Banksiamyces
Encyclopedia
Banksiamyces is a genus
of fungi in the Helotiales
order, with a tentative placement in the family Helotiaceae
. The genus contains four species, which grow on the seed follicles of the dead "cones" (flower spikes) of various species of Banksia
, a genus in the plant family Proteaceae
endemic to Australia. Fruit bodies of the fungus appear as small (typically less than 10 mm diameter), shallow dark cups on the follicles
of the Banksia fruit. The edges of dry fruit bodies fold inwards, appearing like narrow slits. The first specimens of Banksiamyces, known then as Tympanis toomansis, were described in 1887. Specimens continued to be collected occasionally for almost 100 years before becoming examined more critically in the early 1980s, leading to the creation of a new genus to contain what was determined to be three distinct species, B. katerinae, B. macrocarpus, and B. toomansis. A fourth species, B. maccannii, was added in 1984.
and Christopher Edmund Broome
described a species of fungus they named Tympanis toomansis, collected from dead cones of Banksia growing on the banks of the Tooma River
in southern New South Wales
, Australia. Its generic placement was a result of its resemblance to Tympanis
, a genus in the Helotiaceae
family of the Ascomycota
.
Additional collections, then still believed to be T. toomansis, were made from South Australia in 1952, again on dead cones of unspecified Banksia, and also in 1956 on dead cones of Banksia marginata
. In 1957 and 1958, R. W. G. Dennis
redescribed the species, and after consultation with Canadian mycologist James Walton Groves, who had earlier completed a monograph
on the genus Tympanis, transferred the taxon to the genus Encoelia
(Sclerotiniaceae family). Encoelia species are small, tough, brownish discomycetes
that typically grow in clusters on hardwood or woody substrates. Because the original collections were incomplete and certain microscopic features inadequately described, various collections made from Australia were presumed to be variations of the original 1887 collection.
In the 1980s, the availability of fresh specimens of the fungus—collected by Australian mycologist Bruce A. Fuhrer
from the cones of Banksia spinulosa
—prompted Gordon Beaton and Gretna Weste to reexamine the previous collections. Various apparent and microscopic differences were found which suggested that three distinct species were represented in the collections, and further, the species differed enough from other Encoelia species to warrant the creation of a new genus, which Beaton and Weste named Banksiamyces. The three Banksiamyces species they described in 1982 were the original B. toomansis (those initially named T. toomansis), as well as B. macrocarpus and B. katerinae. A fourth species, B. maccannii, was added to the genus by the same authors in 1984. A 2006 study identified two additional taxa
that did not quite meet the description for previously published species; these have been called Banksiamyces aff. macrocarpus and Banksiamyces aff. toomansis. Some existing species were found on other banksia species, so evidence strengthened that the individual Banksiamyces fungi did not exclusively parasitize
only one banksia species, a suggestion proposed by Beaton and Weste in 1982.
Banksiamyces is classified
in the Helotiaceae, a widespread but poorly known family
of fungi, many species of which are saprobic on herbaceous or woody tissues. The placement in the Helotiaceae is tentative, and no molecular analysis has yet been performed that might clarify the phylogenetic relationships of Banksiamyces to other taxa in the Helotiales
order. Based on physical similarity, Wen-Ying Zhuang included Banksiamyces under Encoelia in his 1998 study of the Encoelioideae subfamily of the Helotiaceae. He admitted, however, that he had not examined any specimens.
(stalk), colored dark brown to black with a dark grey center. When dry, the apothecia are covered with a whitish powder. The edges of the cup may be rolled inwards (especially when dry), or be twisted and somewhat flattened. Both the outermost tissue
layer (the ectal excipulum) and the tissue of the stalk are made of fungal cells with brown pigments that can be variously thick- or thin-walled, covered with small particles (granules), and spherical to ellipsoid. The middle tissue layer (the medullary excipulum) of both the cup and the internal tissue layer (the medulla) of the stalk contain a layer of tissue made of hyphae similar to the ectal excipulum. A second layer of tissue is made of hyphae that are translucent
and gelatinous; this layer may be present in either the medullary excipulum, the medulla, or both.
