Diario de cooperj

The genus Geastrum is easy to recognise, but like many fungal groups it is not so easy to identify species within the genus. Names like Geastrum triplex and G. saccatum are used indiscriminately everywhere. Most identifications adopting these names will be wrong.

My former colleague the late Ross Beever was interested in puffballs and their allies, and this current work builds on the collections and observations made by Ross (in the PDD national collection). I wanted to get this preliminary key ‘out there’ so it can be used, hopefully to recognise interesting collections for further work. Like many of my keys it was written for my own benefit, at least in this first instance. I do not know Geastrum very well (which is probably apparent) and the process of developing a key clarifies the concepts needed to separate species in my own mind. I will change this entry as more information becomes available, and I learn more. Like most genera you can only reliably identify species using a combination of macro and microscopic characters.

Good collections of Geastrum are needed for correct identification. By that I mean examples of the fruitbody from bulb stage to fully expanded, making a note of the colour of fresh material (any pinkish/red colouration of the flesh may fade). A fruitbody still attached to a piece of substrate is useful for characterising rhizomorphs. If material in the field looks old and knocked about then don’t bother attempting to identify it. A number of the key features are ephemeral and difficult to observe in weather-worn specimens. Read the terms in the short glossary to work out which features are key.

Geastrum was last revised in New Zealand by Cunningham in 1944 (Gasteromycetes of Australia and New Zealand). He recognised the following 7 species: G. pectinatum, G.plicatum, G. minus, G. limbatum, G. velutinum, G. triplex & G. floriforme. Like many authors he adopted names from the Northern Hemisphere. Although our species look similar to northern hemisphere species, and in many cases are closely related, the current sequence data suggest they are mostly localised in their distribution and probably indigenous. They are found in both natural and modified habitats, but there are some exceptions. G. floriforme might be introduced and is identical to the northern hemisphere species. In NZ this species favours dunes and alpine areas but isn't common in macrocarpa plantations, unlike many UK records where it is found under macrocarpa. On the other hand G. coronatum aff. and G. tenuipes, both like macrocarpa plantations, and G. velutinum aff. likes pine plantations. All of these species are not exclusively limited to these habitats and other Geastrum species occur in the same habitats. 

Because many of our species are near, but not the same as northern hemisphere species the names are followed by ‘aff.’ meaning ‘having and affinity with’ (phylogenetically and morphologically). Most are probably undescribed species. In some cases the sequence data indicate the names cover multiple species within a complex (G. velutinum aff. & G. saccatum aff.). The separate phylogenetic species within these complexes usually have no differentiating morphological characters (that I can find at the moment).

With reference to Cunningham's names his G. pectinatum/plicatum are treated here under the name G. tenuipes. It is possible that collections with a stalk, sulcate peristome and non-ridged apophysis may turn out to be different to collections posessing a ridged apophysis and a new name will be required for that species. Likewise G. minus is generally treated as a synonym of G. quadrifidum and here would key out under G. coronatum aff. It is likely that smaller versions of G. minus auct. NZ (of which there are a few collections) represent a good species which is likely to be G. austrominimum. More collections are needed to genetically characterise G. minus in NZ (in section quadrifida). There have been no confirmed collections since 1926. G. limbatum is now generally treated as a synonym of G. coronatum. A number of collections have been historically identified as G. triplex but on closer examination they represent G. lageniforme, G. australe or G. minutisporum aff. as well as G. triplex aff. The latter species is distinctly larger than any of the others. The presence of a residual fractured collar of pseudoparenchymatous material around the endoperidium has been shown not to be a unique and reliable feature of G. triplex (the so-called collared Geastrum), and I haven't used it as a character. G. triplex was described from Indonesia is a tropical species of south-east Asia. The name is used broadly and imprecisely elsewhere.

A number of names were added subsequent to Cunningham's 1944 treatment. A collection of G. drummondii was incorrectly identified. G. morganii was represented by a single collection which was used in an exhibit and not returned to PDD (duh!) but another collection related to G. morganii has been identified. G. fimbriatum has been recorded quite often but it is possible all these records refer to one of the species in our G. saccatum complex. G. fenestratum is generally regarded as a synonym of G. fornicatum and in New Zealand probably refers to G. setiferum aff.

