Pulveroboletus Bembae
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''Pulveroboletus bembae'' is a species of
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
in the family
Boletaceae The Boletaceae are a family of mushroom-forming fungi, primarily characterised by small pores on the spore-bearing hymenial surface (at the underside of the mushroom), instead of gills as are found in most agarics. Nearly as widely distributed ...
that was first described in 2009. It is known only from the
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
of northern
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
, a region known for its high level of
species diversity Species diversity is the number of different species that are represented in a given community (a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional species abundan ...
. Like all
bolete {{refimprove, date=July 2020 A bolete is a type of mushroom, or fungal fruiting body. It can be identified thanks to a unique mushroom cap. The cap is clearly different from the stem. On the underside of the cap there is usually a spongy surfa ...
s, ''P. bembae'' has fleshy
fruit bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
that form
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s in tubes perpendicular to the ground on the underside of the
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. Caps typically have a visor, or no brim at all. They are popular in casual and informal se ...
. These yellowish tubes form a surface of pores, each about 1–2 mm in diameter. The brownish caps may reach up to wide, and rest atop pale brown stems up to long. The stems have a woolly, whitish yellow
ring Ring may refer to: * Ring (jewellery), a round band, usually made of metal, worn as ornamental jewelry * To make a sound with a bell, and the sound made by a bell :(hence) to initiate a telephone connection Arts, entertainment and media Film and ...
of tissue that is short-lived, and may be absent in older specimens. The
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s of ''P. bembae'' are spindle- or fuse-shaped, and have rough surfaces—a detail observable when viewed with scanning electron microscopy. The fungus grows in a
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l relationship with ''
Gilbertiodendron dewevrei ''Gilbertiodendron dewevrei'' is a species of tree in the family Fabaceae, native to tropical rain forests in Central Africa. It is often the dominant tree species of the Guineo-Congolian rainforest. The timber is traded as ''limbali'', and is us ...
'', the dominant tree species of the Guineo-Congolian rainforest. Other similar ''Pulveroboletus'' species in the area include '' P. annulus'' and '' P. croceus'', which may be differentiated from ''P. bembae'' by a combination of macro- and microscopic characteristics.


Discovery and classification

The specimens of ''Pulveroboletus bembae'' upon which the species description is based were collected in April, 2008 from three locations in
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
: in Ogooue-Ivindo Province at the Ipassa-Makokou Research Station; in the
Minkébé National Park Minkébé National Park is a national park in the extreme northeast of Gabon. It covers an area of 7,570 km2. The WWF recognized it as an area needing protection as early as 1989 and has been actively working towards protecting the forest sin ...
near
Minvoul Minvoul is a town in Woleu-Ntem Province, Gabon. Location It is near the border with Cameroon and is near a tributary of the Ntem River The Campo (in Spanish: ''Río Campo'') or Ntem River is a border river in Cameroon, mainland Equatorial G ...
, and in Bitouga, both locations in the northerly province of
Woleu-Ntem Woleu-Ntem is the northernmost of Gabon's nine provinces. It covers an area of 38,465 km and named after Woleu and Ntem rivers that cross it. The provincial capital is Oyem, which had a total of 60,685 inhabitants in 2013. As Woleu-Ntem i ...
. Until the report of this species and the related '' Pulveroboletus luteocarneus'', 12 species of ''
Pulveroboletus ''Pulveroboletus'' is a genus of fungi in the family Boletaceae. The genus has a cosmopolitan distribution and contains 25 species. Taxonomy The genus was first described by American mycologist William Alphonso Murrill in 1909. He defined specie ...
'' had been reported in
tropical Africa Although tropical Africa is mostly familiar to the West for its rainforests, this biogeographic realm of Africa is far more diverse. While the tropics are thought of as regions with hot moist climates, which are caused by latitude and the tropi ...
. According to Degreef & De Kesel, who described the species in a 2009 publication, ''P. bembae'' belongs to the
section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sign ...
''Pulveroboletus'' of the genus ''Pulveroboletus''. This section, defined by Singer in 1947, is characterized by the presence of a pulverulent-arachnoid veil (covered with fine, powdery wax granules and cobwebby) and fruit bodies that are sulphur-yellow, greenish, or yellowish-brown in color. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
is derived from the word ''bemba'', a name used by the Baka people for the tree ''
Gilbertiodendron dewevrei ''Gilbertiodendron dewevrei'' is a species of tree in the family Fabaceae, native to tropical rain forests in Central Africa. It is often the dominant tree species of the Guineo-Congolian rainforest. The timber is traded as ''limbali'', and is us ...
'' that is associated with the fungus.