The asci
are elongated reproductive structures that bear ascospores, in groups of eight. Banksiamyces species have asci that are cylindrical to club-shaped, and contain a plug at their extreme tips that will absorb color when stained with iodine
. The ascospores may be arranged in one or two rows (uniseriate and biseriate, respectively), or rarely, irregular. The ascospores are ellipsoid, translucent, have a slight curve and may be tapered; most ascospores contain two oil drops. When in the ascus, the ascospores are covered with a translucent mucilage
that is highly refractive
to light. Paraphyses
are filamentous hyphal cells present in the fertile spore-bearing tissue, distributed amongst the asci. The free ends of the paraphyses fork and branch, combining with the tips of the asci to form a translucent to brown pigmented layer of tissue.
B. katerinae is named after G. Beaton's wife, the senior author of the 1982 protologue. It was first discovered in 1964, growing on the seed follicles of dead cones of Banksia ornata
in the Mount Zero Area in the Grampians
in northwestern Victoria
.
B. maccannii, first described in 1984, was found on dead Banksia saxicola
cones. The specific epithet was chosen to honor Ian McCann, for his "discovery of the type collection and ... his years of ecological, educational and conservation work in the Victorian Grampians." The fungus is distinguished from the other Banksiamyces species by its larger asci, larger spores, and tapering paraphyses
tips. Further, the type collection was found fruiting in December and January, compared to winter and autumn for other Banksiamyces.
B. macrocarpus grows on the dead cones of Banksia spinulosa
, and was first collected near Tonimbuk
, Victoria in 1981. It is the type species
of Banksiamyces.
B. toomansis
is the species originally described and illustrated by Berkeley and Broome. The type collection was found on a cone of Banksia marginata on the banks of the Tooma River
of New South Wales
. It has also been recovered from a cone of Banksia sphaerocarpa
from near Busselton in Western Australia, B. nutans
, B. pulchella
, B. speciosa
, and B. occidentalis
, all from Mount Merivale, 20 km (12.4 mi) east of Esperance, B. baxteri
cultivated at Cranbourne Botanic Gardens
, B. integrifolia from the Blue Mountains, and B. marginata from Kangaroo Island. Synonyms
include Tympanis toomansis Berk. & Br., and Encoelia toomansis (Berk. & Br.). Its ascospores can range in shape from elliptical to cylindrical, and have dimensions of 6–10 by 2.5–3 µm
.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of fungi in the Helotiales
Helotiales
Helotiales is an order of the class Leotiomycetes within the division Ascomycota. According to a 2008 estimate, the order contains 10 families, 501 genera, and 3881 species.Helotiales is the largest order of inoperculate discomycetes...
order, with a tentative placement in the family Helotiaceae
Helotiaceae
The Helotiaceae are a family of fungi in the Helotiales order. The distribution of species in the family are widespread, and typically found in tropical areas...
. The genus contains four species, which grow on the seed follicles of the dead "cones" (flower spikes) of various species of Banksia
Banksia
Banksia is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes and fruiting "cones" and heads. When it comes to size, banksias range from prostrate woody shrubs to trees up...
, a genus in the plant family Proteaceae
Proteaceae
Proteaceae is a family of flowering plants distributed in the Southern Hemisphere. The family comprises about 80 genera with about 1600 species. Together with the Platanaceae and Nelumbonaceae they make up the order Proteales. Well known genera include Protea, Banksia, Embothrium, Grevillea,...
endemic to Australia. Fruit bodies of the fungus appear as small (typically less than 10 mm diameter), shallow dark cups on the follicles
Follicle (fruit)
In botany, a follicle is a dry unilocular many-seeded fruit formed from one carpel and dehiscing by the ventral suture in order to release seeds, such as in larkspur, magnolia, banksia, peony and milkweed....
of the Banksia fruit. The edges of dry fruit bodies fold inwards, appearing like narrow slits. The first specimens of Banksiamyces, known then as Tympanis toomansis, were described in 1887. Specimens continued to be collected occasionally for almost 100 years before becoming examined more critically in the early 1980s, leading to the creation of a new genus to contain what was determined to be three distinct species, B. katerinae, B. macrocarpus, and B. toomansis. A fourth species, B. maccannii, was added in 1984.