Modern phylogenetic treatments by Zamora and colleagues (e.g. Zamora et al., 2015. Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota); Persoonia, 34: 130–165 & Zamora et al., 2014. Systematics of the genus Geastrum (Fungi: Basidiomycota) revisited; Taon, 63: 477–497) have opened up the black-box of Geastrum. They provided a phylogenetic framework to place species, and some new characters assist in identifying them. In some cases reliable identification of NZ species, even to the broad ‘species complexes’ treated here, is not easy. In particular the G. saccatum/G. lageniforme groups are difficult to separate without observation of the form of crystals attached to the rhizomorphs attached to the fruitbodies, which are often not collected. 

As mentioned above, a preliminary phylogenetic analysis of a number of NZ collections show they are closely related to some recently named southern hemisphere relatives and I have adopted those names, with the ‘aff.’ because they aren’t the same (to re-iterate what aff. means). Thus G. setiferum was named in 2002 from South America and G. minutisporum also from South America in 2016 and in both cases our species are phylogenetically close and do have different characters. They need new names. The numbers in brackets after a species name in the key refer to the Zamora classification (in the following table)and the position of representative sequences of NZ material in that classification.

It isn’t easy to incorporate photographs into this journal entry so those will have to wait.

NZ Trial Key to Geastrum

The Zamora et al classification of Geastrum, and NZ representatives

Glossary

Apophysis – collar/ring like swelling on stalk or junction of endoperidium and stalk.

Binding litter - The outer mycelial layer can bind litter or not, and this is a key feature. This doesn’t mean lumps of substrate that can be trapped by rays bent backwards. It means fine material firmly embedded within and stuck to the mycelial layer on underside of the rays. It is an indication the closed fruitbody was hypogeous (buried), and not sitting on the surface before it opened. However, soil encrusted layer may flake off, as a whole or in bits, and so I find this a difficult character to be absolutely certain about. The point of attachment of the fruitbody is often smooth, so don’t be fooled by that.

Endoperdium – the globe-like structure in the centre of the earth-star, specifically the skin of that structure. The texture of the surface is a useful feature. In some species it is obviously smooth/polished, but in most species it is superficially matt and under a lens either looks smooth or minutely hairy (tomentose), sometimes with aggregations of white crystals.

Exoperidium – The outer skin of the fruitbody can form three major layers. The outer mycelial layer (which can be simple or double), a middle fibrous later, and an inner pseudoparenchymatous (fleshy) layer. Outer and inner layers are evanescent, flaking off, sometimes lost entirely. Any pinkish colour to the fleshy layer is an important feature. The exoperidium splits into petal-like rays, which fold back.

Fornicate – rays arched downward and tips attached to remains of a separated exoperidial layer forming a basin in the soil. In some species the rays arch downward but without a separated cup layer.

Hygroscopic – You need to wet dried material to see the rays open out and then close up again as it dries. However, dried fruitbodies of hygroscopic species will be closed with the rays entirely covering the endoperidium so it isn’t visible. In non-hygroscopic fruitbodies the closed rays won’t cover over the endoperidium.

Mesoperidum – sometimes seen as a transient layer on surface of fleshy layer and surface of endoperidium, but usually dispersing rapidly.

Peristome – the circular zone around mouth, often different to rest of endoperidium (delineated) in either colour, texture, or raised/sunken. The area can be radially fibrillose, sulcate (ridged/pleated/folded) or smooth. Sometimes the term seems to have been used to refer just to the edge of the stoma rather than the circular zone surrounding the mouth.

Rhizomorphs - the rope-like hyphae connecting the fruitbody to the mycelium in the soil. They often have small (microscopic) crystals stuck to them, and the shape/form of these crystals is very useful fo separating groups. 

Saccate – an endoperidium sitting inside exoperidium at maturity, i.e. the globe is sitting in a bowl, not pushed up and exposed. Conversely the endoperidium looking like it is sitting on an upturned bowl (and then often with a stalk).

Stalk – between the endoperidium and exoperidium. Present or absent, and with or without an apophysis. Sometimes difficult to see without a vertical section.

Stoma – mouth where spores emerge. Edge of the stoma may be regular, fibrillose or ragged.

[2021 March - an updated phylogeny of NZ Strophariaceae sensu lato is loaded on this record https://inaturalist.nz/observations/48212034

One of my earlier posts was ‘The Stropharia, Hypholoma, Pholiota, Leratiomyces, Clavogaster series in NZ’. This post covers the remaining genera in the strophariaceae in a broad sense. The genera included here may or may not reside in the family according to different views. I have deliberately excluded Hebeloma, Psathyloma and Naucoria (but included Galerina).

