News
2023
This study by Putra et al. documents the discovery and identification if Inosperma subsphaerosporum in Indonesia (Borneo). This species was first described from tropical forests in south China, but in Indonesia Is. subsphaerosporum is routinely collected for food and sold in markets there where it is known locally as “jamur lengkuas” and as a delicacy. This is interesting because the species is closely related to others known to produce toxic secondary metabolites (muscarine, ibotenic acid, or muscimol).
Latha and colleagues describe a new interesting species of Pseudosperma, P. indicum,from tropical India. The new species is characterized in part by its association with Dipterocarpaceae and phylogenetic position near two other species – I. neglectum from Thailand, with a Fagaceae association, and I. “actinocephalum” nom. prov. (PBM2863) collected in southern Louisiana under a several hundred-year-old plantation oak tree. It is very helpful from a phylogenetic perspective that these authors consistently sequence rpb2 from these evolutionarily intriguing species from southern India.
Quite a bit of new work is coming out of Turkey describing new Inocybaceae from Mediterranean habitats in west Asia. This one, by Kaygusuz et al., describes a new species of Pseudosperma in association with pine. The phyologenetic position of this species, named P. pamukkalensis, is not exactly clear. It appears on an isolated branch relatively distant from members of the P. rimosum and P. araneosum groups. A possible explanation for this is the highly unusual and large number of autapomorphic inserts near the 5’ end of the 28S gene region with respect to all other Pseudosperma.
A new paper by Furtado et al. document ectomycorrhizas from roots of Guapria opposita (Nyctaginaceae) in Brazilian Atlantic Forest in southern Brazil. One of the ECMs is an Inocybe that appears to share a phylogenetic alliance with what I would call here the I. amazoniensis clade, a new world and largely tropical group of nodulose-spored Inocybes. The ECM root is mostly yellow but with a lovely lilac tip! The Inocybe ITS sequence produced from the root appears unique.
Pošta et al. described a new species of Inocybe – I. istriaca – from Croatia in this work[5] published in the journal Diversity. One of the unusual features of the new species is the large basidia. Inocybe istriaca belongs probably to as of yet unnamed clade including the following species: I. adorabilis, I. urceolicystis (syn. I. pseudoscabelliformis), I. dvaliniana, I. trollii, I. venustissima, and I. chalcodoxantha. Most of these have been very recently described from Europe and feature smooth spores. The greater position of this unusual clade within Inocybe is not entirely clear, however, I. istriaca appears most closely related to what I interpret as Murrill’s I. hebelomoides described from northern Florida. Inocybe olpidiocystis may belong here as well, or possibly with I. subconnexa in sect. Splendentes.
Francesco Dovana and colleagues did a revisionof the I. similis complex and species suggested as similar to it. In this work, the authors found that I. vulpinella is a later synonym of Bresadola’s I. similis and I. immigrans, originally described from eastern Canada by David Malloch, is a later synonym of I. chondrospora from Europe. In addition, the two species are distantly related. Inocybe similis is nested within the I. xanthomelas clade (normally a clade with nodulose-spores and entirely pruinose stipe), whereas I. chondrospora is placed in its own clade, the “/chondrospora CLADE”, together with I. kuberae and an unidentified group of environmental sequences. The I. chondrospora clade may be most closely related to nodulose-spores species I. fibrosoides, I. phaeosticta, and I. quercicola among others. An interesting finding of this study is that the distribution of caulocystidia on the stipe of I. similis varies, so much so that Kuyper recognized them in his different “supersections” of Cortinatae and Marginatae, which were later shown by early phylogenetic studies to be artificial groups. Dovana et al. suggest that development of the velipellis influences the distribution of caulocystidia on the stipe (but which direction?). Note that another European species with slight angular-smooth spores, I. diabolica, has also been recovered in the I. xanthomelas clade. A similar situation is also presented in the I. soluta group.
A new species from Pakistan, I. nigroumbonta, was described to accommodate a nodulose-spored species with phylogenetic affinities to I. subsect. Oblectabiles, a group formerly referred to by Matheny (in Matheny & Bougher 2017 and Aignon et al. 2022) as the I. calida clade. Turns out prior sequenced specimens of I. oblectabilis were misidentified and bona fide I. oblectabilis was united, and thus equated, with the I. calida clade. Esteve-Raventós and colleagues determined that the former Oblectabiles is now referable to as the Pseudohiucla clade. Other species now in subsect. Oblectabiles include I. subradiata from eastern North America and several species from Australia and southeast Asia such as I. sejuncta, I. tumidula, I. lineata, and I. parvibulbosa and its variants. An undescribed species from Indiana, I. sp. IN18, is also in this group. The publication of I. nigroumbonata is in one of those large multi-authored and inclusive articles in FUSE (too large to link here in its entirety), published online 14 Jun 2023 with Lebeuf as the lead author.
