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ISSN:2682-4043
DOI:10.36462/H.BioSci.202505

Research Article
Open Access

1 Department of Biotechnology Research,

Kyaukse, Ministry of Science and Technol-

ogy, Myanmar

* To whom correspondence should be
addressed: sakurasan2007@gmail.com

Editor: Muhammad M. Adeel Arthritis Clinical
Immunology program Oklahoma Medical Research
Foundation, Oklahoma City, United States.

Reviewer(s): Alsayed Alsoudy, Mohammed VI
Polytechnic University, Ben Guerir, Morocco.

Wan-Atirah Azemin, School of Biological Sciences,
Universiti Sains Malaysia, Minden, Pulau Pinang,
Malaysia.

Received: March 25, 2025

Accepted: June 27, 2025

Published: July 26, 2025

Citation: Yu SS, Marr KT, Mon WW, Aye
KS. Molecular identification and screening the
antagonistic and cellulolytic activities of native strain
Bacillus subtilis isolated from Myanmar marine
sediment. 2025 July 26;8:bs202505

Copyright: © 2025 Yu SS et al.. This is an open
access article distributed under the terms of the Cre-
ative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in
any medium, provided the original author and source
are credited.
Data Availability Statement: All relevant data are
within the paper and supplementary materials.
Funding: This work was supported by the De-
partment of Biotechnology Research Department,
Kyaukse, Ministry of Science and Technology,
Myanmar, by the Department of Plant Medicine,
Kyungpook National University, Daegu, South Ko-
rea, and by the International Scholar Exchange Fel-
lowship program, Chey Institute of Advanced Stud-
ies.
Competing interests: The authors declare that they
have no competing interests.

Molecular identification and screening the antagonistic and cellulolytic
activities of native strain Bacillus subtilis isolated from Myanmar
marine sediment

San San Yu*1
><�, Khin Thae Marr1

>< �, Wai Wai Mon1
><�, Khin Swe Aye1

><

Abstract

In order to combat plant diseases and produce profitable agricultural crops, biocontrol
agents derived from bacteria offer sustainable and eco-friendly options. Numerous
fungal diseases have been successfully treated by some advantageous strains of
Bacillus subtilis. The bacterial strain used in this study, designated S1-10, was isolated
from a marine sediment sample. The antifungal qualities of the isolate were evaluated
against eight out of ten plant-pathogenic fungi. The highest cellulolytic activity of the
isolate was observed after four days of incubation. The association of the strain with
Bacillus subtilis was confirmed by phylogenetic analysis based on the sequences of
the gyrB and 16S rRNA regions. The NCBI has received strain S1-10, which has been
assigned accession codes OR708648 for 16S and PV007904 for gyrB.

Keywords: Bacillus subtilis, marine sediment, antifungal effect, cellulolytic activity, phyloge-

netic analysis

Introduction
Marine sediments represent one of the largest habitats on our planet. Their distinctive ecological

characteristics, such as elevated salinity, high pressure, and low oxygen levels, can trigger the activa-

tion of previously dormant genes in marine microbes. This process leads to the emergence of unique

microbes, enzymes, bioactive compounds, and specialized metabolic pathways that have adapted

to these specific environmental conditions. The microorganisms sourced from marine sediments

and their bioactive metabolites hold significant importance and present promising opportunities for

commercial development in sectors such as food, pharmaceuticals, chemicals, agriculture, envi-

ronmental protection, and human nutrition and health [1]. The unique living conditions found in

marine environments compared to terrestrial ones make marine microorganisms highly promising

candidates for producing innovative natural compounds [2]. Marine Bacillus isolates represent a

diverse collection of bacteria, exhibiting both phylogenetic and phenotypic variations within the

broader microbial world. These bacteria are widely distributed throughout the marine environment

and possess remarkable resilience, thriving in challenging conditions like high temperatures, pressure,

salinity, and pH levels.

Typically, Bacillus strains necessitate ample nutrients and space to achieve their maximum

growth rates, a competitive advantage they utilize against other organisms. The vastness of the ocean,

with its constant dilution, compels marine organisms to develop powerful bioactive compounds for

survival, either to outcompete rivals or to defend against smaller predators [3]. Numerous bioactive

compounds, such as cyclic peptides, glycopeptides, lipopeptides, and bacteriocins, are produced

by Bacillus species, which are well known for their wide spectrum of antibacterial properties [4].

