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Strophariaceae sensu lato in NZ

[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

1

Pileipellis a trichoderm. Frb partialy secotioid. Sometimes in plant pots.

Tympanella galanthina

1’

Pileipellis hymeniform or a cutis. Frb secotioid or not.

2

2

Pileipellis hymeniform, often cracking on drying

3

2’

Pileipellis a cutis/trichoderm (but see also Agrocybe pediades)

4

3

Spores with germ pore

Agrocybe

3’

Spores without germ pore (Spores cruciate = Crucispora)

Cyclocybe

4

Stipe absent or laterally attached, small, Crepidotus-like

Deconica section Melanotus

4’

Stipe centrally attached (secotioid or not)

5

5

Spore print black/purplish

6

5’

Spore print brown

12

6

Pileus AND stipe glutinous in wet weather, on dung

Protostropharia semiglobata

6’

Pileus and stipe never glutinous. On dung or not

7

7

Clustered on wood or roots

Hypholoma

7’

Not clustered or if so then on soil

8

8

Well developed ring or ring zone

Stropharia

8’

No ring

9

9

Chrysocystidia absent

The secotioid C. virescens has chrysocystidia, but they are indistinct

10

9’

Chrysocystidia present. Frbs never blueing on bruising.

Hypholoma

10

Frbs red, secotioid or not

Leratiomyces

10’

Frbs brown, grey or blue, secotioid or not

11

11

Frbs usually blueing on bruising, secotioid or not

Psilocybe

11’

Secotioid or semi-secotioid, innately powder blue or yellowish

Clavogaster

12

Chrysocystida present (exceptions in other genera with Chrysocystidia:  Deconica baylissianum & D. novaezeandiae)

Pholiota & Protostropharia

12’

Chrysocystidia absent

13

13

Spores rough

14

13’

Spores smooth

15

14

Frb slender, often in mosses, sometimes on wood, spore print dull brown, taste mild.

Galerina pp.

14’

Frb more robust, on wood, spore print bright brown/orange, taste bitter

Gymnopilus

15

Pileus hygrophanous.

16

15’

Pileus not hygrophanous.

17

16

Growing on wood. Kuehneromyces brunneoalbescens has white flecks on a dark stem

Kuehneromyces brunneoalbescens & Galerina section Naucoriopsis (G. marginata group)

16’

Terrestrial in soil or grasses, or on dung.

Deconica

17

Pileus glabrous

Flammula ‘Matawai’

17’

Pileus with veil in slime, yellow/orange

Pholiota adiposa

 

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 (if at all). 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 rabbits 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.  

1

With ring. Spores with or without germ pore.

2

1’

Without ring. Spores germ pore.

4

2

On living trees. Spores with a pore

Cyclocybe parasitica

2’

Not on living trees. Spores with or without pore

3

3

On wood chips. Pileus pale brown. Often in spring. Spores with a pore

Agrocybe praecox

3’

On soil. Pileus dark brown. Spores without pore.

Cyclocybe erebia

4

Pileus with velvety texture and pilocystidia. On wood chips.

Agrocybe putamimum

4’

Pileus smooth, without pilocystidia, usually on soil (sometimes manured)

5

5

Pileus wrinkled, with olivaceous hue (A. rivulosa without ring)

Agrocybe olivacea

5’

Pileus smooth, yellow/orange

6

6

Not growing frm a sclerotium

Agrocybe pediades

6'

Growing from a buried sclerotium and pleurocystidia with apical fingers

Agrocybe arvalis

 

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.

1

Minute, laterally attached to dead herbaceous stems

2 (section Melanotus)

1’

Not laterally attached

4

2

On dead Astelia leaves, spores > 10um, limoniform

D. citrispora

2’

Not on Astelia, spores smaller and not limoniform

3

3

Spores > 8um on average, Pileus opaque.  On wood. Indigenous D. vorax, introduced D. horizontalis

D. vorax/horizontalis

3’

Spores < 8um on average, Pileus translucent striate, on grasses

D. philipsii

4

Orange/tan, secotioid, Thaxterogaster-like but in alpine herbfields.