Matheny et al. led a collaborative study that made possible a revision of the genus Mallocybe from eastern North America. The work uncovered 16 species of Mallocybe from the region, four of which were described as new. Importantly as well, the study established a useful infrageneric clade/lineage arrangement within Mallocybe, based in part of discovery of two southeast lineages, and also found that most species of Mallocybe had undetectable amounts of muscarine. A number of old names described by Peck and Murrill were revitalized based on studies of their types and collecting efforts at or near type localities (e.g., Gainesville, Florida and Lake Placid, New York) connecting the types with modern reference materials. A key to eastern North American species was also included.
The North American Mallocybe work followed up on a nice and in-depth study of Mallocybe from China. In this work by Hu et al., the authors outlined three major clades within Mallocybe and described three new species, all of which were characterized by elongated cheilocystidia. The three new species cluster in one of the three major clades, which we labeled as the Unicolores clade, together with several eastern North American species now known as M. unicolor, M. multispora, and M. fulviceps. The group is noteworthy because of the east Asian-east North American phylogenetic alliance.
In addition to the discoveries in Mallocybe from east Asia and North America, Saba and colleagues described two new species in this study from pine forests in Pakistan – M. pokistanica and M. pinicola. It was a good year for Mallocybe!
A new smooth-spored species, Incoybe bhurbanensis, is described from Pakistan. It is most closely related to I. demetris, a member of the greater Hystrices-Lactiferae clade. Both are also very closely related to an unclarified North American species known from specimens and environmental samples in the American southwest.
A Hungarian study took place that identified 28 species of Inocybaceae (23 Inocybe, three Mallocybe, two Pseudosperma) from urban habitats in Budapest.
A new smooth-spored Inocybe – I. hamadryadis – is described in Phytotaxa as new from Europe and west Asia. It is most closely related to I. tenuicystidiata from Europe and other species in the greater clade containing sections Hysterices and Lactiferae.
Eberhardt and colleagues, in this revision of species attributed to Hebeloma by W.A. Murrill in North America, addressed and resolved several species of Inocybe, Inosperma, and Pseudosperma. Murrill’s Hebeloma dryophilum was moved to Inocybe and found to match a large number of sequenced specimens from northern California, Oregon, Washington, and B.C. Hebeloma floridanum is now recognized as Inocybe hebelomoides Murrill, non Kühner, the latter now recognized as Mallocybe hebelomoides. Hebeloma praefarinaceum was sequenced and supported in Inocybe near I. metrodii in the I. leiocephala group. Two species described by Peck – I. sterlingii and I. vatricosoides – are considered synonyms of Hebeloma excedens. Molecular annotation of Murrill’s Hebeloma pallidifolium support its placement in Inosperma; in my estimation this can be considered synonymous with Inosperma lanatodiscum. Murrill’s Hebeloma californicum was placed in Pseudosperma, an oak associate described from Stanford University. Unfortunately, molecular annotations could not be obtained from I. cinchonensis from Jamaica.
A new species in the Inocybe cryptocystis group has been described from Pakistan: I. subhimalayanensis.
Inocybe distans is described as new from native habitat in Australia and eucalypt plantations in Spain and Argentina. It is a member of the I. tenax group, which belongs to a hyper-diverse clade of smooth-spored, metuloid, southern hemisphere Inocybe.
2022
A new species, Inocybe viscida, is described in the Inocybe geophylla group from Turkey and published in the Turkish Journal of Botany.
Two new species of Inocybe were described in one of the large inclusive series put out by Persoonia in their ‘Fungal Planet description sheets’. This one has Tan as the lead author (with about 100 co-authors) and includes sheets 1436 – 1477. The two new species are Inocybe longistipitata, described from Pakistan, and I. udicola from Finland. Inocybe longistipitata, save many ITS polymorphic positions, appears identical to the earlier described, but invalidly named, I. quercicola, also from Pakistan. If we equate these two species, then there is no reason to post-validate the name I. quercicola, although no mention is made of this by the authors on their species sheet. The second new species, I. udicola, captures a new taxon with superficial similarities with I. posterula. Good news is that the epitype of I. posterula has been sequenced and found to be distantly related to I. udicola.
He et al. describe the new species, Inocybe carpinicola, from a tropical region in China (Hainan Province). The species is placed phylogenetically in an interesting clade that has not received much attention other than slowly accumulating species over the years now. The clade, which I have previously refered to as the I. stellata group [now sect. Leptocybe], is dominated by nodulose-spored species mainly of tropical origin in south Asia (e.g. China, Thailand, India) and (sub)tropical regions of Australasia (Queensland, New Caledonia). At least two north temperate species belong here as well, I. pseudoasterospora and I. perlucida. If the topology of He et al. is true, then the two north temperate species signify a transition to colder temperate climates compared to a likely Old World tropical ancestor.
Kaygusuz produced a work that documents the discovery of Inosperma bongardii in the Aegean region of Turkey for the first time. This species has been confirmed with molecular data from European areas including Italy, Norway, and Sweden, but previously recorded from west Asia in Lebanon (Is. bongardii has also been recorded with molecular data from Estonia, Finland, and in the U.K. as var. pisciodora). Inosperma bongardii was shown here to be most closely related to an undescribed or unnamed species from China. In Turkey, Is. bongardii occurs in oak forests. In northern Europe (per Jacobsson & Larsson in Funga Nordica), Is. bongardii is reported on calcareous soils in hardwood forests mainly with Fagus and Quercus, but sometimes with Picea or in subalpine forests with Betula.