Chemical pesticides are currently the main tool used by agricultural producers to control or prevent

crop diseases. The accumulation of these pesticide residues in food, soil, and aquatic environments

poses significant threats to human health, the environment, and ecosystems [5]. As a result, there is

a growing demand from consumers and environmental advocates to substitute chemical pesticides

with natural, eco-friendly microorganisms that have innovative modes of action for the sustainable

cultivation of crops [6]. In nature, biomass rich in lignocellulose and agricultural residues is widely

available as a source of carbon. Utilizing microorganisms or their enzymes for the biological

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https://creativecommons.org/licenses/by/4.0/
https://creativecommons.org/licenses/by/4.0/
mailto:sakurasan2007@gmail.com
https://orcid.org/0000-0003-2465-2866
mailto: khinthaemarrmm@gmail.com 
https://orcid.org/0009-0005-9920-9385
mailto:thetmin.wai333@gmail.com
https://orcid.org/0009-0006-1088-936X
mailto:khinsweaye.mdy@gmail.com
http://bioscience.highlightsin.org/


Yu SS et al., 2025 Molecular identification and screening the antagonistic and cellulolytic activities

Figure 1. Colony and microscopic morphology (100×) of (S1–10) isolate on LB

(Luria-Bertani) and N/A (Nutrient Agar)

pretreatment of lignocellulosic biomass provides an environ-
mentally friendly method for processing biomass. The use of cel-
lulases and hemicellulases to hydrolyze lignocellulosic biomass
results in the formation of monomeric sugars that can be easily
fermented by microorganisms [7]. Numerous studies have shown
that both bacterial and fungal species can produce cellulolytic
enzymes. Bacterial species, particularly those from the Bacillus
genus, have been predominantly utilized in industrial applica-
tions due to their ability to secrete a variety of hydrolytic enzymes
[8]. The cellulolytic enzymes derived from B. subtilis have been
extensively researched because of their significant production
capacity and usefulness in biomass valorization [9].

Bacillus subtilis is a catalase-positive, rod-shaped bacterium
that is Gram-positive. Like other members of the Bacillus genus,
it can produce an endospore that enables it to endure extreme
conditions such as high temperatures and dryness. B. subtilis
is regarded as the most extensively studied model organism for
investigating bacterial chromosome replication and cell differ-
entiation among Gram-positive bacteria [10]. Additionally, it is
frequently utilized as an industrial cell factory for synthesizing
vitamins, inositol, acetoin, hyaluronan, and various other chem-
icals. Research in agriculture has shown that incorporating a
suitable amount of B. subtilis can significantly enhance the hu-
mus and carbon levels in compost, thereby improving soil quality
and supporting crop growth [11].

Myanmar’s economy is largely based on agriculture, making
the agricultural sector crucial for business. Thus, enhancing agri-
cultural output, ensuring food safety, and improving soil quality
are fundamental strategies. The agricultural inputs currently in
use are primarily chemical in nature. Poor management practices
on farms and the incorrect application of agrochemicals have led

Figure 2. Detection of cellulase and optimum growth condition

to soil degradation and environmental harm. This study aims to
isolate and identify Bacillus subtilis from marine sediment and
to explore the effective antagonistic and cellulolytic properties of
this bacterial isolate for use as a biofungicide and in composting.

Materials and Methods
Sample collection and strain isolation

Strain S1-10 was isolated from a sample of wet sediment
taken from the sea near Panga Village, located in Thanbyuzayat
Township, Mawlamyine District, Mon State, Myanmar (15° 54′

18′′ N, 97° 43′ 31′′ E). Ten milliliters of a 0.85% saline solution
were used to suspend one gram of the marine sediment, which
was then shaken for half an hour at 30°C. After settling for
two hours, the sample solution was serially diluted (10−1 to
10−4). Using these dilutions, 100 µL was plated onto a 3% NaCl-
enriched nutritional agar medium and cultured for three days
at 30°C [12]. Repeated streaking on the same medium allowed
for the selection and purification of the colonies that emerged.
For future research, the pure culture of strain S1-10 was stored
at −80°C in a 20% (v/v) glycerol solution after being regularly
maintained and incubated at 30°C.