Description

The
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. Caps typically have a visor, or no brim at all. They are popular in casual and informal se ...
is initially convex—sometimes with a small rounded elevation in the center—and flattens out in maturity. It reaches in diameter, and the color is almost uniformly rust-brown to reddish brown, although young specimens have a slightly paler margin (edge). The cap surface is dry and dull, but develops a sheen with age. In older specimens, the texture of the margins is described as ''rimulose''—a condition in which a surface is cracked, but the cracks do not intersect one another to form a network and mark out areas. The
cap cuticle The pileipellis is the uppermost layer of hyphae in the pileus of a fungal fruit body. It covers the trama, the fleshy tissue of the fruit body. The pileipellis is more or less synonymous with the cuticle, but the cuticle generally describes th ...
extends slightly over the edge of the cap and curves downward, and is partly covered with remnants of the
universal veil In mycology, a universal veil is a temporary membranous tissue that fully envelops immature fruiting bodies of certain gilled mushrooms. The developing Caesar's mushroom (''Amanita caesarea''), for example, which may resemble a small white sphere ...
. The
flesh Flesh is any aggregation of soft tissues of an organism. Various multicellular organisms have soft tissues that may be called "flesh". In mammals, including humans, ''flesh'' encompasses muscle Skeletal muscles (commonly referred to as mu ...
at the center of the cap is less than about thick, and gradually becomes very thin towards the margin. It is cream-colored to pale yellow with pale reddish-brown to light brown shades under the cuticle and down the
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
. The yellowish tubes on the underside of the cap are slightly swollen on one side, slightly depressed around the area of attachment to the stem. They are fused to the stem, in an
adnate Adnate may refer to: * Adnation, in botany, the fusion of two or more whorls of a flower * Adnate, in mycology, a classification of lamellae (gills) * Conjoined twins Conjoined twins – sometimes popularly referred to as Siamese twins – are ...
attachment; rarely, some tubes will have a decurrent "tooth" (tissue that runs slightly down the length of the stem) that is less than long. The pores formed by the tube ends are angular to round, and are more elongated near the stem. Their diameters are typically less than 1–2 mm in diameter, and are they are the same color as the tubes, or slightly greener. The
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
is by thick, cylindrical, with a narrow base measuring 2–4 mm, and sometimes attached to yellow
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
. It is solid, but as it ages it becomes stuffed (as if filled with cotton) and eventually almost completely hollow. The stem surface is dull, dry, pale brown, and entirely covered with tiny brown to reddish brown squamules (small scales). The flesh of the stem is cream-colored, streaked with pale reddish brown to light brown from the upper third towards the base, while the base is light brown. The
ring Ring may refer to: * Ring (jewellery), a round band, usually made of metal, worn as ornamental jewelry * To make a sound with a bell, and the sound made by a bell :(hence) to initiate a telephone connection Arts, entertainment and media Film and ...
is located on either the stem or the margin of the cap. This woolly, whitish yellow ring of tissue is fragile and short-lived, and has usually weathered away in older specimens. The odor of the mushroom is described as "mildly fungoid to earthy", and the taste "mildly fungoid".


Microscopic characteristics

The color of the
spore print 300px, Making a spore print of the mushroom ''Volvariella volvacea'' shown in composite: (photo lower half) mushroom cap laid on white and dark paper; (photo upper half) cap removed after 24 hours showing pinkish-tan spore print. A 3.5-centimeter ...
is unknown. The
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s are somewhat spindle-shaped, boletoid (long, lean, and fuse-shaped), with a pronounced suprahilar depression (a surface indentation formed where the spore was attached to the spore-bearing cells, the
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly-c ...
), and typically measure 9.3–11.3 by 3.9–4.7 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
. They are weakly pigmented, and their rough surfaces can be seen under scanning electron microscopy. The spores are
inamyloid In mycology a tissue or feature is said to be amyloid if it has a positive amyloid reaction when subjected to a crude chemical test using iodine as an ingredient of either Melzer's reagent or Lugol's solution, producing a blue to blue-black stain ...
, meaning they will not absorb
iodine Iodine is a chemical element with the symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , and boils to a vi ...
stain from
Melzer's reagent Melzer's reagent (also known as Melzer's iodine reagent, Melzer's solution or informally as Melzer's) is a chemical reagent used by mycologists to assist with the identification of fungi, and by phytopathologists for fungi that are plant pathogens ...
. The basidia are 26.9–39.3 by 9.0–12.0 µm, cylindrical to narrowly club-shaped,
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
(translucent), and have four
sterigmata In biology, a sterigma (pl. sterigmata) is a small supporting structure. It commonly refers to an extension of the basidium (the spore-bearing cells) consisting of a basal filamentous part and a slender projection which carries a spore at the ti ...
(extensions that attach the spores). The
pleurocystidia A cystidium (plural cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that ar ...
(
cystidia A cystidium (plural cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that ar ...
on the gill face) are 57.4–92.6 by 9.4–17.4 µm, spindle-shaped, moderately frequent, and extend beyond the surface of the
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some ...
. They have thin, hyaline walls, and are colored the same as the hymenium, without any crystals or encrustations. The
cheilocystidia A cystidium (plural cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that ar ...
(cystidia on the gill edge), which measure 50.6–75.1 by 12.2–16.1 µm, are more abundant than the cheilocystidia, but otherwise share the same characteristics. The cap cuticle is made of a thin physalo- palisadoderm—a type of tissue where the ends of the
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e reach the same length and form a palisade of cells; these short
anticlinal Anticlinal may refer to: *Anticline, in structural geology, an anticline is a fold that is convex up and has its oldest beds at its core. *Anticlinal, in stereochemistry, a torsion angle between 90° to 150°, and –90° to –150°; see Alkane_st ...
hyphae are 20–40 by 5–8 µm, and support one or two inflated, brownish, spherical to spheropedunculate (somewhat spherical with a stem) terminal elements that are 25–45 µm wide, non-amyloid, thin-walled, and do not have any encrustations. The cuticle of the stem is made of smooth parallel hyphae. The squamules on the cap surface have a physalo-palisadodermic arrangement made of short
anticlinal Anticlinal may refer to: *Anticline, in structural geology, an anticline is a fold that is convex up and has its oldest beds at its core. *Anticlinal, in stereochemistry, a torsion angle between 90° to 150°, and –90° to –150°; see Alkane_st ...
hyphae that support elongated inflated elements of 15–30 by 10–15 µm and some scattered basidia. The flesh is made of hyaline, thin-walled hyphae, measuring 10–15 µm wide, and organized in a parallel fashion. These hyphae do not have an associated mediostratum—a central strand of parallel hyphae from which other hyphae diverge sideways.
Clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rece ...
s are absent in the hyphae of ''P. bembae''.


