Journal of Geological Society of Sri Lanka Vol.

15 (2013), 99-110

OBSERVATIONS AND INTERPRETATIONS ON PAST MICROBIAL

ACTIVITIES AT EPPAWALA PHOSPHATE DEPOSIT
N. DEEPAL SUBASINGHE

Institute of Fundamental Studies, Hantana Road, Kandy, Sri Lanka

E-mail: [email protected]

ABSTRACT

An extensive, economically viable phosphate deposit exists at Eppawala in the north

central Sri Lanka. Textures and certain microscopic features found in this ore body had
led to the speculation of a sedimentary origin for this deposit, although the parent rock is
of igneous origin. Detailed investigations reveal that this deposit is a result of
weathering followed by in-situ diagenesis. This phoscrete-type phosphate deposit has
been developed on an apatite-rich carbonatite body. In certain sections of the ore,
microbial activities were detected under the scanning electron microscope as well as the
light microscope. These microbial activities might have started and ceased during the
formation of secondary phosphates. Currently micro-organisms are preserved in cavities
and silicified areas of the secondary phosphatic matrix. Morphology and the dimension
of different microbial components point towards the presence of more than one group of
microbes. This paper attempts to categorise some of the observed microbes as well as
understand the role played by them, using photomicrographic and mineralogical
evidences.