Keys to genera for fungi are becoming increasingly difficult to construct because in recent years genera are named or redefined based on phylogenetic entities, sometimes without a set of unique morphological characters separating them, or at least not characters that are easily seen. In reality, only keys to species work well. For that reason this key to genera should be used in conjunction with the key to species in the earlier post.

I’ve used micro-features in the key because, even without phylogenetic splitting, naming genera (and certainly species) purely by macro-morphology is often a betting game with poor odds. If you want to name fungi accurately then get a microscope.

Some jargon is used. Google the terms if you don’t know them. Chrysocystidia are a very useful micro-character in this group. It refers to the reaction of the cystidia to KOH solution. Some species will show yellow staining content. The dye Patent Blue is even more effective but difficult to obtain. It is sometimes sold in tablet form as a ‘dental revealer’ but banned in NZ. Beware species that have yellow cystidia anyway, without the addition of KOH, e.g. in Gymnopilus.

The keys, as usual, are rough and need refining. I’ll work on them as I get time.

The Genera

Agrocybe/Cyclocybe & Crucispora

Agrocybe acericola, A. howeana & A. puiggarii are all considered doubtfully present, based on inadequate material.

Agrocybe praecox is very common on wood chips, especially in spring. An endemic fungus recorded rarely (A. olivacea) is suspiciously similar to a recently described wood chip species (A. rivulosa) spreading in the northern hemisphere. Agrocybe arvalis has recently been confirmed as present. It is a species associated with sclerotia and may be confused with Hypholoma tuberosa (or vice versa). Cylocybe parasitica is the new name of Agrocybe parasitica, which itself is very close (same as) the confused pair A. cylindracea/aegerita of the northern hemisphere. An earlier name for C. parasitica might be A. chaxingu, which was obscurely published. Cyclocybe species are phylogenetically very different to the rest of Agrocybe. Interestingly Crucispora naucorioides, described by Egon Horak from New Zealand appears to be related to Cyclocybe based on sequences of a 2nd collection of this rare fungus. Crucispora, as the name suggests, has cruciate spores, unlike all the species below. C. naucorioides is the only species in the genus and has only been collected once in NZ, until recently.

Agrocybe pediades is common in NZ on soil/sand in open areas, especially areas manured (usually with rabbit pellets). It is variable in appearance. Material confirmed by sequences as this species has a surface ixocutis and the hymeniform nature of the partially gelatinised subpellis difficult to discern or absent. For that reason it has frequently been misidentified in NZ as the microscopically similar Deconica subcoprophila, but that species has a darker cap, often radially striate, and above all else has dark brown/violaceous spores, without the typical cinnamon coloration of the gills/spores of Agrocybe. Sometimes it is also confused with Stropharia coronilla , but that has a distinct grooved ring on the stem.  

Deconica

Some species originally placed in Psilocybe have been historically recorded and not include here. D. physaloides is a nom. dub. and Guzman’s concept, according to Noordeloos, is a mixture of species including D. subviscida and D. xeroderma. Relatives of both are known in NZ and material labelled D. physaloides has not yet been examined by me. NZ material labelled Psilocybe subcoprophila by Johnston et al., 1995, is annulate, related to D. merdaria, and included in the key as D. merdaria cf. Other NZ material labelled D. subcoprophila is a misidentification of Agrocybe pediades. 

Psilocybe brunneoalbescens was described from Tasmania under a very broad concept of Psilocybe that incorporated species better placed in several other genera. From the description it might be assumed to fit within the modern concept of Deconica, perhaps like D. montana aff. However, NZ collections identified as P. bruneoalbescens by Gen Gates and David Ratkowsky, and Tasmanian material are similar, and NZ material has sequences which fall within Kueheneromyces. The recombination K. brunneoalbescens was created.  It has a darker brown stem and sweet taste compared to similar Deconica species. It does not have a large germ-pore which is characteristic of northern hemisphere Kuehneromyces spp.