Screening of antagonistic activity and cellulolytic activity
Plant pathogenic fungi were generously supplied by Profes-

sor Dr. Hee-Young Jung from the Department of Plant Medicine
at Kyungpook National University in Daegu, South Korea. The
dual culture technique was employed to assess the efficacy of
a Bacillus strain against these fungi, utilizing Petri dishes mea-
suring (90 × 15 mm) filled with potato dextrose agar (PDA). In
this method, a 6 mm diameter circle containing a suspension of
conidia and mycelium, serving as the pathogen inoculum, was
placed at the center of the dish. The antagonist was positioned
3 cm away from the center. Each experiment was conducted in
triplicate, with control fungal colonies not exposed to the antago-
nists. The inoculated Petri dishes were incubated at 30oC for 14
days. After this incubation period, the inhibition zone between
the phytopathogen and antagonist colonies was observed on the
Petri dish [13].

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Yu SS et al., 2025 Molecular identification and screening the antagonistic and cellulolytic activities

The assessment of cellulolytic activity was conducted using
LB medium supplemented with 1% cellulose powder. Colonies
of the isolate were inoculated onto this medium and incubated
at room temperature for 48 hours. Following incubation, the
plate was treated with 0.1% Congo red for a duration of 15 to 20
minutes, then rinsed with 1 M NaCl for an additional 15 to 20
minutes. The presence of a clear zone surrounding the colony
was noted, indicating cellulase activity [14]. This transparent
area indicates the cellulolytic activity of the isolate. According
to Miller GL’s 1959 description, the 3,5-dinitrosalicylic acid
(DNS) method was used for the quantitative analysis of cellulase.
By adding 3 mL of DNS reagent—which is made up of 1 g
dinitrosalicylic acid, 16 g NaOH, 300 g potassium sodium tartrate,
and distilled water to a final volume of 1 L—to the 1 mL reaction
mixture, the enzymatic reaction was stopped. The optical density
of this mixture was then measured at 540 nm after it had been
boiled for five minutes in capped glass tubes and cooled in cold
water. D-glucose-based calibration curves were used to measure
the cellulase activity.

Analysis of Morphology and Physiology
A variety of agar media, such as nutrient agar (NA; Difco),

marine agar 2216 (MA; Difco), tryptic soy agar (TSA; Difco),
and Luria Bertani agar (LB; Difco), were used to assess the
growth of the isolated bacterial strain S1-10. On LB agar, the
temperature range for growth was determined at 4, 10, 15, 20,
25, 30, 37, 42, and 45 °C. Furthermore, after a 7-day incubation
period, the pH range for growth was evaluated on LB agar at pH
levels ranging from 4.0 to 10.0, in increments of 1.0 pH unit. To
adjust the pH of the medium, 1 M NaOH and HCl were used.
Additionally, growth was observed on nutrient agar after 7 days
in the presence of NaCl concentrations ranging from 1.0% to
10.0% (w/v), with 1.0% increments.

Phylogenetic Analysis
The HiGene Genomic DNA Prep Kit (Biofact, Daejeon, Ko-

rea) was used to extract the genomic DNA of strain S1-10 from
24-hour-old cultures grown on nutrient medium. The methodol-
ogy developed by Weisburg et al. was followed in the polymerase
chain reaction (PCR) procedure used in this study to amplify the
16S rRNA region. Primers 9F (5’-GAG TTT GAT CCT GGC
TCA G-3’) and 1512R (5’-ACG GCT ACC TTG TTA CGA
CTT-3’) were used in this investigation [15].

For phylogenetic analysis and species identification within
the Bacillus group, the DNA gyrase B subunit gene, gyrB, was
selected for examination. DNA sequencing was conducted using
degenerate primers for gyrB, specifically gyrB-F (5-GAA GTC
ATC ATG ACC GTT CTG-3) and gyrB-R (5-AGC AGG GTA
CGG ATG TGC GAG-3) [16]. The PCR product cleaning reagent
ExoSAP-IT (Thermo Fisher Scientific, Waltham, MA, USA) was
used to purify the amplified PCR products.

