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Research Article
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Macrofungi Community in Rubber Plantations and a Forest of Edo State, Nigeria |
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O.O. Osemwegie,
J.A. Okhuoya,
A.O. Oghenekaro
and
G.A. Evueh
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ABSTRACT
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Permanent plots in rubber plantations and a lowland forest, each measuring 25x25 m, were randomly laid out using coloured ribbons and studied twice a month for macrofungi for a period of 14 months. A total of 435 fruit bodies belonging to 93 different species of macrofungi were encountered, 70% of which were identified. Identified taxa were distributed into 4 Classes, 9 Orders and 28 Families with the class Hymenomycetes and family Tricholomataceae as the best represented taxa. Agaric (52%) and polypores mushrooms (31%) were also recorded as the best represented life-forms while wood-based substrates recorded 70% of the total mushroom taxa encountered during the study. The species richness and diversity estimate of 100 randomization accumulation sample order of mushroom abundance data from each of the sampled plots showed that the forest (Plot E) had the best species richness and diversity index values compared to plot A, B, C and D.
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INTRODUCTION
Defining the population of fungi globally has in recent times remained a challenge
to mycologists all over the world (Wood, 1992). Many
mycologists agreed that there are more fungi in the world than reported, especially
in the tropics and sub-saharan Africa and this has resulted in the inconsistency
associated with global fungi estimate reported by Hammond
(1992), Hawksworth (1993) and Crous
et al. (2006). Scanty information abounds on the diversity of African
macrofungi (Mueller et al., 2007; Osemwegie
et al., 2006). In Nigeria, mushrooms are often overlooked in many
biodiversity studies compared to plants and animals (Idu
et al., 2007). Mushrooms in Nigeria are poorly collected, sparingly
studied and relatively underutilized (Osemwegie and Okhuoya,
2009; Labarere and Menini, 2000). Higher plants are
preferred by most Nigerian as sources of food amidst sporadic reports on the
use of some mushrooms as food, food supplements and in folk medicine practices,
especially by the rural populace (Akpaja et al.,
2005; Okhuoya and Akpaja, 2005; Osemwegie
et al., 2006).
In some developed countries of the world, macrofungi are exploited for economic
gains in the areas of food security and foreign exchange earnings via large
scale mushroom cultivation, mushroom export and pharmacopoeia. They are also
used to improve silviculture, agroforestry and agriculture and industries such
as brewing, beverage, enzymes, dye, paper mill, organic acids, hormones and
animal feeds industries (Arora, 1989; Chang
and Miles, 1991; Wainwright, 1992; Wasser,
2007). Mushrooms are also applied in waste management and remediation of
contaminated arable lands and waters (Wasser, 2007).
Several researchers both in Nigeria and abroad have reported that many macrofungi
are potential biological control agents of insects, arthropods and other microorganisms
of bacteria and fungi origin (Roberts and Hajek, 1993;
Boa, 2004; Jonathan and Fasidi, 2005;
Gbolagade et al., 2007).
In Nigeria, mushroom researches have focused more on low-cost cultivation of
many indigenous edible mushrooms, their nutriceutics and ethnomycology rather
than their diversity, taxonomy, biogeography and ecology (Rammeloo
and Walleyn, 1993; Osemwegie and Okhuoya, 2009).
Lodge et al. (1995) remarked that the knowledge
of mushroom composition and ecology are central to efforts establishing proactive
conservation strategies and identifying areas in urgent need of conservation
as well as species in short- and long-term danger of extinction. Studies on
the diversity of wild macrofungi indigenous to Nigeria are regional and biased
to agroecosystems (Osemwegie and Okhuoya, 2009).
This study aimed at identifying the diverse mushroom taxa associated with rubber agroforests and comparing their mushroom community with that of a secondary uncultivated forest within the same ecological zone. MATERIALS AND METHODS Study area: The study area, Rubber Research Institute of Nigeria (RRIN) Iyanomo is located in Ikpoba-Okha local Government Area of Edo State, approximately about 29 km from Benin City (Fig. 1). The geographical, ecological and edaphic characteristics are as enumerated in Table 1.
