Mor. J. Agri. Sci. 2 (1): 9-13, March 2021
9
Inocybaceae and affiliated taxa from West Africa
Hyppolite L. AIGNON1*, Arooj NASEER2, Azize A. BOUKARY1, Nourou S. YOROU1
Abstract
1
Research Unit Tropical Mycology
and Plant-Soil Fungi Interactions,
Faculty of Agronomy, University of
Parakou, Benin
2
Department of Botany, University
of the Punjab, Quaid-e-Azam
Campus, Lahore, Pakistan
*
Corresponding author
hyppoliteaignon@yahoo.com
Received 26/01/2021
Accepted 24/02/2021
Inocybaceae and its affiliated taxa recorded in West Africa were examined by bibliographic survey through available publications coupled with field data collections in
Benin, Burkina Faso, Côte d’Ivoire, Guinea, Mali and Togo. Twenty-one Inocybaceae
and affiliated taxa have been reported in the region. All taxonomic names have been
checked in Index Fungorum and the distribution of Inocybe beninensis, I. flavipes,
I. fuscobrunnea, I. pallidiangulata, Inosperma africanum, I. bulbomarginatum, I.
flavobrunneum, Mallocybe africana, Pseudosperma afrofibrosum, P. beninense and P.
fragilipes in West Africa were reported. In addition, results of Basic Local Alignment
Search Tool search including some undescribed collections from Zambia, such as
PC96082, PC 96204 with PC:0088767 and L4512 Inoc Zam05 that showed similarity
respectively with the collections of Inosperma bulbomarginatum, Mallocybe africana
and Inocybe flavipes from West Africa with >97% similarity.
Keywords: Checklist, Ectomycorrhizal fungi, Inocybe, Distribution, Pseudosperma,
West Africa
INTRODUCTION
Inocybaceae Julich, is monophyletic family with a
worldwide distribution. It encompasses nearly 1,050
species (Matheny et al., 2020) found mainly in forests
(from woodland, gallery or dense forests) and rarely in
grasslands, wetlands or agricultural lands (Ov, 2015).
The species of Inocybaceae are ectomycorrhizal and associate with 23 families of vascular plants (Matheny et
al., 2020). Recently, Matheny et al. (2020) suggested a revised phylogeny for the family Inocybaceae. Today, Inocybaceae contains seven genera, Auritella Matheny &
Bougher, Inocybe (Fr.) Fr., Inosperma (Kühner) Matheny
& Esteve-Rav., Mallocybe (Kuyper) Matheny, Vizzini &
Esteve-Rav, Nothocybe Matheny & K.P.D. Latha, Pseudosperma Matheny & Esteve-Rav and Tubariomyces
Esteve-Rav. & Matheny (Matheny et al., 2020).
However, due to similarity and lack of molecular analyses, a large number of Inocybaceae taxa were classified
in Cortinariaceae Singer (Horak, 1978; Singer, 1986,
1975) and probably all that were recognized then, which
may include taxa that are not recognized now such as
Astrosporina Schröt. In the literature, most of the premolecular classifications merged not only Inocybaceae
with affiliated taxa, Crepidotaceae Singer and Tubariaceae Vizzini. The first taxonomic evaluation within Inocybaceae was based on morphological and microscopic
characters for the distinction of genera earlier (Kuyper,
1986). Beforehand, Jülich (1982) placed Inocybe (Fr.) Fr.
in Inocybaceae together with the genus Astrosporina
which is now known to be polyphyletic and nested
within Inocybe (Matheny, 2005; Matheny et al., 2002).
Except some scarce publications (Aïgnon et al., 2021a;
b; Buyck and Eyssartier, 1999; Matheny and Watling,
2004; Piepenbring et al., 2020) taxonomic documenta©
Moroccan Journal of Agricultural Sciences • e-ISSN: 2550-553X
tion of the family Inocybaceae from tropical Africa is
scant. But, recent mycological sampling in West Africa
reported eleven new species to science that some of
which like Inosperma africanum, I. bulbomarginatum,
I. flavobrunneum and Mallocybe africana have been
published (Aïgnon et al., 2021a;b). Though taxonomic works are still progressing, it is of paramount
importance to provide a state-of-the-art related to the
diversity known in West Africa, in order to establish a
reference guide to detect new species.
In this paper, we provide a checklist of known species of
Inocybaceae and affiliated taxa from West Africa based
on the literature, observation and field data collection
between 2013 and 2018.
