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Coming soon! Changes affecting the genus Inocybe $ %D "C

t may come as news to some readers that in cally the differences are much less apparent (one accordance with developments proposed in a of the reasons why this genus has always been Ipaper published in Mycologia last December considered such a challenge to identify). Basically (Matheny et al., 2019) the genus Inocybe is to be what is going to happen is that these three main revised and divided up into several separate subgenera are now to be elevated to genus level genera. This trait in mycology is one with which conveniently retaining their subgenera epithets we are becoming increasingly familiar and, as the new genus name (though there is a compli- whether we like it or not, is here to stay. Each cation involved – read on). Thus, for example, time a reappraisal such as this occurs the field Inocybe agardhii becomes Mallocybe agardhii, mycology community at large is required to come Inocybe bongardii becomes Inosperma bongardii to terms with a whole new suite of names, not and Inocybe napipes remains Inocybe napipes. only genera names as here in the case of Inocybe That’s the good news. Now for the inevitable but also, though thankfully to a much lesser ‘buts’, the first of which luckily affects us in the degree, species names as well. (It is a relief to UK very little, though there follow some issues many that the common English names remain which are likely to cause some groans, possible unchanged when this happens!) The reason for confusion and inconvenience. this general state of affairs has been well documented in recent years and I will not dwell Seven new genera now in Inocybaceae here upon the mycological transformation worldwide brought about by the inexorable march of molecu- This large and ecologically important mycor- lar development. My aim is to explain to readers rhizal genus comprising over 1000 known species what is happening to Inocybe in simple terms in worldwide is in fact to be split into seven differ- order to smooth the passage for the amateur ent genera – we have been expecting a possible community with regard to naming and continu- split into three genera for some time, but seven?! ing to record. How soon this new system is to be Well, maybe we should be grateful that this is far adopted in the UK will become clear in the fewer than the mind-boggling number of new fullness of time, but rest assured it will happen. genera into which Boletus has been split! As This is an attempt, therefore, to make that already mentioned, three of the new genus names transition easier to grasp when it does. are elevated from previous subgenera and are thus thankfully familiar to those who study the Some familiar subgenus names retained genus; four others, however, are new. Here are Unlike the treatment of genera such as Coprinus, the seven new genera followed by a resumé of Collybia and more recently Boletus where very how the new system will work with a discussion few species ended up being retained within the of each genus in turn: Auritella, Inocybe, original genus, fortunately with Inocybe the bulk Inosperma, Mallocybe, Nothocybe, Pseudosperma, of species will in fact retain that genus name. To Tubariomyces. date the majority of recognised dichotomous keys for the genus have for convenience generally split Inocybe – the genus for all those species the species into three clearly defined and recog- with metuloid cystidia (Fig. 1) nisable subgenera (sometimes called sections or So for us in the UK once our previous subgenera supersections but subsequently below referred to of Mallocybe and Inosperma become genera in as subgenera), these being Mallocybe, Inosperma their own right, what remains will form the new and Inocybe, each with their own set of micro- slimmed-down genus Inocybe (i.e. our previous scopically distinct characters though macroscopi- Inocybe Subgenus Inocybe) comprising those C$ /!$#$ " doi:10.1016/j.fl dmyc.2020.07.007 102544 H"%%ILK species which have metuloid pleuro- and cheilo- know the genus that there are to be three genera cystidia, spores both smooth (usually amygdali- to replace the previous two subgenera of form) and nodulose, stems both cortinate and Mallocybe and Inosperma. I will discuss marginate, also both cylindrical and bulbous (Fig. Mallocybe first, leaving the more contentious 1). This will still be by far the largest and most issue of Inosperma and Pseudosperma till last. widespread genus within the Inocybaceae having an estimated 850 species with representatives Mallocybe (Fig. 2) occurring in all continents apart from Antarctica. The new genus Mallocybe will include all species At a rough estimate 120 of these are British previously in Inocybe subgenus Mallocybe though, as elsewhere in the world, this number (though for the moment with one notable excep- will continue to grow - in fact several new British tion – more on this below). It may come as a discoveries are already in the pipeline. surprise that although Mallocybe is very poorly represented in the UK (currently with only four The four new genera with unfamiliar species and those generally much less familiar to names: Auritella, Nothocybe, Pseudosperma us than Inocybe or Inosperma), this is in fact a and Tubariomyces. sizeable genus with an estimated 55 species All other species within Inocybaceae worldwide, having representatives occurring in Africa, Asia, i.e. those