Keywords: Eppawala, apatite, carbonatite, microbial activity, Sri Lanka

INTRODUCTION sedimentary phosphate deposit. The term

“Phosphorite” is generally used for sedimentary
Eppawala phosphate deposit, located in the
phosphates of marine origin. However, the
Anuradhapura district of Sri Lanka, about 200
phosphate deposits created by terrestrial
km from Colombo, was discovered in 1971
processes, such as in-situ weathering and
(Jayawardana, 1976). This deposit exists as a
diagenesis can also be referred to as phosphorites
dense weathering profile underlain by an apatite
(sensu-lato) despite their original source.
containing carbonate rock. The ore bodies at
Dahanayake and Subasinghe (1988, 1989a,
Eppawala contain up to 42% P 2O5, and citric
1990a, 1991) used the term phosphorite to
acid solubility of different components varies
describe the phosphate deposit at Eppawala in
from 4 to 6%. Owing to intense weathering of the
this context. Eppawala deposit was formed by
apatite-bearing carbonate rock, a phosphate
relative accumulation of primary apatite and
deposit has been formed by relative accumulation
diagenesis of secondary phosphates. However,
of the primary apatite crystals, together with the
since those processes took place in terrestrial
secondary products derived from them
settings rather than marine or lacustrine, it is
(Dahanayake and Subasinghe, 1988; 1989a;
more appropriate to call the deposit as a
Subasinghe 1998).
phoscrete-type phosphate deposit (Dahanayake
and Subasinghe, 1991; Subasinghe, 1998).
The Eppawala phosphate deposit, which is a
secondary deposit formed over a parent
carbonatite rock display some features of a
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In many phosphate deposits in the world, Microbial activities were observed in the
especially in sedimentary ‘phosphorite’ deposits, secondary matrix of the phosphate rock at
microbial processes have played a significant Eppawala. Morphological information obtained
role. According to Hirsch et al. (1995), microbial through electron microscopic observations are
rock degradation is independent of climate, or the main data available on them.
location, or annual seasons. Dahanayake and
Krumbein, (1985) studied the microbial mats and Discovery of microbes in the phosphatic saprolite
emphasised their importance in the formation of prompted a question about the role of the
phosphorites. Soudry and Champetier (1983), microbes. It is necessary to understand whether
reported the microbial processes in the Negev the texture of the saprolite was significantly
phosphorites in southern Israel. Soudry (1987) affected by the microbes (dissolving or forming
presented a detailed account on the same deposit, phosphate minerals and textures) or, whether
and discussed the ultra-fine structures and microbes are playing a passive role and only
possible role of biological activities in the inhabit the areas favourable for them without
genesis of phosphorite. Comparing the structures considerably affecting the already existing
presented by the above workers, Dahanayake and textures.
Subasinghe (1988; 1989b) also suggested some
possible influence of microbial activities in the In this paper, the author presents strong evidence
formation of the terrestrial phosphate deposit at on the presence of microbes and their activities in
Eppawala, Sri Lanka. Soudry (1992) presented the Eppawala Phosphate deposit with the
further analogy on phosphorites from Negev, intention of understanding the nature and the
Israel. The numbers, growth rates and roles they would have played in bringing the
productivity of heterotropic bacteria in sediments phosphate deposit to its current state. This is a
of the continental-margin of Eastern Australia tribute to Prof. Kapila Dahanayake who put
were determined and discussed by Moriarty et al. forward the possibility of the existence of
(1991). Reimers et al. (1990) also presented microbes in the Eppawala phosphate more than 2
evidence for phosphate mineralisation by decades ago (Dahanayake and Subasinghe, 1988;
microbial mat formation. They studied and 1989b).
compared the present day microbial mats with
those found fossilised in Monetary Formation, GEOLOGICAL SETTING AND
California. Working on phosphatic hardgrounds PREVIOUS STUDIES
and stromatolites in Central Greece, The apatite-bearing carbonate rocks at Eppawala,
Pomonipapaioannou and Solakius (1991) occur as massive, discontinuous bodies in a
considered that phosphate was supplied to the Precambrian, high-grade metamorphic terrain.
pelagic sediment by a bacterially controlled Based on the field observations, mineralogy and
precipitation from interstitially circulated petrology, this carbonate rock has been assigned
solutions. an igneous origin, and thus called carbonatite
(Jayawardana, 1976). Later, geochemical work
On the other hand, there are many studies including oxygen and carbon isotopic studies
confirming that microbes, especially those found confirmed the igneous origin Subasinghe 1998;
in terrestrial environments, can solubilise the Weerakoon et al., 2001; Pitawala et al., 2003;
phosphate, thus increasing the availability of Manthilake et al., 2008; Pitawala and
phosphate to the plants (e.g. Panhwar et al., Lottermoser, 2012). Therefore, the term
2012; Khan et al., 2009; Ivanova et al., 2006). In ‘carbonatite’ is used hereafter to indicate the
a petrological context, these microbes have a parent carbonate rock with primary apatite
destructive effect on the phosphatic rock, rather crystals. The lithological units within the
than constructive. carbonatite complex could be grouped into those
of the basement and carbonatite. The economic
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phosphate deposit has been developed as a Secondary material mainly consists of carbonate-
secondary weathering profile over the parent and iron-phosphate varieties (Dahanayake and
carbonatite rock. Very large primary apatite Subasinghe, 1989; Subasinghe 1998) with relict
crystals of up to 1 m in length can be observed primary apatite, magnetite, mica and few other
occasionally in the parent carbonate rock. These minerals. Formation of secondary phosphate
primary apatite grains are large, yellowish green minerals and the observed textures have been
crystals with subhedral to euhedral shape. studied in detail by previous researchers
(Dahanayake and Subasinghe, 1988a; 1988b;
As shown in Figure 1, the phosphate deposit at 1989a; 1989b; 1989c; 1990a; 1990b; Subasinghe
Eppawala is located close to the boundary 1998). Further, Subasinghe (1998) stressed the
between Wanni and Highland complexes. importance of iron in the process of formation of
Granitic gniesses, granites, quartzites, secondary phosphates and discussed the potential
charnockitic gneiss, hornblend-biotite gneiss and use of this natural phenomenon in material
pegmatitic rocks can be observed in the science applications (Subasinghe, 2012).
surrounding area.
Field observations indicate that the phosphatic
In the carbonate rock (the parent rock), calcite, saprolite profile at Eppawala more or less
dolomite and apatite are major minerals, whereas resembles a supergene weathering profile, having
magnetite, spinal, pyrite, and ilmenite occur as loose lateritic material on the top and more
important accessories. In addition, forsterite, consolidated material towards the bottom. It also
phlogopite, enstatite, magnesite, martite, indicates that the phosphatic saprolite is more
diopside, tremolite and few other minerals also resistant to chemical weathering than its parent
occur in smaller amounts. rock or the siliceous saprolite. Borehole data
show that the phosphatic saprolite is always

Figure1. Location and Geology of Eppawala Phosphate deposit (Subasinghe, 1998, modified after
Jayawardena, 1976).
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overlain by weathered material; either phosphatic composition to confirm the identity of a

lateritic soil or siliceous lateritic soil, or, siliceous suspected mineral grain or feature.
saprolite. Also, at no place was siliceous saprolite
or laterite found underneath the phosphatic Some samples were prepared as polished sections
saprolite cover. These observations are explained so that they could be observed under SEM as
if supergene weathering is considered. well as the light microscope, consequently.