Taxonomy
In 1887, English mycologists Miles Joseph BerkeleyMiles Joseph Berkeley
Miles Joseph Berkeley was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology....
and Christopher Edmund Broome
Christopher Edmund Broome
Christopher Edmund Broome was a British mycologist.-Background and education:C.E. Broome was born in Berkhamsted, the son of a solicitor. He was privately schooled in Kensington and in 1832 was sent to read for Holy Orders with the curate of Swaffham Prior in Cambridgeshire...
described a species of fungus they named Tympanis toomansis, collected from dead cones of Banksia growing on the banks of the Tooma River
Tooma River
The Tooma River is a river in the Snowy Mountains of New South Wales, Australia.The Tooma River starts near Mount Jagungal in the Snowy Mountains and drains part of the western side of the range...
in southern New South Wales
New South Wales
New South Wales is a state of :Australia, located in the east of the country. It is bordered by Queensland, Victoria and South Australia to the north, south and west respectively. To the east, the state is bordered by the Tasman Sea, which forms part of the Pacific Ocean. New South Wales...
, Australia. Its generic placement was a result of its resemblance to Tympanis
Tympanis
Tympanis is a genus of fungi in the family Helotiaceae. The genus contains 29 species.-External links:* at Index Fungorum...
, a genus in the Helotiaceae
Helotiaceae
The Helotiaceae are a family of fungi in the Helotiales order. The distribution of species in the family are widespread, and typically found in tropical areas...
family of the Ascomycota
Ascomycota
The Ascomycota are a Division/Phylum of the kingdom Fungi, and subkingdom Dikarya. Its members are commonly known as the Sac fungi. They are the largest phylum of Fungi, with over 64,000 species...
.
Banksia marginata
Banksia marginata, commonly known as the Silver Banksia, is a species of tree or woody shrub in the plant genus Banksia found throughout much of southeastern Australia. It ranges from the Eyre Peninsula in South Australia, to north of Armidale, New South Wales, and across Tasmania and the islands...
. In 1957 and 1958, R. W. G. Dennis
R. W. G. Dennis
Richard William George Dennis, Ph.D. , was an English mycologist and plant pathologist.-Background and education:Dennis was born in Thornbury, Gloucestershire, the son of a schoolmaster. He was educated at Thornbury Grammar School and Bristol University, where he studied geology and botany,...
redescribed the species, and after consultation with Canadian mycologist James Walton Groves, who had earlier completed a monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...
on the genus Tympanis, transferred the taxon to the genus Encoelia
Encoelia
Encoelia is a genus of fungi in the family Sclerotiniaceae.-External links:*...
(Sclerotiniaceae family). Encoelia species are small, tough, brownish discomycetes
Discomycetes
Discomycetes is a former taxonomic class of Ascomycete fungi which contains all of the cup, sponge, brain, and some club-like fungi. It includes typical cup fungi like the scarlet elf cup and the orange peel fungus, and fungi with fruiting bodies of more unusual shape, such as morels, truffles and...
that typically grow in clusters on hardwood or woody substrates. Because the original collections were incomplete and certain microscopic features inadequately described, various collections made from Australia were presumed to be variations of the original 1887 collection.
In the 1980s, the availability of fresh specimens of the fungus—collected by Australian mycologist Bruce A. Fuhrer
Bruce A. Fuhrer
Bruce Alexander Fuhrer is an Australian mycologist. He was awarded the Australian Natural History Medallion in 1989 and is a Life Member of the Ringwood Field Naturalists Club. He has written many books on Australian fungi, including a Comprehensive Field Guide to Australian Fungi. He worked for 25...
from the cones of Banksia spinulosa
Banksia spinulosa
The Hairpin Banksia is a species of woody shrub, of the genus Banksia in the Proteaceae family, native to eastern Australia. Widely distributed, it is found as an understorey plant in open dry forest or heathland from Victoria to northern Queensland, generally on sandstone though sometimes also...
—prompted Gordon Beaton and Gretna Weste to reexamine the previous collections. Various apparent and microscopic differences were found which suggested that three distinct species were represented in the collections, and further, the species differed enough from other Encoelia species to warrant the creation of a new genus, which Beaton and Weste named Banksiamyces. The three Banksiamyces species they described in 1982 were the original B. toomansis (those initially named T. toomansis), as well as B. macrocarpus and B. katerinae. A fourth species, B. maccannii, was added to the genus by the same authors in 1984. A 2006 study identified two additional taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
that did not quite meet the description for previously published species; these have been called Banksiamyces aff. macrocarpus and Banksiamyces aff. toomansis. Some existing species were found on other banksia species, so evidence strengthened that the individual Banksiamyces fungi did not exclusively parasitize
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
only one banksia species, a suggestion proposed by Beaton and Weste in 1982.