Sampled forest and rubber plots: Permanent plots, each measuring 25x25
m were laid out from randomly selected rubber plantations and a lowland secondary
rainforest all connected by a common road. The plots were each approximately
5 m away from the edge of the road. Plot C and D were old (50-55 years old),
no longer being tapped for latex and characterized by thick undergrowth and
broad canopy cover while Plots A and B were younger (38-43 years old) populated
by rubber trees being tapped for latex and weeded once every year. Plot E was
a secondary forest with thick undergrowth and rich tree diversity. The plots
were each surveyed twice a month for mushrooms for a period of 14 month which
ran from 2006 through to 2007 across season gradients. Observed mushrooms were
collected and preserved according to Lodge et al.
(2004). Vouchered mushrooms were kept in the Mushroom Biology Unit of the
Department of Plant Biology and Biotechnology, University of Benin, Benin City,
Edo State for further molecular verification. Encountered macrofungi were photographed
in situ and features such as phenology, smell, habit, colour, nature
of substrate and associations recorded before transportation to the laboratory
for identification.
Table 1: | Geographic
ecological and edaphic characteristics of the study area |
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Identification was based on macroscopic features and nomenclature was carried
out using a variety of field monograph of coloured mushrooms and books (Largent
and Their, 1984; Largent, 1986; Arora,
1991; Mueller et al., 2004; Lincoff,
2005). The number of fruit bodies per sampled data was later subjected to
analysis using Estimates statistical package according to Colwell
(2005) and Chao et al. (2005).
RESULTS AND DISCUSSION A total of 93 different species of macrofungi represented by 435 fruit bodies (abundance value) were encountered during the 14 months period of study from which 70% were already identified and named (Table 2). The identified species were distributed into 28 Families, 9 Orders, 4 Class and 2 Phyla (divisions). The most represented mushroom life-forms encountered in the study area was the fleshy fungi (gilled or agaric mushrooms) and polypores comprising of 52 and 31% species, respectively. The earth-stars, puffballs, tubers and cup fungi were the least represented (Table 3). The family Tricholomataceae and members of the class Hymenomycetes were the best represented taxa (Table 3). The study showed that wood and litter-based substrates supported the growth of 70 and 23% of mushrooms taxa observed during the study, respectively while the soil-based substrate recorded the least (Fig. 1). Mushrooms such as Chlorophyllum sp., Coprinopsis atramentarius (Bull.) Redhead, Vilgalys and Monclavo, Hygrocybe sp. and Pleurotus tuberregium (Fr.) Sing were observed to colonize more than one type of substrate or exhibited flexible substrate propensity (Table 2). | Fig. 1: | Substrate
types and quantitative representation of mushrooms that inhabit the |
Table 2: |
List of macrofungi observed per sampled plot and their
substrate propensity |
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Plot A and B = 38-43 years old; Plot C and D = 50-55 years
old. ****Class, ***Order, **Group, *Family, +: Present, -: Absent, AYR:
All year round, BW: Burried wood, CW: Coarse wood, DL: Decomposing litters,
DW: Dead decaying wood (tree stump and fallen logs), S: Soil, T: Living
tree, TB: Tree branch |
Rubber tree-dominated secondary forest (Plot E) recorded the highest number
of mushroom species (41) and abundance while Plot D which is one of the older
rubber plots surveyed recorded 36 different types of macrofungi (Table
4, 5). The highest number of unshared species (16) was
also recorded for Plot E and the least (4) for Plot B. Plots A and B were observed
to be the most similar recording similarity index values (Chao shared estimate,
Jaccard, Sorensen and Morisita-Horn classic) closer to 1 compared to similarity
indices recorded for other sampled plots (Table 6).
Table 3: | Distribution
and amount of mushrooms across life-forms and taxonomic hierarchy |
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Table 4: | Number
of mushrooms encountered per sampled plots |
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Plot E also recorded the highest biodiversity indices and the least number
of shared mushroom species amounting to 19% of the total mushroom taxa. It was
also most varied in terms of mushroom composition compared to Plots A, B and
C (Table 5).
Wood-substrate colonizers like Auricularia auricular Judae (Bull.) Pat.,
Coprinopsis acuminatus (Romagn.) Redhead, Vilgalys and Monclavo,
Cyathus striatus (Huds.) Willd., Daldinia concentrica (Bolt. ex Fr.)
Ces., Nothopanus sp., Pleurotus squarrosulus (Fr.) Kum., Volvariella
volvaceae (Bull.) Singer and Schzophyllum commune Fr were observed
in all the plots surveyed. They were also observed all the year round. Auricularia
auricular, S. commune, P.tuberregium, Agaricus arvensis
Schaeff., P. squarrosulus and Pluteus cervinus (Schaeff. ex Fr.)