MATERIALS AND METHODS RESULTS AND DISCUSSION

Samples were collected from different regions of Parent material, the apatite-bearing carbonatite
the Eppawala phosphate deposit to represent rock, is shown in Figure 2. Note the white
different weathering stages and varying coloured unweathered carbonatite matrix at the
mineralogy. Partially weathered carbonatite bottom, and the development of brown coloured
samples as well as secondary phosphate with secondary material at the top part that is exposed
detrital primary apatite crystals were included in to meteoric weathering. Some meteoric water
the sampling. More interesting secondary penetrates into the rock through the fissures and
textures with well preserved and relatively less fractures of the rock. They leave a brownish stain
disintegrated samples were found towards the in the rock. Primary apatite crystals are more
southern end of the deposit. Fresh samples were resistant to weathering than the carbonate matrix,
stored in sample bags and brought to the thus getting accumulated when the carbonate
laboratory. Samples were cleaned, dried, surface matrix is partially removed and converted to
sterilised using UV light, and stored in a cold secondary matrix, which is usually light to dark
room to avoid any biological contamination, until brown in colour.
they were used for analysis. Sub samples were
selected from well consolidated samples for SEM
studies.

Handspecimen Studies
Handspecimens were inspected using a hand
lens. Certain samples were cut and polished to
show specific features photographically whereas
some samples were observed with their naturally
broken surfaces.

Scanning Electron Microscopy (SEM)

Selected samples were broken in to small pieces
using gentle fracturing so that natural features
will be clearly seen. These samples were
mounted on metal sample stubs and coated with
either gold or carbon. Samples were examined in
JEOL JXA-840 scanning microanalyser and
JEOL JSM5300 scanning electron microscope at
20KV accelerating voltage and usual settings.
Figure 2. Partially weathered carbonatite rock
Magnification range used for the analyses is from
with small primary apatite crystals (pale green),
10 x to 10,000 x. Qualitative elemental analysis
dissolution cavities and partially developed
was carried out at certain points of interest, using
secondary material (brown colour). Width of the
energy dispersive X-ray analysis. This was
photograph is about 11 cm.
mainly used to identify the major element

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Secondary materials appear as a fine-grained However, microbes are firmly attached to the
matrix to the naked eye. However, under the apatite crystals and spread their filamentous
SEM, they display crypto-crystalline hexagonal material across them indicating their affinity for
apatite crystals, clay minerals, detrital silicate phosphate.
minerals and apatite grains and most
interestingly, some microbial activities at certain At advanced stages of microbial activities, apatite
places, as shown in Figure 3. crystals had been dissolved from all directions, as
shown in Figures 5-7. Diameter of the bacillus
Close observations show that microbial activities material is in the order of 1 µm on average, while
are in several stages of development. As shown that of the filaments is less than 0.5 µ.
in Fig. 4, at initial stages of microbial activities,
bacillus shaped connected chains of microbes are
observed on the apatite crystals. At this stage,
apatite crystals show little or no deterioration.

Figure 3. Microscopic
apatite crystals under
SEM. No microbes
are seen in this
sample and apatite
crystals are intact
with no indication of
dissolution

Figure 4. Initial stages of

some microbial activities
in the secondary matrix of
Eppawala phosphate.
Note the bacterial
material and their
filaments attached to the
hexagonal apatite crystals

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Figure 5. Microbial activities (arrows) cause apatite crystals to dissolve from sideways,
i.e. parallel to prism faces. Compare with Figure 3

Figure 6. Hexagonal cross-sectional view of apatite crystals showing the inter-planar

dissolution, probably due to secretions resulting from microbial activities (arrows)

