (Krish-
namoorthy & Ramachandra 1988), P. corethrurus is known
for its subsurface feeding, which occurs at depths between
5 and 10 cm in shady regions with low salinity and high
475
Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes
organic matter (Kale 1998). It was selected for these ex-
periments because it is locally abundant, it is known to
survive around mining areas in Goa (Ganihar 1996), and
it can tolerate temperatures up to 28oC because of its se-
cretion of a rich mucus that prevents desiccation. It can
also tolerate a wide range of edaphic conditions, namely pH
(5.9–7.2), moisture (50–90%), organic matter (0.5–8.53%),
and soil texture (silt-loam to sandy-loam) (Choudhuri &
Bhattacharjee 1999). The fine prostomium can be ex-
tended and withdrawn and exerts a piston-like action
while making its way into the soil, allowing the worm to
burrow without the use of body segments (Kale 1998),
thus facilitating its movements through soils. The intesti-
nal secretion, the mucus, and other excreta make the soil
slightly compact and cement-like, which may act as an ini-
tial soil binder at mine sites. Pontoscolex corethrurus is an
endemic species and propagates quickly by parthenogen-
esis (Edwards & Lofty 1977).
Experimental Design
Experiment 1. Earthworms were collected and hand sorted
from loamy soil in the fields in and around Panaji city
(15o2925N, 73o4845E). They were placed for 3 days in
large petri dishes (10 per petri dish) containing blotting
paper to empty the gut contents. They were then added to
red clay pots (3 L, 18 cm diameter, 12 cm height). Each pot
was filled with 2.5 kg of air-dried and sieved (2-mm mesh)
iron mine waste from reject dumps and abandoned mines
of Sesa Goa (P) Ltd. at Sanquelim, Goa. Cattle dung col-
lected from fields near Panaji was air dried, finely ground,
and sieved through 2-mm mesh. Various amounts of mine
soil waste and cattle dung (2.5 kg  0 g, 2.475 kg  25 g,
2.375 kg  75 g, 2.325 kg  175 g, and 2.250 kg  250 g)
were added to each pot and mixed thoroughly with the
iron mine waste. This treatment created 0, 1, 3, 5, 7, and
10% (w/w) of organic matter, respectively, in the different
pots. Adequate water was added to each pot to maintain
moisture at field capacity (30%). Each pot was immersed
in a plastic container (4 L, 14 cm height, filled with 2 L of
water) such that the outer surface of the pot was kept cool
to maintain the temperature between 23 and 25oC. Tem-
peratures between 23 and 25oC provide high growth rates
for earthworms (Van Gestel et al. 1992). The pots were al-
lowed to settle for 2 days before introducing earthworms.
Ten healthy and active earthworms were introduced from
the petri dishes to each pot. The pots were then covered
with muslin cloth held on with rubber bands. Five repli-
cates of each treatment were maintained (total of 30
pots).
Each earthworm was weighed to obtain initial wet bio-
mass. The number of survivors and their live mass were re-
corded at 14-day intervals for 96 days. Growth rates were
estimated by taking the body weight of each surviving
earthworm, weighed to the nearest 0.01 mg. Dead earth-
worms were removed from the pots as soon as they were
detected to avoid contamination. Growth rate corrections
476
were made for dead earthworms by deducting the initial
mean weight from the final total weight (Petrusewicz &
Macfadyen 1970). Soil samples were collected and ana-
lyzed for chemical properties before the earthworms were
added and again at the end of the experiment.
Experiment 2. Leaf litter samples of Acacia auriculiformis
(Family: Mimosaceae), Casuarina equisetifolia (Family:
Casuarinaceae), Eucalyptus camaldulensis (Family: Myrt-
aceae), Anacardium occidentale (Family: Anacardiaceae),
Phyllanthus reticulatus (Family: Euphorbiaceae), and Tam-
arindus indica (Family: Caesalpiniaceae) were collected
from the forest floor just after leaf fall. The first three tree
species are native to Australia and widely planted; the latter
three are native to India. The leaf litter was air dried in the
laboratory at room temperature and kept separated by
species. The leaf litter and additional quantities of cattle
dung were then soaked in tap water containing 0.1%
KNO3 and 0.01% K2HPO4 to accelerate the process of de-
composition (Hendriksen 1990). After 24 hr the water was
drained, samples were placed in 2.5-L plastic jars, and the
mouth of each jar was covered with a polythene sheet with
a few slits. Distilled water was supplied periodically to
compensate for evaporation losses. Three replicate jars for
each litter type and cattle dung combination were used.