This study out of China by Deng et al. describs a new tropical member of the genus Inosperma – Is. zonativeliferum – from a Castanopsis-dominated forest. Muscarine was also detected in the new species, which clusters without a dozen others in “Old World tropical clade 2”. This latter clade currently lacks a formal name but includes Inosperma species from Papua New Guinea and south Asia (India, Thailand, Hainan Province). “Old World tropical clade 1” comprises African taxa only at this point, but not all African Inosperma occur in this particular group.
Two of the co-authors from the above study also published a work, in which five new species of Inosperma were described from China – Is. longisporum, Is. nivalellum, Is. sphaerobulbosum, Is. squamulosobrunneum, and Is. squamulosohinnuleum. This increases the number of Inosperma species in China to 17. The authors also conducted secondary metabolite analyses and detected the presence of various neurotoxins in unique combinations depending on the species. For example, Is. nivalellum was muscarine positive, but Is. longisporum and Is. squamulosohinnuleum were found to contain both ibotenic acid and muscimol [very unusual!], whereas Is. squamulosobrunneum contained only muscimol. Consistent with prior reports regarding Is. calamistratum and allies, psilocybin was not detected in any of the new species. The report of ibotenic acid and/or muscimol in Inosperma is the first discovery of these toxins in the Inocybaceae .
Three new species of nodulose-spored Inocybe were formally described after many years from Miombo woodlands in Zambia in one of the large mosaic ‘fungal biodiversity profiles’ articles – this one in the journal Cryptogamie, Mycologie. The three new species are: Inocybe hebes, I. leucophaea, and I. media described by Eyssartier & Buyck. Inocybe leucophaea belongs to a unique group of species from tropical Africa and tropical India, including the Indian species I. snigdha and I. pingala. The phylogenetic positions of the other two species are not yet clear with available rDNA data only. The authors suggest morphological similarities between I. hebes and the originally north temperate I. curvipes, but an affiliation with I. sect. Lacerae does not seem likely based on phylogenetic distance at least. An earlier published phylogenetic and diversification study by Ryberg and myself show I. hebes sister as yet to an unpublished Zambian species, I.”fuscescentipes” (now indicated in the 2022 study as I. subfuscescentipes nom. prov., but any more inclusive alliance is not possible to deduce with 28S data alone). Inocybe media is indicated in association with I. asterospora and allies by Buyck et al., but in published phylogenies I. media is shown as sister to I. napipes with poor support (Ryberg & Matheny 2012) or as sister to the now-published Thai species I. lineata (Matheny et al. 2009). If the latter is true, then I. media might merit consideration in I. subsect. Oblectabiles or possibly with an undescribed group of Inocybe including I. cicatricata and I. bulbinella.
This study out of China by Zhao et al. describes two new species of Pseudosperma and made the new combination P. rubrobrunneum for an Indian name. The authors produced ITS, 28S, and rpb2 data, as well as muscarine analyses, that demonstrate the presence of muscarine in the new species. Both P. fulvidiscum and P. singulare are tropical in distribution (occurring at least in Hainan Province) where they associate with Carpinus.
This work by Kaygusuz et al. resulted in description of a new smooth-spored species of Inocybe – I. kusadasiensis - from the coast of the Aegean Sea in western Turkey. The species is a pine associate found on calcareous soils and is most closely related to species in the I. hirtella group.
Bandini and colleagues generated an important study, including even a table of contents page, published in Integrative Systematics. The work describes 11 new species of Inocybe mainly from Europe. In addition, new typifications are designated for I. griseovelata, I. lacera, I. soluta, I. subcarpta, I. tarda, and I. transitoria. Several synonymies and affirmation of previous suggested synonymies are also proposed. All in all, the taxonomic status for 29 species of Inocybe was assessed in this work. This will be extremely helpful to others going forward with future studies on the groups that include these taxa.
Another useful study by Esteve-Raventós et al. sheds more light on the Inocybe xanthomelas group. As a result, I. vaurasii is described as new accommodating in part the broadly applied name I. xanthomelas. The authors were also able to generate molecular annotations from the types of I. xanthomelas, I. humilis, and I. subrimosa, the latter a late 19th century species described by P. Karsten. Both I. humilis and I. subrimosa have encompassed earlier misapplications of the name I. xanthomelas. Now we just need a key!
Aïgnon et al. describe four new tropical species of Inocybe from west Africa. These include I. beninensis, I. flavipes, I. fuscobrunnea, and I. pallidiangulata. The treatment includes a 3-gene region phylogeny of Inocybe.
Matheny, students, and colleagues published a revision of Inocybe species in the I. grammata group. Four gene regions were studied and separate gene trees presented. The results support description of three new species – I. albodiscoides (long recognized as I. albodisca of the Pacific Northwest) and two fagaceous associated species, I. panamica and I. velicopia, that occur in Central America, one of which is distributed as far north as New York. The I. grammata group contains at least eight confirmed species (more may belong here) and is formally recognized as a I. sect. Albodiscae. The work was published in Brittonia.