without metuloid cystidia, are to be Australia, Europe, New Zealand and N. America. split into six genera but for us in the UK this is However, even well into the DNA age as we are not as fearful as it might at first seem. Two of the now, experienced workers are still finding this six names, Mallocybe and Inosperma, I’ve already suite of species extremely problematic and tricky mentioned above; of the remaining four (those to identify. In the UK we have many unnamed with unfamiliar names) only one will we need to collections which we hope in time will add new become acquainted with: three of these species to our meagre list. It may turn out that in (Auritella, Nothocybe and Tubariomyces) do not some cases identification to species level may occur in the UK nor are likely to in the foresee- only be possible with molecular confirmation and able future, so we can safely eliminate them from that we may have to be content with assigning the new British Inocybaceae scene. some collections to a complex of species. • Auritella occurs only in sub-Saharan Africa, The exception I mentioned above: Our only India and Australia with fifteen species known relatively common member of this group is and is placed somewhere near Mallocybe in the Inocybe dulcamara (known to be a species phylogenetic tree. complex) but this recent paper lists this species • Nothocybe occurs only in India with one under ‘Taxa of uncertain position’ owing to an species so far described and is placed somewhere apparent discrepancy over our modern concept between Inocybe and Pseudosperma in the phylo- compared to Persoon’s original description (1801) genetic tree which (the paper suggests) is ‘questionably a • Tubariomyces occurs mainly in sub-Saharan Mallocybe as widely interpreted but not in the Africa, Australia and (to a much lesser extent) sense of the original author’. So for now this Spain with six species known and is placed species remains in limbo without an official new somewhere near Mallocybe in the phylogenetic name. Where does that leave us recorders? tree. Presumably we continue to record the species This leaves Pseudosperma, the fourth new under the name Inocybe dulcamara until this genus and one which does affect us in the UK problem can be satisfactorily resolved. It would and is likely to cause some confusion. It is proba- seem to me that this taxon (whatever it becomes) bly best explained in combination with will clearly take its place within the genus Inosperma and Mallocybe. Mallocybe though to satisfy the rules of nomen- clature it may need to be given a different species Inosperma, Mallocybe and Pseudosperma – name. the three new genera for those Inocybaceae species in the UK and N. Europe with Inosperma and Pseudosperma (the flies in ‘simple’ cystidia (no metuloids). the ointment?) (Figs 3 & 4) By now it will have dawned on those of you who If asked to name an example of an Inocybe

103545 H"%%ILK from subgenus Inosperma (as was), most mycolo- new genera Inosperma and Pseudosperma will gists familiar with this genus would probably have changed endings if previously they ended in suggest Inocybe rimosa, one of our commonest the letter ‘a’. My apologies to those of you for UK species. The introduction of the new genus whom this elucidation is old hat but suspect there Pseudosperma into the melting pot will, however, may be many who like me have been previously a see I. rimosa (as was) and all closely related bit bemused by such things. species moving to adopt this new genus name, leaving the remainder of what formed subgenus Now back to Inosperma and Pseudosperma Inosperma as the new genus Inosperma. Here, The new genus Inosperma will have about 55 then, is the spot within the new treatment of species with representatives occurring in Africa, Inocybaceae where UK mycologists will find some Australasia, Asia, Europe and N. America with confusion and doubt as to what the new name of roughly 10 of these occurring in the UK. The new their species might be. The situation is further genus Pseudosperma is slightly larger with about slightly compounded by the fact that all those 70 species with representatives occurring in the taxa which will now transfer from Inocybe to one same five continents plus northern S. America of these two new genera but which have species with roughly 10 of these occurring here. names that previously ended in the letter ‘a’ will When I realised that Inosperma would now no now need to change to end in ‘um’. longer include I. rimosa, I found myself wonder- ing where I. maculata and I. cookei (both Latin and Greek derived endings examples of species complexes and traditionally I’m going to digress for a moment here in an included within the Rimosae) would now reside. attempt to explain why these seemingly inconve- Would they also be transferred to Pseudosperma nient and irksome changes in endings need to with the Rimosae or not? So to clarify possible happen and am grateful to Alick Henrici for confusion over the new placement of these and putting me straight over the niceties involved. other species, herewith is the complete list of UK This all comes about because Latin genus names species from Subgenus Inosperma as was but derived from the Greek and ending in ‘cybe’—a with their new names. It might be a useful feminine ending meaning head/cap—require to exercise to annotate whatever keys and reference be followed by a Latin species name with a books you like to use