After 2 months the partially decomposed litter and dung
were air dried and stored at room temperature (until the
experiments were completed). Subsamples were crushed
and analyzed for nutrients.
Fourteen sets of clay pots, each containing about 2.250
kg of mine waste plus a 250-g mixture of partially decom-
posed litter and cattle dung as in experiment 1, were estab-
lished as follows: (1) 6 contained mine waste plus leaf litter
(one pot for each leaf litter type); (2) 1 contained mine
waste plus cattle dung; (3) 6 contained forest soil plus leaf
litter (one pot for each litter type); and (4) 1 contained for-
est soil plus cattle dung. Each mixture was composed of
10% organic matter, as required for the normal survival
and growth of earthworms (Edwards & Lofty 1977; Lee
1985). Litter and cattle dung were mixed thoroughly with
the mine soil. Triplicates of each treatment were main-
tained (total of 42 pots). The control pots in experiment 1
also served as controls in experiment 2, because both ex-
periments were run simultaneously. In addition, forest soil
was assumed as a control in experiment 2.
Ten days after the addition of litter 10 earthworms were
introduced to each pot. Before their introduction each
earthworm was weighed to measure initial wet biomass.
The survival and growth rates of the earthworms were
measured at intervals of 14 days for 96 days. Each week 10 g
of decomposed litter and/or cattle dung was added to the
respective pot to compensate for the loss due to earth-
worm feeding (an average 8–10 mg per 1 g of body weight
was suggested by Edwards & Lofty 1977). Only data col-
lected at the end of the experiments were used for the sta-
tistical analysis, because the corresponding chemical anal-
ysis was performed at the end of the experiments.
Restoration Ecology DECEMBER 2003
 Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes

Physicochemical Analysis of Soil and Litter based principal components analyses (PCAs) were per-
Soil pH and electrical conductivity were measured by pre- formed (growth and survival percentages) versus physico-
paring a soil solution using distilled water (1 g in 20 mL) and chemical parameters. PCA is a multivariate statistical
calibrating the solution with an Elico-pH meter (Elico technique that attempts to explain the observed variations
Company, Pradesh, India). Water-holding capacity was in a large data set on the basis of as few (uncorrelated) at-
measured using sintered crucibles filled with oven-dried tributes as possible. The extracted “factors” are ranked in
sieved (2-mm mesh) soil samples (100 g). After 24 hr soil order of decreasing importance in terms of the amount of
samples were placed over a container filled with water in variation accounted for by each factor. The first factor axis is
such a way that the water from the container just touched positioned close to the mean so that it accounts for the great-
the bottom of the crucible. Then the water percolated est possible amount of variation (the sums of the squared
slowly, and the resulting increase in the weight of the cru- factor loadings are minimized) and each successive factor ac-
cible indicated the moisture-holding capacity of the soil counts for a progressively smaller amount of variation. PCA
sample. Soil textures were estimated by the use of sieves of was conducted using MINITAB 8.3 (Minitab 1991).
different meshes. The soil was passed successively through
a series of sieves with different size (diameter) of meshes Results
(pores). Sieving separated different fractions (different
size particles) from the sample, and then the relative pro- Physicochemical Characteristics of Soil and Litter
portions of each fraction were calculated. A particle diam- Table 1 shows the mean values (n  20) and the significant
eter of less than 0.002 mm is clay, between 0.002 and differences in physicochemical characteristics between
0.02 mm is silt, 0.02–0.2 mm is fine sand, 0.2–2 mm is mine soil and forest soils used in the experiments, namely
coarse sand, and above 2.0 mm is gravel (Arakeri et al. salinity, water-holding capacity, clay content, organic mat-
1967). Specific gravity and porosity were measured by us- ter, Fe3, SiO2 (p  0.001), bulk density, fine sand, Al2O3
ing a pyknometer (Black et al. 1965) and oven-dried soil ( p  0.01), silt, pH, specific gravity, porosity, total chlo-
samples. Soil organic matter was determined as loss-on- ride, and ( p  0.05).
ignition of oven-dried soil over 24 hr in a muffle furnace at Nutrient contents of the resource materials (Table 2) in-
550oC (Allen et al. 1976). To estimate organic carbon the dicate that litter of A. occidentale, C. equisetifolia, and E.