Banksiamyces is classified
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....
in the Helotiaceae, a widespread but poorly known family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of fungi, many species of which are saprobic on herbaceous or woody tissues. The placement in the Helotiaceae is tentative, and no molecular analysis has yet been performed that might clarify the phylogenetic relationships of Banksiamyces to other taxa in the Helotiales
Helotiales
Helotiales is an order of the class Leotiomycetes within the division Ascomycota. According to a 2008 estimate, the order contains 10 families, 501 genera, and 3881 species.Helotiales is the largest order of inoperculate discomycetes...
order. Based on physical similarity, Wen-Ying Zhuang included Banksiamyces under Encoelia in his 1998 study of the Encoelioideae subfamily of the Helotiaceae. He admitted, however, that he had not examined any specimens.
Description
The fruit bodies, or apothecia, of Banksiamyces species are cup-shaped receptacles borne on a stipeStipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
(stalk), colored dark brown to black with a dark grey center. When dry, the apothecia are covered with a whitish powder. The edges of the cup may be rolled inwards (especially when dry), or be twisted and somewhat flattened. Both the outermost tissue
Tissue (biology)
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...
layer (the ectal excipulum) and the tissue of the stalk are made of fungal cells with brown pigments that can be variously thick- or thin-walled, covered with small particles (granules), and spherical to ellipsoid. The middle tissue layer (the medullary excipulum) of both the cup and the internal tissue layer (the medulla) of the stalk contain a layer of tissue made of hyphae similar to the ectal excipulum. A second layer of tissue is made of hyphae that are translucent
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
and gelatinous; this layer may be present in either the medullary excipulum, the medulla, or both.
The asci
Ascus
An ascus is the sexual spore-bearing cell produced in ascomycete fungi. On average, asci normally contain eight ascospores, produced by a meiotic cell division followed, in most species, by a mitotic cell division. However, asci in some genera or species can number one , two, four, or multiples...
are elongated reproductive structures that bear ascospores, in groups of eight. Banksiamyces species have asci that are cylindrical to club-shaped, and contain a plug at their extreme tips that will absorb color when stained with iodine
Iodine
Iodine is a chemical element with the symbol I and atomic number 53. The name is pronounced , , or . The name is from the , meaning violet or purple, due to the color of elemental iodine vapor....
. The ascospores may be arranged in one or two rows (uniseriate and biseriate, respectively), or rarely, irregular. The ascospores are ellipsoid, translucent, have a slight curve and may be tapered; most ascospores contain two oil drops. When in the ascus, the ascospores are covered with a translucent mucilage
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...
that is highly refractive
Refraction
Refraction is the change in direction of a wave due to a change in its speed. It is essentially a surface phenomenon . The phenomenon is mainly in governance to the law of conservation of energy. The proper explanation would be that due to change of medium, the phase velocity of the wave is changed...
to light. Paraphyses
Paraphyses
Paraphyses are part of the fertile spore-bearing layer in certain fungi. More specifically, paraphyses are sterile filamentous hyphal end cells composing part of the hymenium of Ascomycota and Basidiomycota interspersed among either the asci or basidia respectively, and not sufficiently...
are filamentous hyphal cells present in the fertile spore-bearing tissue, distributed amongst the asci. The free ends of the paraphyses fork and branch, combining with the tips of the asci to form a translucent to brown pigmented layer of tissue.
Species
Banksia ornata
The Desert Banksia is a species of shrub in the plant genus Banksia which grows up to 3 m tall. It occurs in western Victoria, and in South Australia, where it is common on the Eyre Peninsula, Kangaroo Island and east of Adelaide, but is apparently absent from Yorke Peninsula.It tends to grow...
in the Mount Zero Area in the Grampians
Grampians National Park
The Grampians National Park is a national park in Victoria, Australia, 235 kilometres west of Melbourne. The Park was listed on the Australian National Heritage List on 15 December 2006 for its outstanding natural beauty and being one of the richest indigenous rock art sites in south-eastern...
in northwestern Victoria
Victoria (Australia)
Victoria is the second most populous state in Australia. Geographically the smallest mainland state, Victoria is bordered by New South Wales, South Australia, and Tasmania on Boundary Islet to the north, west and south respectively....
.