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A B
A B

C D

C D

E F

E F
Figure 7. Filamentous bacterial colonies in advanced stages of microbial activity. B and
C shows enlarged views of the photo A. D-Partially dissolved apatite crystals and some
microbes. E and F show thick microbial colonies. Note the size of the microbes and their
filaments
Figure 7 shows a collection of different views of in dimension. The presence of similar size
microbial activities in the secondary phosphatic filaments suggests that bacterial origin and not
matrix. It is clear that the size of the bacillus- fungal origin. This indicates that the microbial
shaped interconnected microbial material have material commonly observed as secondary
diameters smaller than 1 µm whereas filaments material at Eppawala phosphate deposit are
are also show similar dimensions (Fig. 7 A and filamentous bacteria (actinobacteria).
B). Such dimensions are typical of bacteria.
Usually, fungal materials are several times larger
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Figure 8 shows a special feature observed in the of a detached and fossilised fungal spore. In fact,
secondary matrix. However, this feature was perfect geometrical shape and regular distance
observed in an area rich in siliceous material. The between circular grains are characteristics of
morphological features of Figure 8 may suggest a diatoms (Barber and Hayworth, 1981; Mann,
detached fungal spore. However, for a detached 1999). The existence of this material in a silica
fungal spore to survive the sample preparation rich mineral is another positive factor to support
procedure as well as the high vacuum and high that this could be a valve view of interconnected
voltage in the SEM, it should be a hard material silica-secreting diatoms.
firmly attached to the substrate, the siliceous host
mineral. Further, it should be fossilised, since the When the role of the microbes is considered,
fungal spores do not have hard cell walls. Also, there are two possibilities to consider: Whether
the fungal spores do not have perfectly circular (a) microbes are playing an active role in
shapes or long, relatively thin stems between producing secondary phosphates, as in certain
each spore. Further, no fungal spores are marine deposits, or (b) if they are solubilising
observed in any of the samples despite the fact and consuming some phosphate. The extent of
that some of the samples are densely populated the microbial activities and the overall effect on
with different types of microbes. Significant the phosphate deposit is the other factor to be
morphological and dimensional differences, as considered.
well as other factors as mentioned above strongly
suggest that the microbes shown in Figure 8 Extensive work carried out by the author as well
cannot be displaced conidiophores. as other researchers on the deposit helped
establishing the present state of the phosphate
In addition, microprobe analysis of the host deposit (Dahanayake and Subasinghe, 1988,
mineral as well as the microbial feature reveals 1989a, b, 1990b, 1991a; Subasinghe, 1998;
high silica contents, suggesting that they are Weerakone et al., 2001; Pitawala et al., 2003).
made of quartz or other silica rich mineral. Geochemical, mineralogical and
Therefore, it is unlikely that Figure 8 is an image sedimentological evidence supported by isotope

Figure 8. Another type of microbial material observed in the matrix. Note the perfect circular
shape and regular distances maintained between each circular feature. Area marked with a
square was analysed for elemental composition and found that it is predominantly silicon. A
sketch of the typical conidiophores is shown on the right for comparison. Significant
morphological differences are evident.
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results strongly suggest that the secondary Other possibility is that the above may be
phosphate material had formed as a result of microbes with hard cell walls. It is a known fact
meteoric weathering acting on the parent that desmids and diatoms secrete silica into their
carbonatite rock containing primary apatite cell walls.
crystals.
Remnants of possible microbial colonies were
As there are good explanations for the formation occasionally observed in thin sections, as shown
of secondary matrix through inorganic (non- in Figure 9. These thin sections show the
biological) weathering and in-situ diagenetic hexagonal outlines or ‘ghost structures’ of
activities (Subasinghe, 1998; Tazaki et al., 1986; previously existing apatite crystals. Currently
1987), it can be deduced that the microbial silica is filled in the outlines, which appear as
activities do not play a critical role in producing hexagonal cavities under plane polarised light
and shaping the phosphate deposit at Eppawala. (PPL), while secondary quartz is observable
under crossed polars (Figure 9 B). These
However, it is still interesting to study these observations confirm that the microbes have
microbial activities; firstly, this is the first time played an active role in removing phosphates
such activities were recorded in Sri Lankan when the silica was in abundance. Silicification
phosphates. Dahanayake and Subasinghe (1988) process that followed the microbial activities has
suggested microbial involvement in the preserved the visible evidence of microbial
formation of secondary phosphates, although colonies that existed.
they could not provide direct strong
photomicrographic evidence for their claims at According to available evidence, it is highly
the time, due to technological limitations. possible that microbes observed in Figure 8 were
Secondly, if the observed microbes are fossilised a diatomaceous species. The microbes might
ones, and if their age can be determined, have entered the rock with rain water passing
identifying them would help understand the through pores and fractures developed after the
environmental conditions at the time of their initial stages of weathering and settled in the
formation. Thirdly, if the role of the microbes is small cavities of the secondary matrix.
the solubilisation of the phosphate, there is a Konhauser (1997) reported iron mineralisation by
possibility of producing biofilms utilising the microbes. These bacterial colonies can leave dark
phosphate solubilising microorganisms isolated brown stains in the rock, even after their death.
from this area to increase the availability of Some dark brown stains found in the silicified
phosphate. areas (e.g. Figure 9) of the Eppawala phosphate
could be considered as similar stains left by
Unless specially prepared, biological material microbial colonies.
such as soft tissues containing water cannot
survive under high-vacuum and high voltage Furthermore, although not commonly
electron beams used in the scanning electron encountered, larger filamentous material could
microscopes. Any of the observed features did also be observed in certain samples. As shown in
not show deformation or destruction under the Figure 10, these materials have diameters of the
electron beam, confirming that they were well- order of 10 µm (10-30 µm). This dimension is
preserved hard material. comparable with that of the common fungal
filaments. Backscattered images (Figure 10-B)
This provides two possibilities: Possibility one is show darker colour in these filaments indicating
that these were bacteria-like soft material that that they consisted of lighter elements, possibly
lived, died and fossilised. Since bacteria have no carbon or silicon, compared with the surrounding
hard cell walls, they can only be preserved as phosphatic matrix.
fossils.