chromic acid colorimetric method (Black et al. 1965) was camaldulensis was significantly richer in organic carbon
used by preparing standard curves. Soil samples each than litter of other species; Ca2 was significantly higher in
weighing 5 g were kept overnight in a mixture of one nor- mine soil; Mg2 values were significantly higher in T. in-
mal K2Cr2O7 and H2SO4 and centrifuged for 10 minutes at dica, cattle dung, and E. camaldulensis litter; PO43 values
2,000 rpm. The green chromous color of the supernatant
liquid was read on a colorimeter. Carbon values were esti-
mated from the standard curve (Perur et al. 1972). Table 1. Mean values  SD of physicochemical characteristics
Exchangeable mineral nutrients were estimated as fol- of mine wastes and forest soils used in the experiments.
lows. Phosphorus levels were estimated colorimetrically Mine Waste Forest Soil
by the molybdenum blue method after extraction with Parameters (n  20) (n  20)
Trough’s reagent, calcium, and magnesium by the EDTA Grain size (% by wt.)
titration method (Allen et al. 1976) and inorganic nitrogen Clay*** 13.70  0.82 21.38  1.18
(ammoniacal nitrogen) by the micro-Kjeldahl alkaline Silt* 18.15  2.35 14.67  1.02
permanganate method (Perur et al. 1972). Total chlorides Fine sand** 16.86  2.24 12.08  1.12
were estimated by the titrimetric method. Ore oxides/ele- Coarse sand 39.39  3.32 41.80  3.10
ments were measured by the dichromate method (Fe3), Gravel 11.90  1.23 10.07  0.16
EDTA method (Al2 O3), and gravimetric method (SiO2) Texture Sandy loam Sandy clay loam
pH* 6.8  0.5 6.0  0.6
(Ray et al. 1979). The values of lignin and tannin contents EC (mmhos)*** 0.063  0.009 0.068  0.024
of plant litter were obtained from the literature (Krishna- Bulk density (g/cm3)** 0.93  0.13 0.79  0.30
murthi 1969; Chadha 1985, 1989; Shastri 1989; Phondke 1992). Specific gravity (g/cm3)* 3.02  0.32 2.59  0.11
Porosity (%)* 69.16  1.53 70.84  7.5
Water-holding
Data Analysis capacity (%)*** 29.0  3.6 80.6  6.20
Data were analyzed with analysis of variance (ANOVA) Organic matter
LOI (%)*** 6.38  0.8 10.31  1.72
to test the differences in rate of mineralization between
Chlorides (%)* 34.92  3.2 39.76  2.12
treatments and leaf litter selection. For growth and sur- Iron-Fe3 (%)*** 56.24  2.34 25.31  4.12
vival percentages ANOVA was applied after arcsine Alumina-Al2 O3 (%)** 5.53  0.20 25.4  3.1
transformation (Snedecor & Cochran 1961). The Tukey’s Silica-SiO2 (%)*** 3.20  0.6 24.92  3.14
test was also used wherever the overall F-ratio was signifi-
Parameters show significant differences among forest and mine soils: * p  .05,
cant (Haber & Runyon 1973). The Student’s t-test was ap- ** p  .01, *** p  .001.
plied for initial and final values of minerals. Correlation- EC, electrical conductivity; LOI, loss on ignition.

DECEMBER 2003 Restoration Ecology 477

Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes

Table 2. Mean values  SD of nutrients of different materials used in both experiments.

Organic Ca2 Mg2 Po43 NH4-N

Carbon (%) (mg /100 g) (mg /100 g) (mg /100 g) (mg /100 g)
Resource Materials p  0.01 p  0.001 p  0.01 p  0.05 p  0.01

Iron mine waste 0.71  0.06a 41.40  2.36c 13.15  1.23a 2.50  0.62a 3.36  0.64a
Forest soil 3.80  0.07b 26.14  0.86d 82.14  1.92d 58.12  3.53c 35.41  3.90b
Cattle dung 28.72  1.42c 4.80  0.62a 124.80  1.64f 133.00  4.73e 85.12  1.36d
Acacia auriculiformis 32.31  2.08c 6.41  0.35b 36.48  0.53b 62.40  3.15c 79.52  2.83d
Anacardium occidentale 40.27  2.55d 4.80  0.82a 59.62  2.05c 25.20  1.23b 59.63  1.54c
Casuarina equisetifolia 37.34  2.26d 3.20  0.41a 48.00  3.23b 55.20  2.22c 44.80  2.84b
Eucalyptus camaldulensis 38.06  1.48d 8.00  0.83b 101.76  4.46e 58.00  2.16c 47.04  1.63b
Phyllanthus reticulatus 33.75  1.96c 6.41  0.82b 45.12  1.11b 144.00  4.26e 59.36  1.84c
Tamarindus indica 25.75  0.92c 16.00  1.85c 130.56  2.09f 76.80  1.68d 78.40  2.00d

Values followed by a different letter are significantly different from other values within the same column at the 1% level, determined with Tukey’s T-test.