Kum were some of the indigenous edible mushrooms recorded.
A survey of rubber agroforests and a secondary forest in Rubber Research Institute
of Nigeria recorded 93 different mushrooms which amounted to a total of 435
fruit bodies (abundance value) belonging to 28 families within the 14 months
period of study. Straatsma and Krisai-Greilhuber (2003)
constitute an average of 4.9 fruit bodies per species per month over a total
study area of 3125 m2. The number of mushroom taxa recorded from
the study contrast with similar work done by Shigeki et
al. (1994) in young forests and evergreen broad-leaved forests, Straatsma
et al. (2001) in Swiss forests and Osemwegie
and Okhuoya (2009) in oil palm agroforests of Edo State.
Table 5: | Computation
of biodiversity indices±SD per sampled plot using 100 randomized
sample order |
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Mao
Tau: No. of observed species |
Table 6: | Comparing
the sampled plots mushroom composition using a similarity index
programme in estimates |
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Values
closer to 1 are more similar in species composition |
The differences in the amount and assemblage of recorded mushroom taxa may
be due to variations in sample frequency and time, land area covered during
surveys, the nature of woodland vegetation (homo-or heteroculture; riparian
or lowland or savannah etc.) studied and geographical location. This apparently
proves that many mushrooms such as those that produce hypogeous and ephemeral
fruit bodies may have been missed by the study and remains to be discovered
with further sampling (Lynch and Thorn, 2006). Reports
such as those of Nicholson (2000) and Osemwegie
et al. (2006) on mushroom diversity in Nigeria portrayed agroecosystems
as impoverished in mushroom diversity and overlooked as viable site for mushroom
studies. Flynn et al. (2009) emphasized their
ultimate ecological function of provisioning ecosystem services such as biogeochemical
cycles, soil binding, decomposition, soil conditioning and regulation of ecosystem
balance which supports the well-being of other biotas. These services amongst
which are mycorrhization may be harnessed for the development and management
of agroecosystems in Nigeria as reported in some African countries (Marx
et al., 1993). Bolger (2001) and Loreau
et al. (2001) observed a positive relationship between rich biodiversity
and ecosystem functions while also recognizing the functional value of species
in forming the ideological framework for improving the performance and productivity
and decreasing the input of energy, chemical fertilizers and pesticides in agricultural
systems. Further studies are required to fully understand the relevance of mushroom
community or assemblage to the overall health of rubber plantations. This present
study has provided the insight and bases by the sheer diversity of macrofungi
recorded in the rubber plantations.
A break down of the result obtained from the study showed that the rubber agroforest
plots (A, B, C and D) sampled recorded mushroom diversity whose composition
varied with their relative age and level of human disturbance. Plot E however
recorded the highest number of fruit bodies amounting to 40 species of mushrooms
and this was reflected in the values of Mau Tao (93±4.6), Alpha (36.7±2.9),
Shannon (4.2±0.0) and Simpson (57.3±0.0) diversity indices computed
while Plot A recorded the least. The tree heterogeneity and according to Tsui
et al. (1998) the low level of human disturbances associated with
plot E may be responsible for the higher value of species diversity indices
recorded. It is however, important to note that further study is required to
adequately understand the qualitative and quantitative impact of human dynamics
in structuring the mushroom composition of a vegetation. This result therefore
supports the line of thought that human activities do impinge on mushroom diversity
and stands in agreement with existing scientific mushroom biodiversity findings
that relate mushroom diversity to tree diversity (Sala et
al., 2000; Jumpponen et al., 2004). Lodge
et al. (1995), Laitung and Chauvet (2005)
and Mueller et al. (2007) observed a parallel
relationship between tree diversities and mushroom richness while Hawksworth
(2001) recognized the use of trees in the estimation of global mushroom
diversity. Conversely, the varying degrees of rubber latex-tapping activity
that characterized the other plots studied and their respective tree homogeneity
may be responsible for the relatively low incidence of mushroom taxa and diversity.
A similarity index analysis of the various sampled plots according to Chao
et al. (2005, 2006) showed that Plots A and
B were the most similar in terms of species composition, sharing 74% (23 species)
of their recorded mushroom taxa while Plot E only shared 19% (9 species) of
its total taxa. The reason for this is not yet fully understood but it might
be connected to variations in the overall nature (diversity of trees, other
biota, climate, landscape, productivity or turnover) and intrinsic configuration
(tree girth and distance from one another, physiognomy or vegetation layers,
canopy spread, gaps, fragmentation) of each of the sampled plots.