D. baylissianum

4’

Not secotioid

5

5

On dung or bare soil, litter/thin twigs

6

5’

On rotting wood (not twigs), Tubaria-like

D. xeroderma

6

On dung (or manured soil)

7

6’

On bare soil (not manured), litter, thin twigs

9

7

Pileus remaining hemispherical, orange brown. Stipe without ring. 

8

8

Spores > 14um in length. [not Guzman's concept, which = D. merdicola]

D. argentina ss Johnston

8'

Spores < 13um in length [currently without sequenced NZ material for evaluation] D. coprophila cf.

7’

Pileus becoming flattened, dark brown, stipe with distinct ring or ring-zone.

D. merdaria cf.

9

On decaying grass stems and moss

D. subviscida aff.

9’

On soil, litter

10

10

Spores mainly rhomboidal

D. novaezelandiae

10’

Spores broadly ellipsoid (see also Kuehneromyces brunneoalbescens)

D. montana aff.

 

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. 

1

With mosses

2

1’

On wood, soil or litter

6

2

Spores calyptrate

3

2’

Spores not calyptrate

4

3

Cheilocystidia cylindrical, not swollen at apex

G. neocalyptrata

3’

Cheilocystidia swollen at the apex. Phylogenetically different but morphology difference elusive.

G. subcerina,G austrocalyptiformis SS NZ

4

Pileus < 1cm. Spores rough

5

4’

Pileus > 1.5cm. Spores smooth

G. pumila

5

Pileus with pilocystidia

G. atkinsoniana

5’

Pileus without pilocystidia

G. vittiformis

6

On soil or litter

7

6’

On wood

8

7

On soil, often in plant pots, with ring or ring-zone. Thick walled cystidia, 4-spored

G. nana

7’

In litter, lower stipe dark, covered in white fibrils. 2-spored

G badipes

8 Stipe eccentric G. nothofaginia & G. eccentrica

8'

Stipe central

9

9

Spores smooth. 4-spored. Cheilocystidia swollen apex. Lower stipe with white fibrils

G sideroides

9’

Spores rough. Deadly poisonous group.

10

10

At least some cheilocystidia bifurcate

G. patagonica

10’

Cheilocystidia swollen but none bifurcate

G. marginata cf.

 

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. 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. Separating 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. G. purpuratus was originally described from Australia but unfortunately the name is invalidly published.  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 considerable phenotypic variability until that work is done. 

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.

 

1

Spores with a perispore, sometimes indistinct 

2

1' Spores without a perispore 3

2

Stipe lateral.  Spores 4.5 - 5.0 µm x 3.5 - 4.0 µm.

G. pyrrhum

2’

Stipe central to excentric. Spores 7 - 8 x 5 - 6 µm. G. ferruginosus, (collections without an obvious perispore) may be a synonym. Pileus rusty colour (if orange see G. austropicreus)

G. mesosporus

3

Partial veil a membranous annulus, frbs mostly large, bright orange, usually on pine stumps.

G. junonius

3’

Partial veil a fine cortina or absent. Frbs smaller, bright or not, modified habitats or not

4

4

Gills rust spotted at maturity. Veil present when young on lower stem. Gills pale to start. Spores 7-9 x 4-5

G. penetrans

4’

Gills not rust spotted at maturity. Veil absent. Gills brighter coloured from start. 

5

5

Pileus with blue, purple or reddish, rust brown colours.

6

5’

Pileus with yellow, golden or rusty orange colours. In some collections of G. dilepis the purple can be absent, so see also couplet 7

9

6

Pileus reddish rust brown, margin often lobed, fibrillose, spores strongly ornamented. Stipe sometimes eccentric. Natural habitats. G. mesosporus is a potential synonym.