were significantly higher in P. reticulatus litter and cattle
dung; and NH4-N values were significantly higher in cattle
dung, A. auriculiformis, and T. indica than in other plant
resource materials.
Experiment 1
Earthworms showed a 10–65% increase in survival rate
and 1–17% increase in growth rate in amended mine soil
when compared with control (Fig. 1), and these rates also
showed a significantly positive correlation with the
amount of organic matter (r  0.90, p  0.001, n  30; r 
0.89, p  0.001, n  30, respectively). The amendments
containing 5% or more organic matter showed positive
growth (8–17%) and better survival rates (80–85%) than
the control. ANOVA values indicated significant differ-
ences among the treatments when comparing survival (p 
0.01) and growth rates (p  0.001). The treatment with
Figure 1. Percentage survival and growth rate (means  SD) in
experiment 1 of Pontoscolex corethrurus in iron ore waste amended
with cattle dung and (treatments) (1) 0%, (2) 1%, (3) 3%, (4) 5%, (5)
7%, or (6) 10% organic matter. Bars with different letters are
significantly different at the 1% level determined with Tukey’s HSD
test. (Upper case letters, differences in survival rates; lower case
letters, differences in growth rates.)
10% organic matter showed a significant difference
(Tukey’s test p  0.001) from all other treatments in both
survival and growth rates.
The changes in the levels of the four extractable miner-
als studied are depicted in Figure 2. There was no signifi-
cant difference in initial values of Ca2 among treatments
(p  0.05), but there was a significant difference in the
final values of Ca2 (p  0.001). The increase in the final
values was significant (p  0.001), ranging from 31%
(treatment with 1% organic matter) to 64% (treatment
with 10% organic matter). The initial and final values of
Mg2 were significantly different among the treatments
(p  0.05 and p  0.001, respectively). Manifold increases
(7–8 times) in the final values over the initial values were
observed.
The initial values of PO43 did not differ significantly
(p  0.1) among treatments, but a significant difference
was observed among the final values (p  0.01). The final
values increased significantly (60–70%), especially in
treatments with 7% and 10% organic matter. The initial
and final values of NH4-N differed significantly among the
treatments (p  0.01 and p  0.01, respectively). There
was a 50–55% increase in final values over the initial
values in all treatments (Fig. 3).
In the PCA using a 5 6 matrix (5 replicates and 6
treatments), the first two PCA factors accounted for 95%
of the variance, with individual values of 90% and 5% of
the original variance, respectively. The input variables
were earthworm growth and survival. Results are graphi-
cally represented by the projection of sample pairs onto
two axes defined by the first two pairs of principal compo-
nents axis (i.e., “factor loadings” were generally used on
coordinates). The plane projection of greatest interest is
that defined by the pair of principal components that are
associated with the two components that explain the larg-
est variance. Projection onto this plane will generally show
the samples more discretely separated from each other
than in any other projection (Sokal & Rohlf 1995). The ex-
amination of the first two principal component loading
plot (Fig. 3) indicated three groups of variables, one with
0% organic matter, the second with 1 and 3% organic

478 Restoration Ecology DECEMBER 2003


Figure 2. Initial and final levels of nutrients (means  SD) in
experiment 1: Ca 2, Mg2, PO3, and NH4-N. The bars with
different superscript letters are significantly different at the 1% level,
determined with Tukey’s HSD test. (Upper case letters, differences
in final values; lower case letters, differences in initial values of
nutrients.)
matter, and the third with 5, 7, and 10% organic matter.
The position of the treatments in the coordinates of the
two principal components are determined by the earth-
worms’ activities (survival and growth rates). These activi-
ties increase the rate of mineralization in a similar way in
mine soils.