accordingly in readiness for feminine ending also, in this case ‘a’. Thus this coming change. Inocybe maculata. However, genus names derived from the Greek but ending in ‘ma’ are British species now in the new genus neuter, not feminine, and therefore require to be Inosperma with changed endings in bold: followed by a species name with a Latin neuter Inosperma adaequatum, I. bongardii, I. calamis- ending, in this case ‘um’. It follows therefore that tratum, I. cervicolor, I. cookei, I. erubescens, I. both new genera Inosperma and Pseudosperma kuthanii (= cookei var. kuthanii), I. maculatum, with Greek neuter endings will require species I. pisciodorum (= bongardii var. pisciodorum), names which follow suit, i.e. their previous Latin I. quietiodor (Fig. 3) feminine ‘a’ endings will change to the neuter ‘um’. Thus Inocybe calamistrata becomes British species now in the new genus Inosperma calamistratum and Inocybe rimosa Pseudosperma, also with changed endings becomes Pseudosperma rimosum. Confusing or in bold: what?! The confusion surely lies in the fact that Pseudosperma arenicolum, P. curreyi, P. flavel- the Latin ending ‘a’ is feminine but the Greek lum, P. mimicum, P. obsoletum, P. perlatum, similar ending ‘ma’ is neuter. However, having P. rimosum, P. spurium, P. squamatum, taken this on board I now follow the reasoning P. umbrinellum, P. xanthocephalum (Fig. 4). behind such familiar names as Cystoderma My caution above in quoting only estimated amianthinum and Scleroderma citrinum – names numbers of UK species in the four new genera is which have worried me for years! In case I’ve lost down to possible discrepancies over updating you along the way, herewith a summary: species checklists and dictionaries used for recording in the new genera Inocybe and Mallocybe will with regard to newly discovered UK species have unchanged endings whereas species in the which may be awaiting official description. Also

10454- H"%%ILK in some cases acceptance of varieties as full complexity, particularly one which of necessity species or collections awaiting DNA testing etc. depends so heavily upon microscopic differentia- Keeping up to date with this genus in which new tion between species, it is essential to make use of species are constantly being described is every available character. Splitting up the challenging to say the least. species into artificial subkeys using spore shape and the distribution of caulocystidia will, I A sting in the tail – caulocystidia suspect, long continue. Time will tell. That, I think, covers the nomenclatural changes to the genus. There is, however, one more inter- Reservations esting development in this paper which I’d like to There is no doubt that Matheny et al. (2019) is an bring to your attention concerning the new-look admirable piece of work, full of detail and expla- genus Inocybe. Most keys to the genus in general nation and well worth a read if you’d like more usage today have employed two convenient tools information on the whys and wherefores etc. The with which to split subgenus Inocybe (as was) authors are to be congratulated though I must into more manageable chunks in the form of four admit to finding it technically challenging to (or more) different ‘subkeys’ according to whether follow in some respects and do have a couple of (a) spores are smooth or nodulose and (b) fruit- reservations: bodies are cortinate or not. We’ve known for some • Genera versus subgenera? time now that though it might seem logical on the Though it is clearly a step forward to include all surface to place the nodulose-spored species in a worldwide related genera under the umbrella of separate genus from the smooth-spored—indeed Inocybaceae, thus furthering our knowledge and there have been attempts in the past to do this— understanding, one could argue that this might it is now scientifically proven not to be justified possibly have been achieved just as well by because monophyletic groupings apparently do retaining the seven new genera at subgeneric not correspond to these differences in spore level, still under the umbrella of genus Inocybe, shape. Furthermore it now transpires that thus keeping names with which we are familiar neither do the monophyletic groupings corre- and removing the need to change species endings. spond to whether species are ‘cortinate’ or Furthermore, the new genus names now have no ‘marginate’, i.e. to the distribution of caulocys- obvious verbal link to the word Inocybe (unlike in tidia down the stem, a feature previously consid- the new genera Coprinopsis and Coprinellus for ered possibly even more significant than spore example, where the words resemble Coprinus). shape within the genus. The paper proposes that This might result in confusion, even misunder- in this genus evolutionary ‘development of basid- standing, especially when, for instance, reference iomes is highly varied’ and that the science does books place genera alphabetically. I surmise that, not justify the official separation of species into though it might not have been so accurate scien- Cortinatae and Marginatae as has been done in tifically, if the use of subgenera had been the past and consequently this method ‘should be retained within the original genus Inocybe some abandoned’. of the above difficulties might well have been Initially it comes as a bit of a shock to those of avoided. us ‘brought up’ to respect the vital importance of • Taxonomic Key to Genera of