The full sequence of the 16S rRNA gene was assembled
using SeqMan software (DNASTAR, Madison, WI, USA). Man-
ual adjustment of gaps and terminal ends of the alignment was

Figure 3. A neighbor-joining phylogenetic tree constructed from 16S rRNA

gene sequences illustrates the evolutionary relationships of strain S1–10 and

related members of the Bacillaceae family. Nodes with bootstrap support values

exceeding 50% are marked. The strain that was isolated is highlighted in bold.

Heynadriskxia sporothermodurans M215 served as the outgroup. The scale bar

represents 0.0050 substitutions per nucleotide position.

performed with BioEdit version 5.0.6 to ensure accuracy and
completeness of the gene sequences. Identification of the phy-
logenetically closest sequences and calculation of pairwise se-
quence similarity values were conducted using the EzBioCloud
server and BLAST searches targeting the 16S rRNA gene.

Sequences of the 16S rRNA gene from related taxa were re-
trieved from GenBank, and multiple alignments were carried out
using the CLUSTALW software [17]. A phylogenetic tree was
constructed using the neighbor-joining method implemented in
MEGA 7 software [18]. Evolutionary distances were calculated
using Kimuras two-parameter model, and the reliability of the
tree topology was assessed through bootstrap analysis with 1,000
replicates.

Results and Discussions
Analysis of Morphology and Physiology

Colonies of the S1-10 strain grown on nutrient agar supple-
mented with 3% NaCl appeared flat, white, and were approx-
imately 3–5 mm in diameter after three days of incubation at
30 °C (Figure 1). Significant growth was observed on nutrient
agar, marine agar, TSA, and LB agar. The S1-10 strain demon-
strated the ability to grow in NaCl concentrations ranging from
1% to 10%, within a pH range of 4 to 10, and at temperatures
between 15 °C and 45 °C. Optimal growth conditions were de-
termined to be 3% NaCl, pH 7, and 30 °C. In this research, a
new strain of Bacillus subtilis, named S1-10, was extracted from

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Yu SS et al., 2025 Molecular identification and screening the antagonistic and cellulolytic activities

Figure 4. Neighbor-joining phylogenetic tree constructed from gyrB gene

sequences, illustrating the phylogenetic placements of strain S1–10 alongside the

Bacillus subtilis group. Bootstrap support values exceeding 70% are shown at

the nodes. The isolate is indicated in bold. Heynadriskxia sporothermodurans

M215 served as an outgroup. Bar, 0.050 replacements for each nucleotide

position.

marine sediment in Myanmar and evaluated for its antagonis-
tic characteristics against several phytopathogenic fungi. The
S1-10 strain of B. subtilis showed notable antagonistic effects
on Sclerotinia sclerotium, Botrytis cinerea, Fusarium tricinc-
tum, Rosellinia necatrix, Cytospora sp., Colletotrichum aenigma,
and Colletotrichum fructicola. It demonstrated minimal inhibi-
tion of the plant pathogenic fungus Diaporthe eres and showed
no inhibitory effect on the development of Sclerotium rolfsii
and Botryosphaeria sinensis. These findings are consistent with
earlier research by[19], which reported growth suppression in
various phytopathogenic fungi, with the exception of Sclerotium
rolfsii and Botryosphaeria sinensis. Moreover,[20] noted that the
effect of Bacillus subtilis BS-1 cell-free supernatant (CFS) on
Botryosphaeria dothidea was due to the stimulation of oxidative
damage in the mycelial cells (Figure S1).

The degradation of cellulose by the cellulase enzyme is vital,
as many agricultural byproducts include cellulose, which is a key
element of plant cell walls. This cellulose can be transformed
into glucose, which acts as a key raw ingredient for alcohol man-
ufacturing [21]. The fermentation process has been employed for
a long time, as it increases nutritional value while minimizing
anti-nutritional elements, toxins, and levels of contamination [22].
A successful method for waste management includes the enzy-
matic hydrolysis of lignocellulosic waste, where fermentation
produces reducing sugars that can be utilized for creating desired

metabolites or biofuels [23]. In this research, the peak activity of
cellulase enzyme production and the optimum growth conditions
for the Bacillus subtilis isolate were observed after four days of
incubation (Figure 2).

Phylogenetic Analysis
The 16S rRNA gene of strain S1-10 is 793 base pairs long.