A series of papers documenting or describing new species of Pseudosperma has been published by a series of overlapping Chinese authors. One attribute they all have in common is the presentation of toxin data (presence of muscarine and/or phalloidin), which serves as a useful accompaniment to the usual molecular systematic work. The first, describes the new species P. arenarium from Asia and Europe and confirms the presence of muscarine in this species. The second documents P. umbrinellum from east Asia, which also occurs in Europe and eastern North America, and presents for the first time the presence of the phallotoxin phalloidin (in addition to muscarine) in any Inocybaceae species. The third study describes two new muscarine-containing species of Pseudosperma from tropical regions of China associated with Carpinus (Betulaceae), P. fulvidiscum and P. singulare. Interestingly, the authors suggest the P. araneosum clade, inclusive of the new tropical Asian species and others from tropical regions of south Asia and northern Australia, form the sister group to the rest of Pseudosperma. However, other studies have shown P. pluviorum, from subtropical Queensland, is sister to the rest of Pseudosperma.
Bandini and colleagues produce another large study in Persoonia where 12 new species of Inocybe are described as new and several synonymous pairings are newly revealed.
A new nodulose-spored species, Inocybe squalida, is described from Quebec with an affinity to I. curvipes and I. lacera.
Muñoz and colleagues present a nice revision of species surrounding Inocybe pusio, a suite of species characterized by a lilac or violent colored stipe, the color of which often fades and thus presents difficulties in identification, and smooth spores. Three new species are described and two new combinations are made. Variation in the distribution of caulocystidia on the stipe was noted (some absent or restricted to the extreme apex, whereas in others caulocysitida can be readily found over the upper fourth or third of the stipe].
Five new species belonging to Mallocybe and Pseudosperma are described as new from north China by Mao and colleagues. The authors use a combination of rpb2, ITS, and 28S data to circumscribe the species phylogenetically. The two new Mallocybe species are found in the “core” clade within Mallocybe, whereas the three new Pseudosperma are scattered through the tree for this genus.
2021 – addendum
An additional 20 papers were published dealing with Inocybaceae systematics in 2021. Together with the 8 papers summarized below, this raises the number of studies published to 26 for 2021.
Two new species of Inocybe were described in one of the large inclusive Fungal Plant description sheet series (too large a file to link here): Inocybe corsica and Inocybe nivea. When I first saw the image of fruitbodies of I. corsica, described by Esteve-Raventós and colleagues from the Mediterranean, I was immediately reminded of a look-alike common in urban areas in Knoxville, Tennessee and in many other locations throughout eastern North America. Turns out I. corsica is closely related to but distinct from this species, as well as the ‘smooth-spored’ I. diabolica. All occur in the very diverse group referred to as the I. xanthomelas clade. In the same series, Larsson described a new species in the I. geophylla group – I. nivea, characterized by the association on calcareous ground with Salix in the subarctic (Svalbard and Jan Mayen) and unique phylogenetic position. Many other novelties in the I. geophylla group await to be described.
Parnmen and colleagues put together a very informative study of the identification, phylogenetic distribution, and muscarine analysis of 26 Inocybaceae collections involved in mushroom toxicity among 61 patients over a 9-year period in Thailand. Of the 26 samples, one appeared close to Inocybe fibrosoides but without typical muscarinic symptoms. All the others clustered in Pseudosperma and Inosperma with many species of Inosperma revealed. No cases attributable to Auritella, Mallocybe, Nothocybe, and Tubariomyces were found. A remarkable aspect to this study was that no single species could be identified with a sequenced described species. In Pseudosperma, I’m only aware of two described species of the genus found in Thailand – P. neglectum and P. keralense. Several others have been described from India and are to be expected in Thailand as well. More work and collaborative efforts are needed to diversity of southeast Asian Inocybaceae . However, during the same year, a Chinese team described as new Is. hainanense, which is the same as Inosperma sp. D152 and D89 involved in the toxicity cases. The Chinese team, led by Deng et al., also detected a relative high amount of muscarine in Is. hainanense.
Aïgnon continues to publish studies from his PhD dissertation at the University of Parakou in Benin. The latest work presents a review of Inocybaceae documented from West Africa, including 10 West African nations, and documents 19 species of mainly nodulose-spored Inocybe and various species of Mallocybe, Pseudosperma, and Inosperma. Species of Auritella and Tubariomyces have also been recorded from Africa but outside the study range.
Bandini et al. produce a landmark study describing 18 new species of smooth-spored Inocybe with only the stipe apex pruinose, if at all. The new species are couched in a taxonomic revision of relevant types mainly from Europe and presented in a phylogenetic context of ITS and 28S data. The study is noteworthy because the Inocybaceae diversity of central Europe was presumably relatively well studied and considered relatively well-known. Despite the novelties, the authors also suggest a dozen synonymies, which help clean up the taxonomy a great deal.
