''Banksiamyces'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

fungi 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 ...

in the order

Helotiales Helotiales is an order of the class Leotiomycetes within the division Ascomycota. The taxonomy within Helotiales has been debated. It has expanded significantly as genomic techniques for taxonomical identification have become more commonly used. ...

, with a tentative placement in the family

Helotiaceae The Helotiaceae are a family of fungi in the order Helotiales. The distribution of species in the family are widespread, and typically found in tropical areas. There are 117 genera and 826 species in the family. A question mark after the genus ...

. The genus contains four species, which grow on the seed follicles of the dead

infructescence Infructescence (fruiting head) is defined as the ensemble of fruits derived from the ovaries of an inflorescence. It usually retains the size and structure of the inflorescence. In some cases, infructescences are similar in appearance to simple fru ...

s or "cones" of various species of ''

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. ''Banksias'' range i ...

'', a genus in the plant family

Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...

endemic to Australia.

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 ...

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.

Taxonomy

In 1887, English mycologists

Miles Joseph Berkeley Miles Joseph Berkeley (1 April 1803 – 30 July 1889) was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology. Life Berkeley was born at Biggin Hall, Benefield, Northamptonshire, and educated at R ...

and

Christopher Edmund Broome Christopher Edmund Broome (24 July 1812 – 15 November 1886) 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 ...

described a species of fungus they named ''Tympanis toomansis'', collected from dead infructescences ("cones") of ''Banksia'' growing on the banks of the

Tooma River Tooma River, a perennial stream that has had some of its flow diverted as a result of the Snowy Mountains Scheme, is part of the Murray catchment within the Murray–Darling basin and is located in the Australian Alpine region of New South Wale ...

in southern

New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...

, Australia. Its

generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...

placement was a result of its resemblance to ''

Tympanis ''Tympanis'' is a genus of fungi 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 class ...

'', a genus in the family

of the

Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...

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 ''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 Arm ...

''. In 1957 and 1958, R. W. G. Dennis redescribed the species, and after consultation with Canadian mycologist

James Walton Groves James Walton Groves (October 18, 1906 - May 6, 1970) was a Canadian mycologist born in Kinburn, Ontario on October 18, 1906 to John James and Laura Groves. He displayed an early interest in education and learning. He taught himself the alphabet f ...

, who had earlier completed a

monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...

on the genus ''Tympanis'', transferred the taxon to the genus '' Encoelia'' (family

Sclerotiniaceae The Sclerotiniaceae are a family of fungi in the order Helotiales. Many species in this family are plant pathogens. Genera * '' Asterocalyx'' * ''Botryotinia'' * ''Botrytis'' * '' Ciboria'' * '' Ciborinia'' * '' Coprotinia'' * '' Cudoniopsis'' ...

). ''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 Bruce Alexander Fuhrer OAM (31 December 1930 – 31 March 2023) was an Australian botanist and photographer, specialising in cryptogams. His photographic collection of fungi numbers more than 3000 species. Early life Born in Woollahra, Sydney, ...

from the cones of ''

Banksia spinulosa ''Banksia spinulosa'', the hairpin banksia, is a species of woody shrub, of the genus ''Banksia'' in the family Proteaceae, native to eastern Australia. Widely distributed, it is found as an understorey plant in open dry forest or heathland ...

''—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 In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

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 Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

of fungi, many species of which are

saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ( ...

on herbaceous or woody tissues. The placement in the Helotiaceae is tentative, and no

molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...

analysis has yet been performed that might clarify the

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

relationships of ''Banksiamyces'' to other taxa in the

order. Based on physical similarity, Wen-Ying Zhuang included ''Banksiamyces'' under ''Encoelia'' in his 1998 study of the Encoelioideae subfamily of the Helotiaceae. He conceded, however, that he had not examined any specimens.

Description

The fruit bodies, or

apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...

, of ''Banksiamyces'' species are cup-shaped receptacles borne on a stipe (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 A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...

that can be variously thick- or thin-walled, covered with small particles (''granules''), and spherical to

ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface; that is, a surface that may be defined as the ...

. 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 In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable scattering of light. On a macroscopic scale (one in which the dimensions a ...

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 An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or ...

, 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 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 ...

. 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 is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of m ...

that is highly

refractive In physics, refraction is the redirection of a wave as it passes from one medium to another. The redirection can be caused by the wave's change in speed or by a change in the medium. Refraction of light is the most commonly observed phenomeno ...

to light. Paraphyses are

filamentous The word filament, which is descended from Latin ''filum'' meaning " thread", is used in English for a variety of thread-like structures, including: Astronomy * Galaxy filament, the largest known cosmic structures in the universe * Solar filament ...

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

Distribution of ''Banksia'' species (the host of ''Banksiamyces''), grouped into western species, ''

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Taxonomy

Description

Species

References