DECEMBER 2003 Restoration Ecology
Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes
Figure 3. Plot of loadings on the first two components, PC-1 and
PC-2, of experiment 1. The variables are 0%, 1%, 3%, 5%, 7%,
and 10% organic matter.
Experiment 2
The survival and growth rates of earthworms in different
media containing litter and mine soil waste and litter and
forest soils are shown in Figure 4. Of the media compared
earthworms in forest soils exhibited highest survival rates
in cattle dung (100%), P. reticulatus (95%), and A. occiden-
tale (92%). A similar outcome was observed on mine soils.
On mine soils the best supporting materials for earthworms
were cattle dung (85% survival rate), P. reticulatus (60%
survival rate), and A. occidentale (60% survival rate). In
forest soils earthworms showed the highest growth rates
on cattle dung (150%), P. reticulatus (67%), and T. indica
(50%). The highest growth rates attained in mine soil were
with cattle dung (8%), P. reticulatus (2.5%), and A. occi-
dentale (2.5%). In both mine soil waste and forest soils
earthworms appeared to prefer cattle dung over any other
litter. After cattle dung P. reticulatus was the most preferred
litter material (Fig. 4). Earthworms did not survive well in
media containing litter of E. camaldulensis, C. equisetifolia,
and A. auriculiformis (32–40%); in fact, earthworms showed
negative growth in E. camaldulensis (4.2%) and A. auricu-
liformis (25%) by losing body weight due to starvation.
ANOVA indicated significant differences in the sur-
vival (p  0.001) and growth (p  0.001) rates of earth-
worms in mine soils with different litter. Similarly, survival
479
Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes
Figure 4. Percentage survival and growth rate (means  SD) in
experiment 2 of Pontoscolex corethrurus in partially decomposed
litter and cattle dung (Aa, A. auriculiformis; Ao, A. occidentale; Ce,
C. equisetifolia; Ec, E. camaldulensis; Pr, P. reticulatus; Ti, T. indica;
Cd, cattle dung). Bars with different superscript letters are
significantly different at the 1% level, determined with Tukey’s HSD
test. (Upper case letters, differences in survival and growth rates in
forest soils; lower case letters, differences in growth rates in
mine soils.)
rates (p  0.001) and growth rates (p  0.001) of earth-
worms showed variation in forest soil with different leaf
litter. Tukey’s HSD test indicated clearly that cattle dung,
P. reticulatus, and A. occidentale were significantly (p 
0.001) better than other litter in terms of survival and
growth of earthworms (Fig. 4).
The first two principal components accounted for 87%
of the variance with individual values of 73 and 14% origi-
nal variance, respectively. Examination of the first two
principal component loading plots (Fig. 5) indicated two
groups of variables, the first group containing A. auriculi-
formis, E. camaldulensis, and C. equisetifolia and the sec-
ond group containing A. occidentale, P. reticulatus, T. in-
dica, and cattle dung. In this case projection of the sample
points to the plane defined by components 1 and 2 permits
the separation of the samples into two groups by earth-
worm activities. The position of the treatment in the coor-
dinates of the two principal components, marked by the
earthworms’ activities (survival and growth rates) due to
their preference, is seen in Figure 5.
480
Figure 5. Plot of loadings on the first two principal components,
PC-1 and PC-2, in experiment 2. The variables are the litter of
partially decomposed Aa, A. auriculiformis; Ao, A. occidentale; Ce,
C. equisetifolia; EC, E. camaldulensis, Pr, P. reticulatus; Ti, T. indica;
and Cd, cattle dung.
Discussion
Abandoned iron ore mines of Goa are highly stressed hab-
itats with no soil profile and a disturbed hydrological re-
gime. The environmental conditions, especially the physic-
ochemical characteristics, make them unsuitable for the
immediate growth of earthworms, thus delaying initiation
of the organic cycle. When left alone recovery in these
habitats requires several years (50 years) (Ganihar
1990). Earthworms are important processors of soil and
nutrients in the undisturbed areas around mines (Ganihar
1996). Earthworm introduction on abandoned mines, after
making suitable amendments with organic matter (5–
10%), will initiate an organic cycle and potentially hasten
the process of restoration.