In a follow-up to their 2020 study also published in German, Bandini and colleagues describe an additional 13 species of Inocybaceae mainly from central Europe (3 species of Inosperma, 3 species of Pseudosperma, and 7 species of Inocybe). The species are distinguished from others by comparative morphology and ITS and at times 28S data.
In a series of fungal biodiversity profiles, Buyck and collaborators formally describe 3 new species of Inosperma from Zambia in south-central Africa. It’s nice to see these published at last, as I have been familiar with these taxa for some time now having published their DNA sequences more than a decade ago in a historical biogeographic study of Inocybaceae . In my opinion, however, one of the new species, Is. boeticum, is doubtfully different from Is. bulbomarginatum described earlier from Benin. In any case, the phylogeny of Inosperma is fascinating because of the rich assemblage of Old World tropical taxa from Africa, Asia, and Australasia that form a grade from which temperate species emerged.
Caiafa et al. produce one of the most interesting Inocybaceae studies that documents novel sequestrate forms from Patagonia. Up to this study, only four sequestrate taxa had been documented in the Inocybaceae from across the world. However, Caiafa et al. describe four new sequestrate species, including one that is strikingly truffle-like (not just secotioid). Seven of the eight sequestrate taxa are known only from the southern hemisphere. It is not clear if the species are vectored by animal mycophagists, but it’s not unreasonable to predict these taxa have lost the ability to produce muscarine.
Cho et al. document seven species of Inocybaceae from the Korean peninsula. Some were known previously only from southeast Asia (I. caroticolor, I. stellata) or Pakistan (Inosperma shawarense) and others from Europe (e.g., I. grammatoides, Mallocybe malenconii). These are important range extensions for these taxa.
After an earlier revision of the Inocybe leiocephala complex, Cullington confirms the presence of two species in the complex in the U.K. – I. lindrothii (new confirmation) and I. fuscescentipes (recent confirmation).
Dakalopoulos et al. establish ectomycorrhizal associations between Mallocybe heimii and two different plant genera – Cistus and Pinus – from sand dunes along a sea shore in the Mediterranean region. An rDNA phylogenetic analysis also supports the synonomy of M. arenaria with the priorable M. heimii.
Deng and colleagues describe a new species of Inosperma from Hainan Province in China characterized by (sub)globose basidiospores and a furfuraceous to fibrillose stipe covering. The species is placed in an Old World tropical clade representing taxa known only from south Asia and Australasia.
In a second study Deng and colleagues describe an additional two new species of Inosperma – Is. muscarium and Is. hainanense – from tropical China and demonstrate that both contain prodigious amounts of muscarine. The two species are also rather closely related to the known toxic species from India, Is. virosum.
Lately, some novel species of Inocybe and allies have been published in the Fungal Planet series as Fungal Plant description sheets. One such species was described in the I. lacera group as I. norvegica from Norway and Sweden by Larsson and Vauras. The I. lacera group contains several distinct phylogenetic species from North America that have probably not yet been described, and Peck’s I. infelix can probably be resurrected to apply to one of them. Inocybe pluppiana, I. impexa, I. helobia, and I. curvipes are all members of the I. lacera group. Possibly I. goniopusio as well, which may be sister to the rest.
A new species has been described in Mycotaxon from Pakistan, this one as Pseudosperma albobrunneum from Himalayan conifer forests. The species appears to be most closely related to Ps. dulcamaroides, Ps. mimicum, Ps. squamatum, and allies, so distantly related to the Ps. rimosum complex.
Another new species was described from Pakistan, Inocybe quercicola, with nodulose spores and an entirely pruinose stipe and marginate bulb. It is mostly closely related to I. fibrosoides from Europe and I. decemgibbosa, among others. However, the name was not validly published (the holotype was not specified) and subsequently replaced with the later description of I. longistipitata. It and the others mentioned here are allied with Inocybe subsect. Praetervisae.
A new species from Italy, Inocybe mediterranea, has been described in the I. cincinnata group by Marchettii and colleagues. An ITS phylogeny suggests a very close relationship between I. mediterranea and I. tiburtina and I. gracillima and I. obscuroides.
Bandini and colleagues describ a new nodulose-spored species with a blackening or graying stipe in the Inocybe xanthomelas group. There are a number of species in this group endemic to Europe. The authors refer to I. sect. Marginatae, but this is merely a taxonomic reference. Phylogenetically, sect. Marginatae will likely be restricted to the clade including its type, I. asterospora. The xanthomelas group is distantly related to this.
Two new species of Inocybe are described from Europe in one of the large publications referred to as “Fungal Plant description sheets” in Persoonia. The first of these, I. corsica, is yet another new species described in the I. xanthomelas group. The second is a member of the I. geophylla group and is called I. nivea; it occurs in the arctic with Salix.
Li and colleagues document a tropical species originally described from Malaysia, Inocybe squarrosolutea, in south and central China, and describe a new species, I. squarrosofulva, from subtropical beech forest in south China. Both are yellow, highly toxic with prodigious amounts of muscarine, nodulose-spored, and sister taxa. Both occupy a poorly resolved phylogenetic position within Inocybe but are sister to an undescribed and similarly bright yellowish fulvous nodulose-spored species from Cameroon.