MATERIALS AND METHODS
Study area
West Africa is defined here to include 15 countries (Benin, Burkina Faso, Côte d’Ivoire, Cape Verde, Gambia,
Ghana, Guinea, Guinea-Bissau, Liberia, Mali, Niger,
Nigeria, Senegal, Sierra Leone and Togo). The recently
undertaken sampling trips and collection sites in Benin,
Burkina Faso, Côte d’Ivoire, Guinea, Mali and Togo are
indicated in Figure 1.
Data compilation
The data were assembled from literature related to Inocybaceae from West Africa. All scientific names have
been cross-checked against Index Fungorum (IF) and
synonymous names have been separated.
Data from literature were coupled with field observations. We made field surveys from 2013 to 2018.
Specimens were collected in Benin (Wari Maro forest
www.techagro.org
Aignon et al.: Inocybaceae from West Africa
10
reserve, Ouémé Supérieur forest reserve, Okpara forest
and Toui-Kilibo forest reserve). Additional surveys were
carried out in Burkina Faso (Toussianbandougou gallery
forest, Dan gallery forest and Niangoloko forest reserve),
Mali (Farako forest reserve), Guinea (Kouraouletediene
forest reserve, Levari forest, Baroforest reserve, Moussaya forest reserve, Haut Niger National Park, Telaya
forest, Tindo forest and Côte d’Ivoire (Kekrekouakoukro
forest and Kouadianikro forest). The sampling sites are
presented in Figure 1.
DNA extraction, PCR and sequencing
On the recent collection of Inocybaceae from Benin,
Burkina Faso, Côte d’Ivoire, Guinea, Mali and Togo,
DNA was extracted from dried badisiomata using
a QIAGEN® plant mini kit and for the Polymerase
chain reaction (PCR) amplification, three nuclear gene
regions such as Internal Transcribed Spacer (ITS),
portions of the large subunit ribosomal DNA gene
(28S) and RNA polymerase II subunit (RPB2), were
amplified using the following primers, ITS1F and ITS4
for the ITS region (Gardes and Bruns, 1993; White et
al., 1990), LR0R, LR7, LR5 and LR3R for 28S (Cubeta et
al., 1991; Rehner and Samuels 1995 Vilgalys and Hester,
1990,) and bRPB2-6F, bRPB2-7.1R for the most variable
region of RPB2 (Matheny 2005). PCR products were
cleaned and sequenced at Macrogen Inc. (Macrogen
Europe B.V., Amsterdam, Netherlands) using the same
primers as those used for PCR. All sequences of the
new species are deposited on GenBank (Aïgnon et al.,
2021a;b) and the new sequences were subjected to a
BLAST search (97% similarity) and relevant related
sequences retrieved from GenBank.
RESULTS
Inocybaceae and affiliated/Allied taxa in west
Africa
Twenty-one Inocybaceae and affiliated taxa have been
reported from West Africa. Table 1 summarizes the different species of Inocybaceae as well as the affiliated taxa
of West Africa.
Existing data show that Inocybaceae and affiliated taxa
have been recorded in four countries from West Africa i.e.,
Ghana, Nigeria, Senegal and Sierra Leone and collected
in forests dominated by Adenia lobata Eng, Afzelia africana Sm. ex Pers., Afzelia bella Harms, Afzelia bracteata T.
Vogel ex Benth, Anthonotha crassifolia Baill. and Uapaca
chevalieri Beille. But, recent sampling in Benin, Burkina
Faso, Côte d’Ivoire, Guinea, Mali and Togo has shown that
the species of Inocybaceae are found in the vegetations
dominated by Isoberlinia doka Craib & Stapf, I. tomentosa
(Harms) Craib & Stapf, Uapaca togoensis Pax, U. guineensis
Müll. Arg., Monotes kerstingii Gilg and Berlinia grandiflora
Hutch. & Dalziel.
At the 97% threshold, after BLAST searched at NCBI
(www.blast.ncbi.nlm.nih.org), Inocybe sp. PC96082
from Zambia present high similarity with the collections
Côte d’Ivoire
Figure 1: Map of West Africa showing the studied areas and recent sampling sites
Mor. J. Agri. Sci. 2 (1): 9-13, March 2021
11
of Inosperma bulbomarginatum, same for the collections
PC 96204 and PC:0088767 from Zambia with the collections of Mallocybe africana from West Africa, as well as
the collection L4512 Inoc Zam05 from Zambia with the
collections of Inocybe flavipes from West Africa which
appear to be the same species.