Inocybaceae such features to be advised now to abandon them! The paper provides a basic key to the new genera It should be mentioned, however, that this is not which I suspect, however, will not be found very a new theory. As long ago as 1980 it was one user-friendly in the UK. Looking for help in supported by Alessio, whose much revered getting to grips with Inosperma versus monograph did not recognise the occurrence of Pseudosperma—an issue of particular relevance caulocystidia as significant or as a means of to British users—I found this key unhelpful and identification in the genus. Personally, reading confusing. Earlier above I touched on the fact between the lines, I do not consider that this that the new genus Inosperma now contains all should herald the end of workers using this species, if you like, ‘left over’ after the Rimosae method as an identification aid despite phyloge- have been transferred to the genus netic reasoning suggesting the contrary. When Pseudosperma. To my mind this creates a diffi- devising a key to a genus of this size and culty in that amongst those remaining are two

10554( H"%%ILK morphologically very different suites of species, falling on the second syllable, in bold). I’ve contin- namely the Cervicolores clade and the Maculata ued with this though many others (no doubt more clade, together with a couple of other British traditionally) favour I-know-sigh-be, in line with species which don’t fit particularly comfortably Con-know-sigh-be (Conocybe) and Cly-toe-sigh- with either of these. If you know the genus well, a be (Clitocybe). I’ve also inconsistently used Inn- glance at the new British Inosperma species in oh-sperma and Mall-ossibee. It seems to me the list above will show you what I mean: yes, now, however, that at least some link between they all lack pleurocystidia and have simple thin- the original genus name and the new generic walled cheilocystidia, but then so do all the names would be retained if a more consistent Rimosae. Furthermore the species within this phonetic pattern of pronunciation were to be new genus share little else that I can ascertain adopted. I suggest, therefore, that the following with which to separate them off ‘en block’ from might help to achieve this: the Rimosae. This is born out by the fact that it has been necessary in the key provided to use I-know-sigh-be; I-know-sperma; Mall-oh-sigh- first one set of characters to split off the Maculata be and Sued-oh-sperma. clade, then in a later couplet another set for the Cervicolores - not ideal. Conclusion The phylogeny may point towards splitting off The complete list of proposed new combinations this somewhat miscellaneous group of species as and names with which the paper concludes is a has been done here, but there is much to be said welcome resource, not least because in quite a in favour of the previous system where the few cases it confirms taxa previously considered Maculata clade resides in close proximity to the by some as varieties; are now elevated to full Rimosae with which it has clear morphological species as a result of DNA research. Over all I links. As an amateur I would respectfully suggest support the forthcoming changes despite some that this appears to be a case of phylogeny unfor- reservations discussed above together with the tunately conflicting with morphology resulting in inconvenience that all relevant literature will an unsatisfactory situation for (a) those strug- now be rendered out of date. When a single genus gling to devise a workable key and (b) those is as large as the Inocybe but has within it such struggling to get to grips with following that key. clearly defined sections to which we have been referring for years, it makes sense to upgrade Thoughts on pronunciation those names to give them more significance. No Normally I’m not one to have strong views on doubt it won’t be long before other large genera our pronunciation of Latin-based fungus names, are given similar treatment. being of the opinion that it doesn’t matter that much as long as the listener knows to which Thanks species one is referring. The much quoted and I am indebted to Fernando Esteve-Raventós, a loved Ella Fitzgerald/Louis Armstrong classic co-author of this new vision of Inocybe, who has ‘You say tomAto and I say tomarto’ springs to patiently answered my many queries not only in mind here (but not its concluding line ‘Let’s call relation to this paper but also on many other the whole thing off’!). As I touched on under Inocybe topics over the past decade. I also thank ‘Genera versus subgenera’ above, the introduc- Alick Henrici for helpful suggestions and advice. tion of the new genus names will in effect remove their verbal link with the word Inocybe (just as Leccinum and Suillus now have no verbal link with Boletus) and this is surely a disadvantage References especially for those starting out in mycology. I Alessio C.L. (1980): Inocybe. Iconographia suggest therefore that some conformity over Mycologica Vol. XX1X Supplementum 111 pronunciation might go some small way towards Tridentini. Matheny P. B., Hobbs, A. M., Esteve- retaining that link. Raventós, F. (2019). Genera of Inocybaceae: I was initially introduced to Inocybe (back in New skin for the old ceremony. Mycologia 1993) pronounced phonetically as I-nossibee DOI: 10.1080/00275514.2019. 166906. (similar in rhythm to Rhinoceros with the stress

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