A review of chemistry and toxicology of major bioactive substances in Inocybe was produced by a European research group. While helpful to summarize available data, it is disappointing the review missed our work that published new toxicology data for 30 species of Inocybaceae sampled world-wide in 2013 by Kosentka et al. and more recent work about a new putative hallucinogenic species of Inocybe from eastern North America in 2019.
2021
A research group in Benin led by a graduate student, Hyppolite Aïgnon, has been actively lately and published three studies all in 2021 on west African Inocybaceae . I will summarize these in succession below, in addition to several others from Europe describing new species of Inocybe.
The first describes three new species of Inosperma from tropical Africa. Inosperma is of particular interest because the genus is characterized by a grade of Old World tropical lineages from which the rest of Inosperma emerged such as the better known In. calamistratum and In. maculatum and allies. Inosperma africanum is also identified as the sister group to the rest of the genus with strong support and thus represents an important discovery.
The second study by Aïgnon and colleagues describes the first species of Mallocybe from Africa, M. africana. However, its phylogenetic position conflicts depending on the gene regions analyzed. rDNA sequences suggest an alliance with another tropical species from south Asia, M. errata. However, analysis of rpb2 only sequences suggest M. africana is sister to the rest of Mallocybe, which as above, would be an important discovery, if accurate. Other gene sequences will be needed to resolve this conflict. Bart Buyck first discovered M. africana some years ago in Zambian Miombo woodland with a student of his, G. Eyssartier. I’ve been aware of this species for some years thanks to Bart, but it went undescribed until re-discovered in west Africa by the Benin research group. One note of caution about the phylogenetic analysis presented here: the tip in the tree figure labeled as M. althoffiae (type) comprises an unusual ITS sequence (possibly chimeric) and the 28S is a contaminant of Ps. sororium from the western U.S., results reported earlier by Matheny et al. (2020).
The third study by Aïgnon et al. (2021) presents a review of Inocybaceae taxonomic diversity as documented from west Africa. The review reports about 20 species in the family from west Africa, the minority of which, not surprisingly, are undescribed. Moreover, many species from other parts of tropical Africa are “known unknowns” and also undescribed. Much systematic and taxonomic work remains to be done!
Dovana and colleagues (2021) describe a new smooth-spored species, Inocybe cervenianensis, from Italy, which is placed phylogenetically in a recently recognized new group, the I. flavoalbida clade. This group is characterized in part at least by their smooth spores and entirely pruinose stipes. The group constitutes several European species such as I. amblyospora and I. pseudoreducta, I. praecox from western North America, and I. flavoalbida from Papua New Guinea and northern Australia.
Another Italian study published the new species Inocybe messapica from oak woods in Italy, which was also confirmed from Estonia based on environmental sequences. The species was classified, based on morphological taxonomy, in I. sect. Splendentes, species with smooth spores and an entirely pruinose stipe and typically absence of a cortina and bulbous stipe base. However, sect. Splendentes has been shown to be non-monophyletic by Kropp et al. (2010).
Mešič et al. (2021) have described a new species, Inocybe brijunica, from Croatia, which is placed in the monophyletic group I. sect. Hystrices, a group characterized by smooth-spored taxa such as I. hystrix, I. melanopoda, I. glabripes, I. aeruginascens, and I. chondroderma (the ‘PDAB’ Inocybe) among others. Inocybe brijunica is unusual because of the conspicuous orange to orange-brown membrane-like material present above or near the base of the stipe.
2020 - addendum
As an addendum to 2020 notes of Inocybaceae publications, three other studies were also produced raising the number of published 2020 works to 21 for the year at least.
Cervini and colleagues describe a new species of Inosperma, Is. vinaceum, from Italy with morphological similarities to Is. rhodiolum and the eastern North American Is. vinaceobrunneum. One possible additional species may be undescribed in this group, together with Is. adaequatum and Is. erubescens; this species is a western U.S. entity that associates with oaks in California.
Fan and Tolgor produced a study including new rpb2 sequences for two new smooth-spored species of Inocybe described from northwestern China in Gansu Province. The new species are I. muricellatoides and I. gansuensis. Inocybe muricellatoides, as the epithet suggests, shares affinities with the European I. muricellata, I. hirtella var. bispora, and the American I. microteroxantha. Inocybe gansuensis is a member of the I. flocculosa group, a clade strongly supported as such by rpb2 data.
Xu and colleagues document a case of muscarine poisoning involving Chinese samples of Inocybe serotina. The two human patients survived. Inocybe serotina was originally described by C.H. Peck from New York and is known from sandy shores and sand dunes. Inocybe bulbosa and I. ammophila, two additional American names, are considered synonymous with I. serotina. Murrill’s I. praefarinacea seems similar as well. The phylogeny shown in the paper indicates that I. serotina is in the I. splendens group (I. sect. Splendentes Singer) together with I. splendentoides, I. terrifera, I. alluvionis, and I splendens.