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A B

Figure 9. Hexagonal outlines of previously existing apatite crystals preserved in dark

secondary material. A.- hexagonal shaped cavities-ghost structures-left by removal
of primary apatite crystals (Under PPL). B. Under CPL quartz infillings can be
observed. Some dark filamentous materials inside the hexagonal outlines indicate
possible existence of microbes during the removal of apatite.
Actino-bacteria were reported in a variety of
environments including rocks and building monuments of six historic Scottish buildings and
materials. Suihkoa et al. (2007) reported hyaline ruins. Morphological features of some of the
fungal species, diatom algae as well as microbes reported by them have close
filamentous bacterial growths on samples taken resemblance to those found at Eppawala samples.
from the inner and outer surfaces of stone SEM images of the microbial mats reported by
Reimers et al. (1990) also closely resemble some
of those found at Eppawala.

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 Journal of Geological Society of Sri Lanka Vol. 15 (2013), 99-110

A B
Figure10. Fungal filaments in the phosphatic matrix. A- Secondary emission image; B- under Back
Scattered Electrons (BSE), filaments appear darker. Note the scale
Secondary matrix contained detrital primary solubilising the phosphates, rather than initiating
apatite crystals as well as crypto-crystalline any precipitation process.
grains. When the conditions were favourable, CONCLUSIONS
microbes could have acted to solubilise the
Microbial activities detected at the Eppawala
apatite crystals while increasing the
phosphate deposit are relatively younger and play
concentration of silica through their cell wall
a passive role when it comes to the formation of
secretions. It is quite possible that these silica
new mineral textures. The extent of the activities
secretions had acted as nucleation material for
is not fully evaluated; however, the observations
the silicification process at Eppawala.
indicate that they are confined to certain sectors
of the deposit. Dimensions and morphological
Although the shape of the microbial filaments
characters of the microbes indicate the presence
resembles those of fungi, the size factor suggests
of several species including action-bacteria,
that these are filamentous bacteria
diatoms and fungi.
(actionobacteria) rather than fungi. Actino-
bacteria (filamentous bacteria) require carbon
dioxide for their growth (Round et al., 1990).
ACKNOWLEDGEMENTS
Weathering of the carbonate matrix releases CO2, Postgraduate Research Institute for
which could facilitate the growth of such Sedimentology (PRIS) at University of Reading,
microbes. Therefore, it is possible that the UK provided the SEM facilities for this study.
microbes observed in Eppawala samples are Professors K. Dahanayake and V.P. Wright are
actinobacteria. thanked for their support and encouragements in
various ways. Discussions with Prof. S.A.
Natesan and Shanmugasundaran (1989) Kulasooriya and Dr. R. Ratnayake, were
experimented with cultured cyanobacteria under extremely useful in recognising different types of
controlled conditions to study their extracellular microbes observed and in understanding their
phosphate solubilisation, especially in the soil role.
environment. Most of the reported microbes in
terrestrial environments solubilise the phosphate
REFERENCES
rather than causing precipitation. Therefore,
together with the microscopic evidence, it can be Barber, H.G. and Hayworth, E.Y. (1981). Guide to the
Morphology of the Diatom Frustule. Freshwater
safely concluded that the role of the microbes at Biological Association, pp.112.
the Eppawala phosphate deposit is towards Dahanayake, K. and Krumbein, W.E. (1985).
Ultrastructure of a microbial mat-generated