The results of this study clearly indicate that a minimum
of 5% organic matter was required for maximum survival,
and growth continued to increase at the highest level of
amendment of 10%. Similar results were observed by Coelho
(1990) on iron mines of Goa amended with litter of Mangi-
fera indica as a source of organic matter in his experiments
with Pheretima orientalis. Mineralization rates of ex-
changeable cations were significant (p  0.01) in mine
soils after the introduction of P. corethrurus in the present
study. This is similar to the results observed by Kale and
Krishnamoorthy (1980) for Ca2 and by several studies of
Mg2 (Lee 1985), PO43, and NH4-N (Haimi & Huhta
1990; Haimi & Einbork 1992; Binet & Trehen 1992; Curry
et al. 1995; Saetre 1998 in natural soils; Emmerling &
Paulsch 2001 in mine wastes).
Significant differences were observed in survival and
growth rates of earthworms between mine soil and forest
soil that were amended with various combinations of leaf
litter and cattle dung. Earthworms grow better in the pres-
ence of cattle dung, but the best growth seems to occur in
Restoration Ecology DECEMBER 2003

forest soil amended with cattle dung. However, for resto-
ration purposes this would require that a prohibitive quan-
tity of cattle dung be introduced on mine wastes. The al-
ternative is to select plant species that can act as pioneer
species to facilitate the recovery of earthworms on mines.
In turn this would help to restore the ecology of mines.
The leaf litter that earthworms preferred was P. reticu-
latus, perhaps due to its richness in nutrients like PO43
(144 mg/100 g) and its lower concentrations of lignin (8–
10%) and tannins (10–12%) (Krishnamurthi 1969). The
other preferred litter sources were T. indica, which is rich
in NH4-N (78 mg/100 g), with 8–10% lignin and 8–10%
tannins (Chadha 1989), and A. occidentale, which is rich in
organic carbon (40%), although tannin content was 20%
and lignin content was even less (10%) (Chadha 1985).
The least preferred plant litters were C. equisetifolia, with
10–15% tannin and 23% lignin (Phondke 1992); E. cam-
aldulensis, with 10–12% tannins and 12–15% lignin (Shas-
tri 1989); and A. auriculiformis, with 23% tannins and 25%
lignin (Chadha 1985), even though these litters all are
moderate in nutrients, mainly NH4-N and PO43. The lat-
ter three are Australian in origin, so the native litters were
preferred by the earthworms. The lack of preference for
these litter sources is probably due to the high contents of
lignin and tannins as suggested by Tian et al. (1995). Simi-
larly, Hendriksen (1990) examined the preference of leaf
litter in relation to its C:N ratio, lignin concentration, and
initial and final concentrations of polyphenolics before
and after decomposition. He found that the polyphenolic
concentration is more important in determining litter pref-
erence. Studies with Lumbricus terrestris have shown that
tannins, a group of polyphenols, have a major influence on
palatability (Satchell 1967; Daniel 1991) and have a direct
effect on the decomposition of highly palatable plant ma-
terials like dandelion leaves. Substances like lignin and
tannins make litter unsuitable for earthworm consumption,
reducing the rate of decomposition by microbes. Apart from
lignin and tannins the essential oils present in the leaves of
E. camaldulensis (1.5–2.5%) may act as antibiotics to mi-
crobes and prevent their colonization on litter, thus check-
ing the rate of decomposition. Similarly, litter of A. auricu-
liformis and C. equisetifolia are richer in lignin and tannin,
making them unsuitable for earthworms. Even the living
phyllode of A. auriculiformis is distasteful to herbivores,
including arthropods (Ganihar 1998).
In conclusion, the present study suggests that (1) 5–10%
organic matter (cattle dung) amendment is required for in-
creased survival and growth rates of earthworms on iron
ore mines sites, and (2) the plant species suitable for
revegetation on abandoned iron ore mines are the local
native species P. reticulatus, T. indica, and A. occidentale,
being rich sources of NH4-N and PO43, with minimal levels
of lignin, tannins, and essential oils in their litter. These
species can support earthworm fauna immediately and
help initiate the organic cycle, thereby hastening the pro-
cess of rehabilitation of soil organisms in this region.
These pot trial experiments eliminated a number of stress
DECEMBER 2003 Restoration Ecology
Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes
factors (e.g., drought, heat, field compaction, etc.) threat-
ening earthworm survival. However, systematic field trial
experiments will likely provide more insight into the po-
tential problems in meeting restoration objectives. Before
undertaking soil amendments and revegetation programs
on a large scale, pot and field trials are needed to predict
the suitable measures for complete recovery.