2020
Overall, 2020 was a very productive year for research on systematics of Inocybaceae , particularly by European biologists. At least 18 papers were published, including description of 40 new taxa and a few range extension reports and contemporaneous descriptions of rare or unusual species. The year also saw a rather significant shift to recognition of 7 genera in the family, which appears to have been readily embraced. In 2020 alone, Inocybaceae systematists described 13 new species of Pseudosperma, 24 new species and 1 new variety of Inocybe, and 2 new species of Mallocybe. Almost all of this work is emanating from Europe and Asia with two studies from North and South America. I would estimate now that worldwide Pseudosperma contains ca. 83 species, Inocybe ca. 875 species, and Mallocybe ca. 57 species.
So, the first paper I’ll mention is here pertains to my work with Fernando Esteve-Raventós with help from a research technician, Alicia Hobbs, whereby we arranged a new systematic framework within Inocybe that establishes 7 genera within the family (previously, 3 genera and 4 clades were recognized by many authors). Arguments pro and con are presented in the paper that ultimately led to our decision to restructure Inocybe in the large sense, a process that actually began more than a decade ago when the genera Auritella and Tubariomyces were described. Penny Cullington drafted a response to the newly proposed Inocybaceae in Field Mycology.
Sesli and Bandini (2020) report the first occurrence of the recently described central European species Inocybe nothomixtilis from east Asia (Turkey).
Yu et al. (2020) described a new species of Pseudosperma – P. citrinostipes – that associates with the conifer Keteleeria (Pinaceae) in southwest China. The species appears to be sister to the bulk of the Ps. rimosum complex (but note the position of an additional undescribed species provisionally named Ps. “burrneicothurnata” from Texas). Attempts to distinguish these species morphologically is inherently difficult, but molecular data have the resolving power to distinguish most.
Jabeen and Khalid (2020) describe a new Pseudosperma from Pakistani Himalayan conifer forest called P. flavorimosum. The species falls in clade A of Pseudosperma or the P. rimosum complex but dwells with several undescribed or poorly clarified species from the southeast U.S. and Papua New Guinea.
The third study of 2020 to describe three new species of Pseudosperma comes from central Europe by Bandini and Oertel in Czech Mycology. In it, the authors also have sequenced the isotypes of P. aureocitrinum (autonomous) and P. copriniforme (synonymous with P. perlatum), a welcoming contribution. Of the new species, Ps. napaeanum is most closely related to the P. umbrinellum group. Pseudosperma amoris occupies a rather long branch but may be related to Ps. dulcamaroides, which would be a nice discovery since the former occupies its own lineage within the genus. Lastly, Ps. amabile is most closely related to Ps. rimosum, Ps. sororium, Ps. melliolens, and Ps. holoxanthum. Epitypification of Ps. rimosum is still needed, and the it may be possible to sequence the type of Ps. sororium to more firmly establish its identity.
The fourth Pseudosperma systematics paper was done by Saba et al. (2020) out of Pakistan. Here they describe three new species associated with Pinus based on ITS, 28S, and the mt 12S locus (mitochondrial small subunit rDNA). Pseudosperma pinophilum is another new species very close to Ps. rimosum in the strict sense but distinguished molecularly. Pseudosperma triaciculare and Ps. brunneoumbonatum form a grade giving rise to the Ps. rimosum complex. It’d be interesting to see how these species relate to Ps. citrinostipes mentioned above. The authors also made five new combinations in Inosperma, Mallocybe, and Pseudosperma.
Amazingly, in a work by Cervini et al. (2020), the fifth Pseudosperma systematics paper presented detailed descriptions, illustrations, and phylogenetic analyses for four Italian species previously published in Index fungorum – Ps. melle um, Ps. conviviale, and Ps. ponderosum (in the rimsoum complex) and Ps. salentinum in the Gracilissimum clade. All four species are characterized by a honey-like smell and Quercus association but do differ subtly by other morphological features and, of course, phylogenetic relatedness.
17 species of Inocybe were detailed here in this study by Ditte Bandini and colleagues (2020), published in German in Mycologica Bavarica, 13 of which were described as new and two species raised from variety to species rank. The work was primarily carried out in western Europe. The table in the paper indicates that many sequences of new species described by E. Ludwig from Germany should be available soon.
Wartchow (2020) produced a short paper here that describes a new nodulose-spored species, Inocybe cavalcantiae, from the Atlantic Forest in northern Brazil. Note the Neotropicsa are dominated by the presence of nodulose-spored species over smooth-spored ones in Inocybe.
Saba and Khalid (2020) described the new species Mallocybe velutina, which was discovered in conifer forests in association with pine in Pakistan. This is an important discovery because it adds another species to a distinct clade within Mallocybe that includes M. heimii and M. tomentosula. rDNA analyses of mine also place the tropical species M. errata and M. africana in this grouping, but there could be some concern this is due to a long branch artifact as rpb2 only analyses place M. africana as sister to the rest of Mallocybe (see discussion of M. africana above).