Its sequence shows a 100% match with several species, including
Bacillus subtilis, Bacillus halotolerans, Bacillus amyloliquefa-
ciens, Bacillus safensis, Bacillus velezensis, Bacillus mojaven-
sis, and Bacillus stercoris. A phylogenetic tree built using the
Neighbor-Joining method indicates that strain S1-10 is grouped
on the same branch as Bacillus subtilis, Bacillus stercoris, Bacil-
lus tequilensis, and Colidifontibacillus erzurumensis (Figure 3).

Therefore, the gyrB gene, which encodes the B subunit of
DNA gyrase, was selected as an alternative marker for phylo-
genetic analysis and species identification within the Bacillus
group. The NCBI BLAST results showed that the isolate shares
100% identity with Bacillus subtilis. A phylogenetic tree con-
structed using the Neighbor-Joining method again places strain
S1-10 alongside Bacillus subtilis strains. Similar clustering pat-
terns were also observed in trees generated using the Maximum
Likelihood and Maximum Parsimony methods (Figure 4).

Conclusions
The indigenous Bacillus strain found in the marine sediment

of Myanmar was classified as Bacillus subtilis through gene
analysis of 16S rRNA and gyrB. The isolated strain exhibited an-
tagonistic effects on the mycelial growth of the plant pathogenic
fungi Sclerotinia sclerotiorum, Botrytis cinerea, Fusarium tricinc-
tum, Rosellinia necatrix, Cytospora sp., Colletotrichum aenigma,
and Colletotrichum fructicola. It demonstrated weak inhibi-
tion against the plant pathogenic fungus Diaporthe eres and
showed no inhibition on the growth of Sclerotium rolfsii and
Botryosphaeria sinensis. The Bacillus subtilis S1-10 isolate may
be used as an effective strain for composting agricultural waste
due to its cellulolytic and antagonistic properties.

Supplementary
Figure S1: Screening the antifungal activity of isolate (S1-

10) by showing fungal growth inhibition compared with the
control and treatment plates. Red box indicates the isolate (S1-
10).

Reference
1. Yao H, Liu S, Liu T, Ren D, Yang Q, Zhou Z, et al. Screening

of marine sediment-derived microorganisms and their bioac-
tive metabolites: a review. World Journal of Microbiology
and Biotechnology. 2023;39(7):172.

2. Voser TM, Campbell MD, Carroll AR. How different are ma-
rine microbial natural products compared to their terrestrial
counterparts? Natural Product Reports. 2022;39(1):7-19.

3. Sayem SA, Manzo E, Ciavatta L, Tramice A, Cordone A,
Zanfardino A, et al. Anti-biofilm activity of an exopolysac-

Highlights in BioScience Page 4 of 5 July 2025|Volume 8

http://bioscience.highlightsin.org/


Yu SS et al., 2025 Molecular identification and screening the antagonistic and cellulolytic activities

charide from a sponge-associated strain of Bacillus licheni-
formis. Microbial cell factories. 2011;10:1-12.

4. Elshaghabee FM, Rokana N, Gulhane RD, Sharma C, Pan-
war H. Bacillus as potential probiotics: status, concerns, and
future perspectives. Frontiers in microbiology. 2017;8:1490.

5. Damalas CA, Eleftherohorinos IG. Pesticide exposure, safety
issues, and risk assessment indicators. International journal
of environmental research and public health. 2011;8(5):1402-
19.

6. Islam MT. Potentials for biological control of plant diseases
by Lysobacter spp., with special reference to strain SB-K88.
Bacteria in agrobiology: plant growth responses. 2011:335-
63.

7. Jin X, Song J, Ma J, Liu GQ. Thermostable β-xylosidase
from Aspergillus fumigatus: Purification, characterization
and potential application in lignocellulose bioethanol pro-
duction. Renewable Energy. 2020;155:1425-31.

8. Alokika V, Kumar V, Singh B. Biochemical characteristics
of a novel ethanol-tolerant xylanase from Bacillus subtilis
subsp. subtilis JJBS250 and its applicability in saccharifi-
cation of rice straw. Biomass Conversion and Biorefinery.
2021:1-13.