Galerina

The moss/non-moss groups are phylogenetically different and will be split sometime, but Galerinas in general form multiple poorly resolved phylogenetic clades and more work is needed. The moss Galerinas are poorly known in NZ and names I’ve used will probably change. They are little more than sequence-informed guesses. We don't have the northern hemisphere species like G. hypnorum, G. calyptrata etc. These names are widely misused on iNat. We do have looka-like species. Amongst the wood inhabiting Galerina species with a ring we have G. patagonica (sensu Australasia) which is recorded from Australia and New Zealand. G. patagonica, as the name suggests, was described from South America but no sequences have been published to confirm the equivalence of the Australasian species assigned to this name. Australasian material tends to have a central nipple on the cap, but not always. There are 2 or 3 species within a complex with others known from Pacific Islands and the Subantarctic Islands. G. marginata cf. in the key is unrelated to the northern hemisphere species and may comprise two closely related species both found in NZ and Australia. One of these versions corresponds to Alec Wood's 2001 concept of G. rudericola with low spore ornamentation. I think it unlikley it is the same as the original version from North America.  The real G. marginata is deadly poisonous and contains the same toxins as the death-cap. G. patagonica and G. marginata cf. probably also contain the same toxins. 

Gymnopilus

Gymnopilus junonius is one of the most frequently reported fungi in New Zealand due to its preference for growing in large clusters on cut stumps of Pinus radiata. Our species is the genuine G. junonius first described from Europe with sequences the same as collections from Australia, South America, and Europe. Asian material, under the name G. spectabilis, is a different species and the North American taxa under this name have recently been revised (Thorn et al, 2020). G. ventricosus from the home of Pinus radiata in California is certainly not the species present in NZ. Published sequence data is not yet available for the reported US blueing variants of ‘G. junonius’, which no doubt represents a different species. Many of our P. radiata-associated fungi are European in origin perhaps because our original P. radiata came from Australian stock, itself based on European stock and not direct from California.

All Gymnopilus species grow on wood, and the G. purpuratus complex often on wood chips. G. purpuratus was originally described from Australia but unfortunately the name is invalidly published. The related G. luteofolius (luteifolius) has been recorded in both New Zealand and Australia, often associated with (pine) wood chips but sequence data indicate this North American name has been misapplied elsewhere. A supported ‘species complex’ clade contains sequenced material labelled G. purpuratus, G. suberis, G. terricola, G. subpurpuratus, G. purpureosquamulosus, G. norfolkensis, ‘G. luteofolius’ (=G. suberis), G. igniculus, G. aeruginosus and G. hispdus. Certainly, ITS sequence data show NZ material in a clade with all material named G. purpuratus from Australia, Cook Islands and Switzerland, and genuine North American G. luteifolius in a different clade outside the complex. The similar G. dilepis forms a well-supported sister-clade and incorporates collections with several names (including another version of G. purpuratus).  Stable morphological characters separating the species in this group are difficult to assess. G. dilepis, described from Sri Lanka, seems to have dominant orange/yellow colours and in dry conditions any purple is present only as a scabrous colouring towards the cap centre, although it is said to have more diffuse magnta/purple colours under wet conditions. It usually has a distinct ring zone/remnants on the stem. At least that is what sequence verified collections imply. G. dilepis is often described with a purple cap which would make it difficult to separate from G. purpuratus, but I believe many of those identifications are incorrect. G. dilepis and G. purpuratus are certainly phylogeneticaly distinct and the shape of the cheilocystidia quite different in the two species.  Separating the similar G. luteifolius is more problematic. One opinion separates G. luteofolius with pleurocystidia from G. purpuratus without. NZ material of G. purpuratus does not possess pleurocystidia supporting the use of this name.   We have one poor NZ collection of a faintly blueing Gymnopilus species as a weed in a mushroom farm and it has a sequence in this complex. For this taxon I have tentatively settled on the name G. megaspora (having started with G. cyanopalmicola). Sequence data indicate it is closely related to (same as) European collections of G. igniculus and (one group of) material labelled G. purpureosquamulosus. There is no doubt that a multi-gene analysis, more collections, and a better assessment of stable morphological characters are required to clarify this species complex globally. In my opinion it is likely (and preferable) that we recognise fewer species with infra-specific biogeographical variation, and with considerable phenotypic variability.

Several species can exhibit off-centre stems and the character is not diagnostic, e.g. for G.crociphyllus described as eccentric-stemmed.

Gymnopilus allochrous is an unpublished name and I have not found a description. The characters in the key come from material with a sequence identical to one named G. allochrous from Australia.

[Jan 2019] The key doesn't work well because the character 'perispore' creeps back in after being discarded. In addition we have the second collection of a species closely related to the Australian G. tyallus/eucalyptorum which needs including. The presence of G. flavus in NZ, growing with exotic grasses, requires confirmation with deposited and sequenced material, but is possible.