B. maccannii, first described in 1984, was found on dead Banksia saxicola
Banksia saxicola
Banksia saxicola, commonly known as the Grampians Banksia is a species of tree or shrub in the plant genus Banksia. It occurs in Victoria in two distinct populations, one in The Grampians and the other on Wilsons Promontory....
cones. The specific epithet was chosen to honor Ian McCann, for his "discovery of the type collection and ... his years of ecological, educational and conservation work in the Victorian Grampians." The fungus is distinguished from the other Banksiamyces species by its larger asci, larger spores, and tapering paraphyses
Paraphyses
Paraphyses are part of the fertile spore-bearing layer in certain fungi. More specifically, paraphyses are sterile filamentous hyphal end cells composing part of the hymenium of Ascomycota and Basidiomycota interspersed among either the asci or basidia respectively, and not sufficiently...
tips. Further, the type collection was found fruiting in December and January, compared to winter and autumn for other Banksiamyces.
B. macrocarpus grows on the dead cones of Banksia spinulosa
Banksia spinulosa
The Hairpin Banksia is a species of woody shrub, of the genus Banksia in the Proteaceae family, native to eastern Australia. Widely distributed, it is found as an understorey plant in open dry forest or heathland from Victoria to northern Queensland, generally on sandstone though sometimes also...
, and was first collected near Tonimbuk
Tonimbuk, Victoria
Tonimbuk is a bounded rural locality in Victoria, Australia. Its Local Government Area is the Shire of Cardinia. At the 2006 Census, Tonimbuk had a population of 275.-References:...
, Victoria in 1981. It is the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
of Banksiamyces.
B. toomansis
Banksiamyces toomansis
Banksiamyces toomansis is a species of fungus in the Helotiaceae family. It was first described as Tympanis toomansis by Miles Joseph Berkeley and Christopher Edmund Broome in 1886, and transferred to the genus Encoelia in 1957 by R.W.G. Dennis. Gordon William Beaton transferred it to Banksiamyces...
is the species originally described and illustrated by Berkeley and Broome. The type collection was found on a cone of Banksia marginata on the banks of the Tooma River
Tooma River
The Tooma River is a river in the Snowy Mountains of New South Wales, Australia.The Tooma River starts near Mount Jagungal in the Snowy Mountains and drains part of the western side of the range...
of New South Wales
New South Wales
New South Wales is a state of :Australia, located in the east of the country. It is bordered by Queensland, Victoria and South Australia to the north, south and west respectively. To the east, the state is bordered by the Tasman Sea, which forms part of the Pacific Ocean. New South Wales...
. It has also been recovered from a cone of Banksia sphaerocarpa
Banksia sphaerocarpa
Banksia sphaerocarpa, commonly known as the Fox Banksia or Round-fruit Banksia, is a species of shrub or tree in the plant genus Banksia . It is generally encountered as a 1–2 m high shrub, and is usually smaller in the north of its range...
from near Busselton in Western Australia, B. nutans
Banksia nutans
Banksia nutans, commonly known as Nodding Banksia, is a species of shrub native to the south coast of Western Australia in the genus Banksia...
, B. pulchella
Banksia pulchella
The Teasel Banksia is a species of small shrub in the plant genus Banksia. It occurs on the south coast of Western Australia from Fitzgerald River National Park east to Israelite Bay....
, B. speciosa
Banksia speciosa
The Showy Banksia is a species of large shrub or small tree in the plant genus Banksia. It reaches up to 8 m in height...
, and B. occidentalis
Banksia occidentalis
The Red Swamp Banksia or Waterbush is a species of shrub or small tree in the plant genus Banksia. It occurs on the south coast of Western Australia in three disjunct populations: at Augusta, around Albany and in the Esperance area.A 1980 field study at Cheyne beach showed it to be pollinated by...
, all from Mount Merivale, 20 km (12.4 mi) east of Esperance, B. baxteri
Banksia baxteri
The Baxter's Banksia , also known as Bird's Nest Banksia, is a species of shrub in the plant genus Banksia. It occurs along the south coast of Western Australia between Albany and Esperance....
cultivated at Cranbourne Botanic Gardens
Royal Botanic Gardens, Cranbourne
The Royal Botanic Gardens, Cranbourne, is a division of the Royal Botanic Gardens, Melbourne. It is located in the Melbourne suburb of Cranbourne, about 45 km south-east of the Melbourne city centre....
, B. integrifolia from the Blue Mountains, and B. marginata from Kangaroo Island. Synonyms
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
include Tympanis toomansis Berk. & Br., and Encoelia toomansis (Berk. & Br.). Its ascospores can range in shape from elliptical to cylindrical, and have dimensions of 6–10 by 2.5–3 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
.