9. Deka D, Bhargavi P, Sharma A, Goyal D, Jawed M, Goyal
A. Enhancement of cellulase activity from a new strain
of Bacillus subtilis by medium optimization and analy-
sis with various cellulosic substrates. Enzyme research.
2011;2011(1):151656.

10. Errington J, Aart LTvd. Microbe Profile: Bacillus subtilis:
model organism for cellular development, and industrial
workhorse. Microbiology. 2020;166(5):425-7.

11. Duan M, Zhang Y, Zhou B, Qin Z, Wu J, Wang Q, et al.
Effects of Bacillus subtilis on carbon components and mi-
crobial functional metabolism during cow manure–straw
composting. Bioresource Technology. 2020;303:122868.

12. Ntozonke N, Okaiyeto K, Okoli AS, Olaniran AO, Nwodo
UU, Okoh AI. A marine bacterium, Bacillus sp. isolated from
the sediment samples of Algoa Bay in South Africa Produces
a Polysaccharide-Bioflocculant. International journal of en-
vironmental research and public health. 2017;14(10):1149.

13. Aquino-Martínez JG, Vázquez-García LM, Reyes-Reyes
BG. Biocontrol in vitro e in vivo de Fusarium oxysporum
Schlecht. f. sp. dianthi (Prill. y Delacr.) Snyder y Hans. Con
hongos antagonistas nativos de la zona florícola de Villa
Guerrero, Estado de México. Revista mexicana de fitopa-
tología. 2008;26(2):127-37.

14. Hendricks CW, Doyle JD, Hugley B. A new solid medium
for enumerating cellulose-utilizing bacteria in soil. Applied
and environmental microbiology. 1995;61(5):2016-9.

15. Weisburg WG, Barns SM, Pelletier DA, Lane DJ. 16S ribo-
somal DNA amplification for phylogenetic study. Journal of
bacteriology. 1991;173(2):697-703.

16. Munir S, Li Y, He P, He P, He P, Cui W, et al. Bacillus
subtilis L1-21 possible assessment of inhibitory mechanism
against phytopathogens and colonization in different plant
hosts. Pak J Agric Sci. 2018;55(4):996-1002.

17. Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Hig-
gins DG. The CLUSTAL_X windows interface: flexible
strategies for multiple sequence alignment aided by quality
analysis tools. Nucleic acids research. 1997;25(24):4876-82.

18. Kumar S, Stecher G, Tamura K. MEGA7: Molecular Evolu-
tionary Genetics Analysis Version 7.0 for Bigger Datasets.
Molecular Biology and Evolution. 2016;33(7):1870-4. Used
to assess evolutionary distances via Kimuras two-parameter
model and bootstrap analysis (1,000 replicates).

19. Nalisha I, Muskhazli M, Nor Farizan T. Production of bioac-
tive compounds by Bacillus subtilis against Sclerotium rolf-
sii. Malaysian Journal of Microbiology. 2006;2(2):19-23.

20. Fan Y, Liu K, Lu R, Gao J, Song W, Zhu H, et al. Cell-free
supernatant of Bacillus subtilis reduces kiwifruit rot caused
by Botryosphaeria dothidea through inducing oxidative stress
in the pathogen. Journal of Fungi. 2023;9(1):127.

21. Qu Y, Zhu M, Liu K, Bao X, Lin J. Studies on cellulosic
ethanol production for sustainable supply of liquid fuel in
China. Journal of Biotechnology. 2006;1(11):1235-40.

22. Evans E, Musa A, Abubakar Y, Mainuna B. Nigerian indige-
nous fermented foods: processes and prospects. In: Myco-
toxin and food safety in developing countries. IntechOpen;
2013. .

23. Singhania RR, Sukumaran RK, Pillai A, Prema P, Szakacs
G, Pandey A. Solid state fermentation of lignocellulosic
substrates for cellulase production by Trichoderma reesei
NRRL 11460. Indian Journal of Biotechnology. 2006;5:332-
6.

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	Abstract
	Introduction
	Materials and Methods
	Sample collection and strain isolation
	Screening of antagonistic activity and cellulolytic activity
	Analysis of Morphology and Physiology
	Phylogenetic Analysis

	Results and Discussions
	Analysis of Morphology and Physiology
	Phylogenetic Analysis

	Conclusions
	Supplementary