DISCUSSION
Uptill now, only five species of Inocybaceae, Inocybe
ghanaensis (Pegler 1969; Piepenbring et al., 2020),
Mallocybe africana (Aïgnon et al., 2021b), Inosperma
africanum, I. bulbomarginatum, and I. flavobrunneum
(Aïgnon et al., 2021a) are reported and well described
from West Africa to our knowledge, but many taxa
are still without description and wait to be published
Inocybe beninensis, I. flavipes, I. fuscobrunnea, I. pallidiangulata, Pseudosperma afrofibrosum, P. beninense,
P. fragilipes. However, some name is not validated such
as Inocybe sp. “gbadjii” registered in index fungorum but
it is not consequently described due to lack of diagnosis
and type (Boa, 2004). In addition to these species there
are many undescribed collections described as Inocybe
spp. from Burkina-Faso, Guinea, Nigeria and Senegal
(Redhead, 1968; Thoen and Ducousso, 1989; Bâ et al.,
2012; Piepenbring et al., 2020). Recent findings (Aïgnon
et al., 2021a;b) has increased the diversity of Inocybaceae
in West Africa to more than eighteen species (Table 1).
However, the Central and East Africa regions contain
Table 1: Checklist of Inocybaceae and affiliated taxa from West Africa
Family
Species
Crepidotus
applanatus
Authority
(Pers.) P. Kumm.
Host Plant
Distribution
Sierra Leone
Field observation:
Collected from dead
trunks of Adenia lobata
(Beeli, 1938)
-
Sierra Leone
-
(Holden, 1970)
Nigeria
Dead decaying wood
(Tree stump and fallen
logs), Decay trees, Dead
decaying wood (Tree
stump and fallen logs)
(Beeli, 1938)
(Osemwegie and
Okhuoya, 2011)
(Osemwegie et al., 2006)
(Osemwegie et al., 2010)
Benin
Field observations: on
soil
In press
Crepidotus mollis (Schaeff.) Staude -
Inocybe
beninensis
Aïgnon, Yorou & Isoberlinia doka
Ryberg
and I. tomentosa
I. flavipes
Aïgnon, Yorou & Isoberlinia doka and Benin, Togo
Ryberg
I. tomentosa
Field observations: on
soil
In press
I. fuscobrunnea
Aïgnon, Yorou &
Burkina Faso,
Berlinia grandiflora Côte d’Ivoire
Ryberg
In press
I. ghanaensis
Pegler
Field observations: on
soil
Field observations: on
bare soil
Field observations: on
soil
-
(Boa, 2004)
-
Ghana
Burkina Faso,
I. pallidiangulata Aïgnon, Yorou &
Berlinia grandiflora Côte d’Ivoire
Ryberg
Inocybaceae
References
Adenia lobata
Crepidotaceae
Crepidotus caspari Velen.
Observations
Inocybe sp.
-
Benin
Inocybe sp.
Afzelia africana
Senegal
Inocybe sp.
Afzelia bella
Senegal
Inocybe sp.
Anthonotha
crassifolia, Uapaca
chevalieri
Senegal
Inocybe sp.
Afzelia africana
Senegal
Inocybe sp.
Afzelia africana
Senegal
Burkina
Aïgnon, Yorou & Isoberlinia doka and Benin,
Faso,
Guinea,
Ryberg
Berlina grandiflora. Côte d’Ivoire, Togo
Aïgnon, Yorou & Isoberlinia doka, I. Benin,
I.