The study also recorded 52% agaric and 31% polypore fungi, respectively while
other mushroom life-forms such as earth stars, puffballs and tubers were scanty.
Agaric and polypore fungi are mostly saprotrophic and capable of biodegrading
many recalcitrant organic-based substrates (Lynch and Thorn,
2006). This inherent attribute coupled with their intrinsic enzyme spectrum
and dynamics which according to Schmit (2005) consequently
broadens accessible substrate-based options, may be the reason for their high
representation. The high level of accessible energy resources (cellulose, hemicellulose
and lignin) fixed in diverse wood-based substrates in the various sampled plots
may have also accounted for the 70% wood-inhabiting mushrooms recorded during
the study. Consequently, the volume of wood and its distribution within the
sampled plots may have accounted for the high incidence of unshared species
(16) observed in Plot E as compared with Plot B which recorded 4 unshared species.
This result supports research findings illuminating wood-based substrate as
a major determinant of mushroom diversity in woodland vegetations (forest and
agroforests) in both temperate and tropical regions. Although, little is known
about variations in the pattern of wood resource utilization by different species
of macrofungi, these factors in addition to the nature of substrate chemistry
and microenvironment may have impacted more on the distribution of mushrooms
than species richness in both agroforest and forest systems. The high (31%)
incidence of fleshy (agaric) mushroom life-forms recorded during the study correlates
positively with increased representation of members of the family Tricholomataceae
most of which were litter mushrooms. Chlorophyllum species, C. atramentarius,
P. tuberregium and Hygrocybe species were observed to fruit on
both soil and wood substrates. This wider substrate colonization propensity
observed amongst some of the macrofungi may have also played a fundamental role
in the higher incidences of polypore and agaric mushroom life-forms recorded
during the study.
Auricularia auricular, C. acuminate, C. striatus, Daldinia
concentrica (Bolt.) Ces. and DeNot., Nothopanus sp., P. squarrosulus
and S. commune were observed throughout the study area, overlapping boundaries
of sampled plots. This characteristic may be attributed to the availability
of widely distributed rich nutrient-based substrates (wood debris). In addition,
this observation is in concert with of Ozinga et al.
(2009) that the dynamism rather than the mechanism of their spore dispersal
in space (long-distance travel) and/or time (dormancy or rest period) can determine
the biogeographic spread of mushroom taxa.
The relatively large number of unidentified species incurred by the study was due to dearth of previous studies, expert mushroom taxonomists, revised mushroom diversity data especially on Nigerias mycoflora and foreign technical supports (Osemwegie and Okhuoya, 2009). Researchers and mushroom scientists are challenged to inventory the nations mushroom heritage and explore the grey areas of mushroom taxonomy, ecology and biotechnology studies in Nigeria without prejudice to any vegetation.
Agroecosystems were hitherto perceived as poor in mushroom diversity are by
this study recognized as good alternative and sustainable sources of mushroom
resources with unprecedented utilitarian values. The study recorded popular
edible and medicinal mushrooms such as A. arvensis, A. auricular,
Macrolepiota sp., P. tuberregium, P. squarrosulus, Pluteus
cervinus, S. commune and V. volvacea in rubber agroforests.
It also lends credence to claims that rich tree diversity facilitated luxuriant
growth of mushroom-forming fungi which ab initio provide ecosystem services
and ecological energy-balance. Furthermore, the study recognized the superiority
of forests over agroforests in terms of mushroom assemblage, diversity, abundance
and species richness. The contribution of mushrooms to woodland systems was
conceptualized by Lawton (1994) and Giller
and ODonovan (2002), who reiterated the need to conserve and preserve
national indigenous mushroom flora as a tool in the whole complex process of
forest and agroforest management.
ACKNOWLEDGMENTS We thank the managements of the University of Benin for providing part of the grant for the research and the Rubber Research Institute of Nigeria (RRIN) for approving the use of her rubber plantations and forest for the research. The contributions of Dr. O.S Isikhuemhen of North Carolina State University and Dr. Cathie Aime of Louisiana State University in the identification of some of the mushroom taxa cannot also go on appreciated.
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