Acknowledgments
I thank the Principal and Prof. S. M. Shetty for laboratory
facilities and M/s Sesa Goa (P) Ltd. for allowing me to col-
lect iron mines wastes and for analyzing ore elements,
namely iron, manganese, aluminum, and silicon. I am in-
debted to Prof. S. K. Saidapur, Prof. Raghavendra Gadag-
kar, Dr. S. N. Harkantra, and Dr. S. G. Dalal for help with
analyzing the data with PCA; Dr. S. M. Karisiddaiah for
valuable suggestions in revising the manuscript; Dr. M. P.
Tapaswi for help with the literature survey; and Prof.
(Miss) Isabel Vaz for valuable suggestions. Thanks are also
due to Shri. S. K. Kulkarni, Shri. Prakash Pednekar, and
Shri. Mohan Naik for their technical assistance. This re-
search was supported by a grant (6/21/87-ER) from the De-
partment of Environment, Forest and Wildlife, Govern-
ment of India, New Delhi. I thank Sheila Kee and two
anonymous referees for their critical review and construc-
tive suggestions that improved the manuscript immensely.
LITERATURE CITED
Allen, S. E., M. H. Grimshow, J. H. Parkinson, and C. Quarnby. 1976. Chem-
ical analysis of ecological materials. Blackwell Scientific, Oxford.
Arakeri, H. R., G. V. Chalam, P. Satyanarayan, and R. L. Donahue. 1967.
Soil management in India. Asia Publishing House, Bombay.
Binet, F., and P. Trehen. 1992. Experimental microcosm study of the role
of Lumbricus terrestris (Oligochaeta; Lumbricidae) on nitrogen
dynamics in cultivated soils. Soils and Biochemistry 24:1501–1506.
Black, C. A., D. D. Evans, L. E. Ensminger, J. L. White, and F. E. Clark,
editors. 1965. Methods of soil analysis. Part 1. Physical and mineral-
ogical properties, including statistics of measurement and sampling.
Agronomy Series No. 9 American Society for Agronomy, Wisconsin.
Butt, K. R., J. Frederickson, and R. M. Morris. 1995. An earthworm culti-
vation and soil inoculation technique for land restoration. Ecological
Engineering 4:1–9.
Chadha, Y. R. 1985. The wealth of India (raw materials). Volume I A.
Publication and Information Directorate, CSIR, New Delhi.
Chadha, Y. R. 1989. The wealth of India (raw materials). Volume X SP-W.
Publication and Information Directorate, CSIR, New Delhi.
Chaudhauri, P. S., and G. Bhattacharjee. 1999. Earthworm resources of
Tripura. Proceedings of National Academy of Science, India 69(B),
II:159–170.
Coelho, F. E. 1990. Rehabilitation of earthworms on iron mine sites of
Goa. Ph.D. thesis. Goa University, Goa, India.
Curry, J. P. 1988. The ecology of earthworms in reclaimed soils and their
influence on soil fertility. Pages 251–261 in C. A. Edwards and E. F.
Neuhauser, editors. Earthworms in waste and environmental man-
agement. SPB Academic Publishing, The Hague.
Curry, J. P., and K. E. Boyle. 1995. Restoring soil fauna in reclaimed land,
with particular reference to earthworms in cut over peat. Acta Zoo-
logica Fennica 196:371–375.
Curry, J. P., D. Byrne, and K. E. Boyle. 1995. Earthworm population of a
481
Nutrient Mineralization by P. corethrurus on Iron Ore Mine Wastes
winter cereal field and its effects on soil and nitrogen turnover. Biol-
ogy and Fertility of Soils 19:166–172.
Daniel, O. 1991. Leaf-litter consumption and assimilation by juveniles of
Lumbricus terrestris L. (oligochaeta, Lumbricidae) under different
environmental conditions. Biology and Fertility of Soils 12:202–208.
Edwards, C. A., and J. R. Lofty. 1977. Biology of earthworms. Chapman
& Hall, London.
Emmerling, C., and D. Paulsch. 2001. Improvement of earthworm (Lum-
bricidae) community and activity in mine soils from open cast coal
mining by the application of different organic waste materials. Pedo-
biologia 45:96–407.
Ganihar, S. R. 1990. Impact of mining on the faunal composition of Goa.
Ph.D. thesis. Goa University, Goa, India.