Psilocybe

It is with hesitation that I say anything about Psilocybe. For obvious reasons species within the genus receive considerable scrutiny around the world. My knowledge is limited by comparison. Nevertheless, as a taxonomist I do have some comments I’ll throw out there.

Psilocybe alutacea was described from Tasmania on cow dung and, as Allan Rockefeller has pointed out, was misidentified as P. fimetaria in New Zealand. Sequence data indicate the same species is present in Argentina misidentified as P. pelliculosa. P. alutacea is surely not native to New Zealand but the real origin remains unclear. Sequence data indicate we may have several poorly defined species in the P. cyanescens complex. I think the P. cyanescens complex needs more work, globally. The current sampling of gene loci (ITS, LSU, EF-a) and proposed key morphological separation characters do not adequately resolve the differently named entities within the clade. Certainly, data based on ITS alone is not definitive and obscured by the presence of multiple different ITS PCR dominant copies (haplotypes) within the same species. I do think there may be closely related real ‘species’ involved, rather than over-splitting of a single broad species (P. cyanescens) because the secotioid P. weraroa sits within the group and is clearly morphologically quite different. However, having said that, the evidence suggests a recent divergence of P. weraroa, indicating how simple the gene switch must be from open mushroom to closed pouch. The current phylogenetic data indicate there might be two very similar but different semi-secotioid species, in addition to secotioid variability in P. weraora . These variants are called informally P. 'subsecotioides' but that has not been published and so not attributable to any specific taxon. They require more work to disentangle them from P. weraroa and confirm as real species. Some species do seem to have been historically over-lumped by several workers. Chang et al, 1992 lumped P. tasmaniana, P. australiana and P. eucalypta into P. subaeruginosa, but the equivalence of dung inhabiting and wood inhabiting species does not have much support, and I agree with other observers on this point. I also need more convincing that the current concept of P. subaeruginosa is the same as Cleland’s original concept. It might be correct, or it might not. Cleland did not designate a type for this species but lists some collections he assumed to be the same species. The original description says it occurs in grass with one collection on dung, which differs considerably from the common (but not universal) modern interpretation of a wood chip or myrtaceae litter associated species. Guzman & Watling studied Cleland’s material, including Cleland 13251, which they called the ‘type’ and demonstrated the presence of unusual and characteristic brown pleuro- and cheilocystidia. That specimen is currently missing from Cleland's collections at Adelaide. A formal lectotype AD 5603 (=Cleland 13256) from Belair National Park Australia has since been designated (Chang & Mills 1992), but without published details of the habitat and substrate of this critical collection which would allow us to narrow down the accepted species concept. Their lectotype apparently does not have brown cystidia. That treatment has received subsequent criticism (e.g. Stamets 1996). No examined NZ material labelled P. subaeruginosa has brown cheilo or pleurocystidia and no sequenced NZ material is so far identical to Australian collections under that name. Likewise at least some recently reported collections of Psilocybe tasmaniana on wood chips in Australia and NZ are not likely to be Roy Watling’s Australian species described on dung “cap to 2cm and without umbo or papilla” and is probably undescribed. Psilocybe allenii was described relatively recently and occurs convincingly in New Zealand based on sequence data. It is considered closely related to P. azurescens and P. cyanescens. A recent sequence of the type of P. azurescens is different to other sequenced taxa under that name. All of which just shows the current application of names in this clade is a bit random and requires more work. All the wood chip species fall out at my couplet 5. Species differentiation based on spore size, cheilocystidia length and cap morphology (used in some keys) does not work for me. If somebody can give me a key that works for all the named species in the P. cyanescens group I would be happy to test it. I think more sequence sampling of the dung, soil and wood species in southern Australia and Tasmania, and epitypification against sequenced and well described material is needed to resolve the use of several names of closely related species.

A phylogenetic tree including New Zealand species may be found attached to this observation ... https://inaturalist.nz/observations/1462023

Stropharia

S.rugosoannulata can have pale forms. S. caerulea is the correct name for many NZ records identified as S. aeruginosa (with others being Cortinarius rotundisporus!). S. cyanea auct is the same as C. caerulea (Bolton’s original concept of Agaricus cyanea is unclear). S. 'Kennedy' may be the same as, or closely related to S. formosa from Tasmania.