bulbomarginatum Ryberg
tomentosa
Aïgnon, Yorou & Isoberlinia doka and Benin,
I. flavobrunneum Ryberg
I. tomentosa
Togo,
Aïgnon, Yorou & Isoberlinia doka and Benin,
Mallocybe
Burkina Faso,
Ryberg
africana
I. tomentosa
Côte d’Ivoire
Inosperma
africanum
P. afrofibrosum
P. beninense
P. fragilipes
Aïgnon, Yorou &
Isoberlina doka
Ryberg
Aïgnon, Yorou &
Isoberlina doka
Ryberg
Aïgnon, Yorou &
Isoberlina doka
Ryberg
Benin
Benin
Benin
(Pegler, 1969)
(Holden, 1970)
In press
Field observations:
Appeared on lateral
(Bâ et al., 2012)
roots of Afzelia africana
Field observation:
Appeared near of
(Redhead, 1968)
Afzelia bella
Test synthesis
mycorrhizae in vitro or
in semi-axenic condition
Rootlets fixation in
formaldehyde-acetic acid
Rootlets fixation in
formaldehyde-acetic acid
Field observations: on
soil
Field observations: on
soil
Field observations: on
soil
Field observations: on
soil
Field observations: on
soil
Field observations: on
soil
Field observations: on
soil
(Thoen and Ducousso,
1989)
(Thoen and Bâ, 1989)
(Thoen and Bâ, 1989)
(Aïgnon et al., 2021a)
(Aïgnon et al., 2021a)
(Aïgnon et al., 2021a)
(Aïgnon et al., 2021b)
In press
In press
In press
Aignon et al.: Inocybaceae from West Africa
12
high diversity of Inocybaceae. The diversity of Inocybaceae is also high in the Central and East Africa regions
where Auritella encompasses three species such as Auritella hispida Matheny & T.W. Henkel, Auritella spiculosa Matheny & T.W. Henkel and Auritella aureoplumosa
(Watling) Matheny & Bougher from Cameroon, as well
as two undescribed species Inocybe sp. TU112047 and
Inocybe sp. TU112061 from Gabon (Matheny et al., 2017,
2012). In Tubariomyces, there is an undescribed species
Tubariomyces sp. 2 BB6018 (PC), from Zambia (Vizzini
et al., 2013). Inosperma misakaense (Matheny & Watling)
Matheny & Esteve-Rav. is known from Zambia (Aïgnon
et al., 2021; Buyck and Eyssartier, 1999; Gardens, 2017;
Matheny and Watling, 2004). As well as, there are several
undescribed species from Zambia including Inocybe sp.
PC 96042, Inocybe sp. PC 96039, Inocybe sp. PC 96081,
Inocybe sp. PC 96095, Inocybe sp. PC 96204, Inocybe sp.
PC 96111, Inocybe sp. PC 96013, Inocybe sp. PC 96083,
Inocybe sp. BB3233, Inocybe sp. BB6018 and Inocybe sp.
PC 96073 (Matheny et al., 2009).
Ectomycorrhizal symbiosis is proven between some
species of Inocybaceae and the roots of native trees. For
example, Anthonotha crassifolia and Uapaca chevalieri
are partner trees of some unidentified Inocybaceae species in the semi-deciduous forest of Senegal (Thoen and
Ducousso, 1989), likewise for Afzelia bella in the tropical forest of Nigeria (Redhead, 1968) and also Afzelia
africana in wooded areas and gallery forest in Senegal
(Thoen and Bâ, 1989). In addition, these ectomycorrhizal host trees are found in various vegetations such as
Zambesian and Sudanian woodlands, semi-deciduous
forests, Guineo-Congolean dense forests and gallery
forests. The presence of Inocybaceae species in these
vegetations is remarkable, and there is little doubt that
other African species of Afzelia, Anthonatha and Uapaca
are also ectomycorrhizal associated with Inocybe but it is
difficult to confirm without consequent analysis.
Analysis of the recent collections (Aïgnon et al., 2021a,b)
have shown a wide distribution of Inocybaceae species in
vegetations dominated by Uapaca spp., Isoberlinia spp.
and Berlinia grandiflora with large distributed in Benin,
Burkina Faso, Guinea, Senegal and Togo (Thoen and Ducousso, 1989; Moyersoen and Fitter, 1999; Newbery and
Stoll, 2013). The same is true for Monotes kerstingii often
mixed with stands of Isoberlinia spp. (Sanon et al., 1997).
It is then clear that Inocybaceae have not been sufficiently
evaluated in West Africa and so there is need to study the
biology and distribution of these species to determine
specific host trees for conservation interventions.
Acknowledgements
We are highly indebted to National Geographic Society (grant No.
CP 126R-17) for financial assistance for collection of many of the
specimens analyzed in this paper. We are also indebted to the Swedish Research Council for Environment, Agricultural Sciences and
Spatial Planning (grant No. 226-2014-1109) for funding molecular
analysis. Microscopic studies have been possible thanks to the microscope equipment granted to Research Unit Tropical Mycology
and Plant-Soil Fungi Interactions (MyTIPs) by German Academic
Exchanges Service (DAAD grant N° PKZ 300499). We are grateful
to P. Brandon Matheny, Martin Ryberg, Sana Jabeen and Boris A.
Olou for their valuable comments and suggestions to improve the
manuscript.
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