Ganihar, S. R. 1996. Earthworm distribution with special reference to
physico-chemical parameters. Proceedings of Indian National Sci-
ence Academy B 62:11–18.
Ganihar, S. R. 1998. Early successional pattern of faunal recolonization on
rehabilitated iron mine sites at Sanquelim, Goa. Tropical Ecology
39:255–272.
Haber, A., and R. P. Runyon. 1973. General statistics. 2nd edition. Addi-
son-Wesley Publishing Co., London.
Haimi, J., and M. Einbork. 1992. Effects of endogenic earthworms on soil
processes and plant growth in coniferous forest soil. Biology and
Fertility of Soils 13:6–10.
Haimi, J., and V. Huhta. 1990. Effects of earthworms on decomposition
processes in raw humus forest soil: a microcosm study. Biology and
Fertility of Soils 10:178–183.
Hendriksen, N. B. 1990. Leaf-litter selection by detritivore and geopha-
gous earthworms. Biology and Fertility of Soils 10:17–21.
Julka, J.M. 1993. Distribution pattern in Indian earthworms. Pages 27–31
in The Zoological Survey of India, ed. Earthworm resources and
vermiculture. Zoological Survey of India.
Kale, R. D. 1998. Earthworm—Cinderella of organic farming. Prism
Books Pvt Ltd., Bangalore, India.
Kale, R. D., and R. V. Krishnamoorthy. 1980. The calcium content of the
body tissues and castings of Pontoscolex corethrurus (Annelida, Oli-
gochaeta). Pedobiologia 20:309–315.
Krishnamoorthy, R.V., and S. Ramachandra. 1988. Population structure
of earthworms in woodlands Karnataka. Proceedings of the Indian
Academy of Sciences (Animal Science) 97:335–362.
482
Krishnamurthi, A. 1969. The wealth of India (raw materials). Volume VII.
Publication and Information Directorate CSIR, New Delhi.
Lee, K. E. 1985. Earthworms: their ecology and relationships with soils
and land use. Academic Press, Sydney, Australia.
Minitab. 1991. Minitab reference manual release: 8.3 for PC Minitab Inc.
State College, Pennsylvania.
Perur, N. G., C. K. Subramanian, G. R. Muhr, and H. E. Ray. 1972. Soils
fertility evaluation to help Indian farmers. Mysore State Depart-
ment of Agriculture, Bangalore, India.
Petrusewicz, K., and A. Macfadyen. 1970. Productivity of terrestrial ani-
mal principles and methods. IBH Handbook No.13. International
Biological Program. Blackwell Scientific Publications, London.
Phondke, G. P. 1992. The wealth of India (raw materials). Volume 3. Pub-
lication and Information Directorate CSIR, New Delhi.
Ray, A. K., M. R. Rairar, and S. C. Pande. 1979. Manual of procedures for
chemical and industrial analysis of ore mineral and ore dressing
products. Government of India Publications, New Delhi.
Saetre, P. 1998. Decomposition, microbial community structure and
earthworm effects along a birch-spruce soil gradient. Ecology 79:
834–846.
Satchell, J. E. 1967. Lumbricidae. Pages 259–322 in A. Burges and F. Raw,
editors. Soil biology. Academic Press, London.
Shastri, B. N. 1989. The wealth of India (raw materials). Volume III, D-E.
Publication and Information Directorate CSIR, New Delhi.
Snedecor, G. W., and W. G. Cochran. 1961. Statistical methods. Allied
Pacific Private Limited, Bombay.
Sokal, R. R., and F. J. Rohlf. 1995. Biometry. The principles and practice
of statistics in biological research. 3rd edition. W. H. Freeman and
Company, San Francisco.
Tian, G., L. Brussard, and B. T. Kang. 1995. Breakdown of plants residues
with contrasting chemical composition under humid tropical condi-
tions: effects of earthworms and millipedes. Soil Biology and Bio-
chemistry 73:277–280.
Van Gestel, C. A. M., E. M. Driven-Van Breemen, and R. Baerselman.
1992. Influence of environmental conditions on the growth and re-
production of earthworm Eisenia andrei in an artificial soil substrate.
Pedobiologia 36:107–120.
Vimmerstedt, J. P., and J. H. Finney. 1973. Impact of earthworm intro-
duction on litter burial and nutrient distribution in Ohio strip mine
spoil banks. Soil Science Society of America Proceedings 37:388–391.
Restoration Ecology DECEMBER 2003