109
N. Deepal Subasinghe/ Observations and Interpretation on the Past Microbial Activities at Eppawala Phosphate Deposit

phosphorite, Mineralium Deposita. Vol. 20(4), Panhwar, Q. A., Radziah O., Naher U. A., Zaharah A.
p.260-265. R., Sariah M. and Mohd Razi I. (2012). Root
Dahanayake, K. and Subasinghe, S.M.N.D. (1988). colonization and association of phosphate-
Development of Recent Stromatolite Structures solubilizing bacteria at various levels of triple
and Phosphatic Enrichment in Precambrian supper phosphate in aerobic rice seedlings.
Marble of Sri Lanka. Economic Geology, Vol. 83 African Journal of Microbiology Research. Vol.
(7), p. 1468-1474. 6(10), p. 2277-2286.
Dahanayake, K. and Subasinghe, S.M.N.D. (1989a). A Pitawala, A. and Lottermoser, B.G. (2012).
Modern Terrestrial Phosphorite - an Example Petrogenesis of the Eppawala carbonatites, Sri
from Sri Lanka. Sedimentary Geology, Vol. 61, Lanka: A cathodoluminescence and electron
p. 311-316. microprobe study, Mineralogy and Petrology,
Dahanayake, K. and Subasinghe, S.M.N.D. (1989b). Vol. 105(1-2), p. 57-70.
Secondary Phosphate Mineralization in a Karstic Pitawala, A., Schidlowski, M., Dahanayake K. and
Environment in Central Sri Lanka. Mineralium Hofmeister, W. (2003). Geochemical and
Deposita , Vol. 24, p. 169-175. petrological characteristics of Eppawala
Dahanayake, K. and Subasinghe, S.M.N.D. (1990a). phosphate deposit, Sri Lanka. Mineralium
Variable Mineralogy and Solubility of a deposita, Vol.38(4), p. 505-515.
Terrestrial Phosphorite Resulting from Pomonipapaioannou, F. and Solakius, N. (1991).
Differential Weathering Phenomena - an Phosphatic hardgrounds and stromatolites from the
Example from Sri Lanka. Chemical Geology, Vol. limestone shale boundary section at Prossilion
84(1/4), p. 42-44. (Maastrichtian-Paleocene) in the Parnassus-Ghiona
Dahanayake, K. and Subasinghe, S.M.N.D. (1990b). zone, Central Greece. Palaeogeography
Formation of Phosphatic Coated Grains and Palaeoclimatology Palaeoecology, Vol. 86(3-4), p.
Peloids by Grain Diminution of Precambrian 243-254.
Apatite Crystals. J. Geol. Soc. India., Vol. 35, Reimers, C.E., Kastner, M., and Garrison, R.E.
p.189-197. (1990). The role of bacterial mats in phosphate
Dahanayake, K. and Subasinghe, S.M.N.D. (1991a). mineralization with particular reference to the
Heterogeneity of the Eppawala Phosphate Monterey Formation. In: Burnett, W.C., and
Deposit. Ceylon Journal of Science., Vol. 1, p.1- Riggs, S.R. (eds.), Phosphate Deposits of the
7. World, Vol. III: Neogene to Modern
Dahanayake, K. and Subasinghe, S.M.N.D. (1991b). Phosphorites. Cambridge University Press, p.
Mineralogical Petrological and Solubility 300-311.
Variation of the Eppawala Phosphate Deposit in Round, F.E., Crawford, R.M. and Mann, D.G. (1990).
Sri Lanka – a Case for Selective Mining for The diatoms. Biology and morphology of the
Fertilizers. Fertilizer Research. , Vol. 28, p. 233- genera. Cambridge University Press, pp. 747.
238. Soudry, D. and Champetier, Y. (1983). Microbial