Similar species

Although the fruit bodies of ''P. bembae'' are roughly similar to those in ''
Xerocomus ''Xerocomus'' is a genus of poroid fungi related to ''Boletus''. Many mycologists did not originally recognize the distinction between the two genera and placed ''Xerocomus'' taxa in genus ''Boletus''. However, several molecular phylogenetic stu ...
'', species in this genus do not have the powdery veil characteristic of ''P. bembae''. Two similar species in the same area include '' P. annulus'' and '' P. croceus'', described in 1951 by Belgian mycologist
Paul Heinemann Paul Heinemann (February 16, 1916 – June 18, 1996) was a Belgian botanist and mycologist. Heinemann specialized in African mycology. In his long career, he published 435 names, including 2 families, 6 genera, 346 species, and 40 varieties. His co ...
, based on specimens collected in the Congo. Although the identity of these two species is not fully clarified because of insufficient collections, ''P. bembae'' differs from both in its larger cystidia, its cream-colored flesh with pale reddish-brown to light brown tones under the cap cuticle (compared to white in ''P. annulus'' and ''P. croceus''), its yellow mycelium (white in ''P. annulus'' and ''P. croceus''), and differences in ecology.


Habitat and distribution

The species has been found growing in small groups in the Guineo-Congolian rainforest. This forest is dominated by the single
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
tree species ''
Gilbertiodendron dewevrei ''Gilbertiodendron dewevrei'' is a species of tree in the family Fabaceae, native to tropical rain forests in Central Africa. It is often the dominant tree species of the Guineo-Congolian rainforest. The timber is traded as ''limbali'', and is us ...
''. Not only does this tree provide food in the form of edible seeds for a wide variety of large mammals, it forms
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l relationships with ''P. bembae''. This is a mutually beneficial relationship where the hyphae of the fungus grow around the roots of the plant, enabling the fungus to receive moisture, protection and nutritive byproducts of the tree, and affording the tree greater access to soil nutrients. The ectomycorrhizal symbiosis is thought to contribute to the success of the dominant species, by allowing it access to nutrients otherwise unavailable. The
Congolian forests The Congolian rainforests are a broad belt of lowland tropical moist broadleaf forests which extend across the basin of the Congo River and its tributaries in Central Africa. They are the only major rainforests which absorb more carbon than they ...
encompass an
ecoregion An ecoregion (ecological region) or ecozone (ecological zone) is an ecologically and geographically defined area that is smaller than a bioregion, which in turn is smaller than a biogeographic realm. Ecoregions cover relatively large areas of l ...
known for its
species richness Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative a ...
and
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
, which is spread across four countries:
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the C ...
,
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
,
Republic of Congo The Republic of the Congo (french: République du Congo, ln, Republíki ya Kongó), also known as Congo-Brazzaville, the Congo Republic or simply either Congo or the Congo, is a country located in the western coast of Central Africa to the w ...
, and the
Central African Republic The Central African Republic (CAR; ; , RCA; , or , ) is a landlocked country in Central Africa. It is bordered by Chad to the north, Sudan to the northeast, South Sudan to the southeast, the DR Congo to the south, the Republic of th ...
.


References


External links

* * — contains original description {{Good article bembae Fungi of Africa Fungi described in 2009