Mallocybe crassivelata was described by Dovana and colleagues (2020) in a Persoonia ‘fungal planet sheet’. The species is known thus far from Italy, Slovenia, and Spain, and is distinguished, in part, by the presence of a heavy white veil, similar in some respects to a few other species also with this condition (e.g., M. leucoblema, M. leucoloma, and possibly M. pallidotomentosa). Interestingly, M. crassivelata is shown to be the sister group (with strong support) to a clade of three environmental sequences from root systems of Dryas in arctic Alaska.
Bandini et al. (2020) describe another new species of Inocybe with smooth spores and a cortina from central and northern Europe – I. woglindeana, which they ascribe to I. sect. Tardae based on morphology.
An unusual Inocybe, I. mytiliodora,with an odor of mussels or shellfish was recently reported as new to the U.K. under birch on calcareous ground by Hobart and Henrici in a Field Mycology article (here). Inocybe mytiliodora was originally described from Finland, observed in nearby Estonia, and reported from western North America with Salix in a subalpine, calcareous, wet meadow in Wyoming. The authors of the FM article also point to the synonymy of I. pedemontana, which extends the species into northern Italy. I have produced ITS and rpb2 sequences from the type and North American material, and both are very similar. As suggested by Jukka Vauras I. mytiliodora is closely related to I. appendiculata, which differs by the odor of rotten meat but becoming spermatic where cut. Both of these are closely related to smooth-spored species I. picrosma, I. tenebrosa, and I. luteifolia (and possibly I. kauffmanii as well), but these are distinguished by their entirely pruinose stipes (see the global rpb2 Inocybaceae tree published as a supplemental file in Matheny and Kudzma (2019).
Inosperma virosum (syn. Inocybe virosa), a tropical species that occurs in south Asia, underwent a detailed metabolic profile by Latha and colleagues (2020). The metabolites methyl palmitate and phytol, in addition to muscarine, were identified by gas chromatography and liquid chromatography. Bioassays were also explored. The species is most closely related to others in Inosperma that also are distributed in the south Asian tropics and Australasia (New Guinea). These species probably associate with Dipterocarpaceae and Fagaceae (Castanopsis), and one from New Guinea associates with Gnetum but best not to eat ‘em!
The species Inocybe antoniniana is described as new by Bandini et al. (2020) in a paper published in Sydowia. This Eurasian species is placed in the traditional sect. Marginatae, but this section is not monophyletic. Phylogenetic analysis places I. antoniniana in a large inclusive clade of species that include I. diabolica, I. xanthomelas, I. intricata, I. phaeocystidiosa, I. krieglsteineri, and I. invadens among others. This strongly supported clade currently does not have a name. This clade is distantly related to the I. mixtilis and I. praetervisa groups.
Inocybe melleiconica was originally described by Grund and Stuntz from spruce forests in Nova Scotia but probably occurs elsewhere in southeast Canada (such as Ontario) and perhaps the northeast U.S. It’s a smooth-spored species with an entirely pruinose stipe often with a bulbous base. Vauras and Larsson (2020) recently documented its presence with Salix in the Scandinavian alpine zone. An alpine version of I. pararubens was also discovered and described as I. pararubens var. padjelantae. The study was published in Karstenia and can be found here.
Cripps and colleagues (2020) report ten species of nodulose-spored Inocybe from the Rocky Mountain alpine zone. Of these, four are described as new. These alpine species associate with Salix, Betula, and Dryas and have broad intercontinental geographic distributions perhaps influenced by dynamic migration patterns of their plant associates. Interestingly, seven of the ten species also occur below tree line in boreal or montane habitats. The study was published in Mycologia and can be found here.
Dovona et al. (2020) provide a contemporary description of a rare species described originally from Italy in 1905 – Inocybe similis. Based on molecular annotations of the type (good work!) and contemporary collections, surprisingly, this smooth-spored to subangular-spored species with affinities to I. sect. Splendentes is most closely related to I. flavobrunnescens, which the authors treat in I. sect. Marginatae, an artificial group of nodulose-spored species.
In this work by Linas Kudzma and myself, we described five new species of Inocybe s. lat. from eastern North America. The first of these – Inocybe carolinensis – is unusual because of the overall reddish coloration; it’s most closely related to I. tahquamenonensis in the I. sect. Inocybe but now known only from two locations at high elevations in the Southern Appalachians. Two species of I. sect. Lactiferae are described, the first of which is probably a hallucinogenic species, I. glaucescens, originally from New Jersey; and the second, I. dulciolens, is basically the east North American version of the European species I. fraudans. Several other unique species-level lineages across North America have been detected in sect. Lacterferae. These await further clarification. Inocybe friabilis -now Pseudosperma friabilis – is a new species in the Gracilissima clade, which forms the sister group to the Ps. rimosum complex. Lastly, I. vinaceobrunnea – now Inosperma vinaceobrunneum – was described to accommodate Hesler’s application of the name I. jurana to a species that it most closely related to the European In. rhodolium and In. adaequatum.
2017
Here’s a study I missed that was published in 2017 by Brugaletta and colleagues and describes a new species of Mallocybe (as Inocybe siciliana) from a riparian corridor under willows in Sicily.