Hirsch, P., Eckhardt, F.E.W. and Palmer, R.J. (1995). processes in the Negev phosphorites (Southern
Methods for the study of rock-inhabiting Israel). Sedimentology., Vol. 30, p. 411-423.
microorganisms: A mini review. Journal of Soudry, D. (1987). Ultra fine structures and genesis of
Microbiological Methods, Vol. 23( 2), p.143-167. the campanian Negev high-grade phospharites
Ivanova, R., Bojinova, D. and Nedialkova, K. (2006). (Southern Israel). Sedimentalogy, Vol. 34, p.
Rock phosphate solubilisation by soil bacteria. 641-660.
Jour. Univ. Chemical Technology and Soudry, D. (1992). Primary Bedded Phosphorites In The
Metallurgy, Vol. 41(3), p. 297-302. Campanian Mishash Formation, Negev, Southern
Jayawardana, D.E. de S. (1976). The Eppawala Israel. sedimentary Geology, Vol. 80(1-2), p.77-88.
carbonatite complex in North – West Sri Lanka Subasinghe, N.D. (1998). Formation of a phosphate
(Ceylon). Geological Survey Department, deposit through weathering and diagenesis – an
Economic Bulletin, No. 3. example from Sri Lanka. PhD Thesis (Unpubl.)
Khan, A.A., Jilani, G., Akhtar, M.S., Naqvi, S.M.S. University of Reading, UK.
and Rasheed, M., (2009). Phosphorus Subasinghe, N.D. (2012). Electron microscopic
Solubilizing Bacteria: Occurrence, Mechanisms studies on phosphate binding processes in the
and their Role in Crop Production, J. Agric. Biol. presence of iron. Advanced Materials
Sci., Vol. 1(1), p. 48-58. Research, Vol. 343-344, p. 307-311.
Konhauser, K. O. (1997). Bacterial iron Suihkoa, M.L. Alakomia, H.L., Gorbushinab, A.,
biomineralisation in nature. FEMS Microbiology Fortunec, I., Marquardtb, J. and Saarelaa, M.
Reviews, Vol. 20, p. 315–326. (2007). Characterization of aerobic bacterial and
Mann, D.G. (1999). The species concept in diatoms. fungal microbiota on surfaces of historic Scottish
Phycologia, Vol.38, p. 437-495. monuments. Systematic and Applied
Manthilake, M.A.G.M., Sawada, Y. and Sakai, S. Microbiology, Vol. 30(6), p. 494–508.
(2008). Genesis and evolution of Eppawala Tazaki, K., Fyfe, W.S. and Dissanayake, C.B. (1986).
carbonatites, Sri Lanka, Journal of Asian Earth Weathering of Phosphatic Marble to Exploitable
Sciences, Vol. 32, p. 66–75. Apatite Deposit, Sri Lanka. Applied Geochemistry
Moriarty, D.J.W. Skyring, G.W. O'Brien, G.W. and Vol. 1, p. 287-300.
Heggie, D.T. (1991). Heterotrophic bacterial Tazaki, K., Fyfe, W.S. and Dissanayake, C.B (1987).
activity and growth rates in sediments of the Weathering of Apatite under Extreme conditions of
continental margin of eastern Australia, Deep Sea Leaching. Chemical Geology, Vol. 60, p. 151-162.
Research Part A. Oceanographic Research Weerakone, M.W.K., Miyazaki, T. Shuto, K. and
Papers, Vol. 38(6), p. 693-712. Kagami, H. (2001). Rb-Sr and SM-Nd
Natesan, R. and Shanmugasundaran, S. (1989). Geochronology of the Eppawala metamorphic
Extracellular phosphate solubilization by the rocks and carbonatite, Wanni Complex, Sri
cyanobacterium Anabaena ARM310. Biosci., Vol. Lanka. Gondwana Research, Vol. 4(3), p. 409-
14(3), p. 203-208. 420.
110