G. ferruginosus

6’

Pileus or stipe with purple or cyanescent, sometimes fading, margin not lobed, spores more finely ornamented.

7 (purpuratus complex)

7

Pileus and stipe weakly cyanescent when bruised, fading.  Spores 8-11 µm x 6-8 µm. 

G. megaspora

7’

Pileus and stipe not cyanescent. Blue/green/purple colours innate. Spores smaller, 6-8.5 µm x 4-6 µm

8

8

Pileus usually brown, not scabrous. Purple colour persistent often dominant. Spores 7.5-8.7 µm x 4.8-5.7 µm

G. purpuratus

8’

Pileus usually orange, purple colours, if present at all, as scabrous cap centre. Ring remnant often present. . Spores. 6.3-7.2 µm x4.5-5.1 µm

G. dilepis

9

Spores 6.0 - 8.0 µm x 4.8 - 5.6 µm (and with an indistinct perispore). Pileus smooth.

G. austropicreus

9’

Spores without a perisporium

10

10

Spores 4.8 - 7.2 µm x 3.2 - 4.8 µm.

G. crociphyllus

10’

Spores mostly larger to 9 µm x 6um

11

11

Spore ornamentation very fine. Cheilocystidia abundant. Pileus yellowish brown, often with white flecks on perimeter. Stipe often with white silky coat.

G. allantopus

11’

Spore ornamentation medium. Cheilocystidia sparse. Colour bright orange/yellow, reminiscent of G. jononius.

G. allochrous ined.

 

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 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. They also did not mention the presence of brown cystidia for their lectotype. That treatment has received subsequent criticism (e.g. Stamets 1996). No examined NZ material labelled P. subaeruginosa has brown cheilo or pleurocystidia and no sequences 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 considered closely related to P. azurescens and P. cyanescens. Recent sequences obtained from the type collection of P. azurescens indicates they are probably the same species and so P. allenii is perhaps a later synonym of P. azurescens. Other sequenced material labelled P. azurescens are misidentifications (of multiple species). All of which just shows the current application of names in this 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

 

1

On dung, stem with veilar remnants, sometimes bluing, similar to P. fimetaria. Deconica argentina/coprophila look may look a bit similar but without blueing. If cap with central nipple see P. angulospora.

P. alutacea

1’

Not on dung

2

2

Frb secotioid

P. weraroa

2’

Frb not secotioid

3

3

On soil

4

3’

On wood/wood chips

5

4

12 µm x 8um on average. Pileus 7-10 mm, with sharp umbo. In grassland.

P. semilanceata

4’

8 µm x 5 µm on average. Pileus 15-55mm, without sharp umbo. Phylogenetically and morphologically close (same as?) P. zapotecorum. Occasionally on wood.

P. aucklandiae

5 Spore length < 10um (9-9.5 x 6-7 µm on average) 6
5' Spore length > 10um. On wood chips in modified habitats (and probably all introduced). A species complex (exc. P. tasmaniana) with currently unreliable separating characters and species concepts. P. cyanescens, P. allenii, P. subaeruginosa, P. tasmaniana
6 On rotting wood in natural habitats. Cap without central papilla. Spores not angular in outline P. makarorae
6' On wood chips, greenhouse, plant-pots. Spores angular in outline. P. angulospora

 

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.

1

Pileus wine red to brown, smooth, frb large, often over 12cm

S. rugosoannulata

1’

Pileus other colours, smaller, under 8cm

2

2

Pileus with blue/green colours

S. caerulea

2’

Pileus yellowish or brown

3

3

Pileus yellowish, smooth, in grass, modified habitats. Stem with grooved ring.

S. coronilla

3’

Pileus brown, velutinate, unmodified habitats, stipe with rhizoids

S. ‘Kennedys Bush’

Posted by cooperj cooperj, October 06, 2016 03:33

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