Notes: Instantly ID-ed as the not-Breckonii A. breckonii by Darv.
Habitat: in soil under Bishop Pine (Pinus muricata). Young specimens partially submerged in duff.
Habit: solitary to scattered.
Pileus 3.5-8.5cm, rounded becoming convex to broadly convex to plane or irregular in maturity, surface slightly viscid when moist, creamy to pale yellow, golden yellow, with a darker, pale greyish- to pale purplish-brown disc at maturity, margin decurved to slightly incurved when young, plane to uplifted in age, striations present (1mm apart, up to 8mm in length) in drier and/or older specimens. context white except for a pale, bright yellow layer immediately beneath the pileus cuticle, more concentrated beneath the disc, universal and partial veil remnants present, UV remnants splitting into a calyptra, patches and/or warts, PV remnants forming an appendiculate margin when young, composed of short, fine, cottony tufts of tissue.
Lamellae adnate to adnexed, then free, leaving ephemeral decurrent lines on stipe apex, close, white .5cm broad, lamellulae present in two series. margin considerably darker (deep golden yellow) upon drying.
Stipe 5-14cm x 1-2cm (in center), enlarged at apex (2.5-3cm) with a tapering, somewhat rooting basal bulb (2.5-3.5cm). tissue discoloring slightly with age and/or handling. outer stipe context with pale, bright yellow tints. interior stuffed/pithy in youth and hollow in age.
Annulus inferior, thin, membranous, evanescent, concolorous with stipe and volva
Volva collar-like and distinctly limbate with scaly/fibrillose zones above the rim that fade in age.
Pileipellis: a partially gelatinized cutis of long, slender, radially appressed, infrequently branched, septate hyphae of fluctuating thicknesses with rounded apices. clamps not observed.
Basidia: clavate, four-sterigmate, unclamped, up to up to 56.5×12.5*
Spores subglobose to broadly elipsoid, half smooth, half distinctly pockmarked (crassospores), inamlyoid w/ pronounced apiculus.
8-11×6-9μm (x=9.4×7.5μm, Q= 1-1.571μm, Qm=1.251μm, m=30, s=1)
9×9 ; 1
9×8 ; 1.125
10×8 ; 1.25
9×8 ; 1.125
10×8 ; 1.25
9×6 ; 1.5
10.5×8 ; 1.313
10×9 ; 1.111
10×8 ; 1.25
9×7 ; 1.286
10×7 ; 1.429
8×7 ; 1.143
8×7 ; 1.143
9×7 ; 1.286
10×8 ; 1.25
10.5×7 ; 1.5
9×7 ; 1.286
9×8 ; 1.125
9×7 ; 1.286
9×6 ; 1.5
9×8 ; 1.125
9×8 ; 1.125
9×8 ; 1.125
10×8 ; 1.25
9×7 ; 1.286
11×7 ; 1.571
10.5×9 ; 1.1667
9×7 ; 1.286
9×7 ; 1.286
9×7 ; 1.286
*Basidia being unusually difficult to locate, these measurements represent the upper limit of basidioles observed in several crush mount of gills. The max length is likely a few microns short of what it would have been measuring similarly-sized basidia with fully-formed sterigmata.
|I’d Call It That||3.0||0.00||0|
sum(score * weight) /
(total weight + 1)
|I’d Call It That||3.0||4.94||1||(amanitarita)|
sum(score * weight) /
(total weight + 1)
material sent. packaged and labeled as instructed. the entire collection’s in there. Dennis put it all into a heat-sealed bag before I had the chance to suggest it be split in half.
in the image in question.
The subhymenium expands a lot during the development of the mature fruiting body. It adds cells and individual cells of the subhymenium can produce more than one basidiole/-um. The latter is what I see going on in the picture indicated by Danny.
Mike Wood (I think) has something on his site about a clamp looking like the handle of a teacup. I think that’s a helpful statement. Now add to that that the teacup handle crosses the boundary between two cells. But the clamp doesn’t sit outside the boundaries of the two cells like a teacup handle hobo straddling two cars on a train (I’m thinking of segments of a hypha here that I imagine running horizontally across your mental image like a train)…part of the clamp is like a wedge driving down between two segments of the hypha. The wedge goes about halfway across the septum dividing the two cells. If you go into 2-D now, imagine the two sides of the wedge and the part of the “regular septum” below the wedge…and you see a “Y”. OK, now go into low quality animation; and rotate our image. The Y is lying on its side. The hyphal segment above the clamp morphs (poof!) into a basidium. The hyphal segment below the clamp morphs into a cell of the subhymenium. Now we are returning to the topic at hand. Stay in 2-D….
The base of the basidium is now fixed. The basidium is very unlikely to change into anything else. Check it out. The base of the basidium has a distinct bend in it…an obtuse angle. The bottom “leg” of the Y is part of the basidium’s base, and one of the “arms” of the Y is also part of the basidium’s base. They make a very open “V.” Go back to 3-D. The base of the basidium is formed of two planes that meet at an obtuse angle. This is going to be true in a clamped basidium whether or not the clamp proliferates. If the clamp turns into another basidium-supporting cell (could happen), then the base of the basidium is going to be the way it was before the clamp became a shape shifter.
Bottom line: You’re not so much looking for a second basidiole as you are looking for basidia with bases that are formed by the two planes meeting at an obtuse angle.
Look for the 3-D “V.” Remember. It’s not a sharp “V.” It’s obtuse.
Final thought. If a species is a member of the pantherinoid group, it is very unlikely you will see a proliferated clamp, because there weren’t any clamps to turn into other things. Clamps are very rare in gemmatoid taxa, but you might run into one. At any rate, if we have been right about this guy so far (and taking into account the information provided below that apparently genetic information also suggests this taxon is gemmatoid) then the likelihood of seeing clamps or the ghosts of clamps is pretty low.
not so hard once you know exactly what you are looking for/at.
just above the reticle. there’s a significantly smaller basidiole-like protrusion beside (and parallel to) the basidiole. they share a common base. proliferated clamp? if so, there were tons of these.
concise, clear, and directly relates to this obsie.
why google when a user knows a working definition that fits this discussion?
why not share and enlighten?
even my “Dictionary of the Fungi” doesn’t have a definition, way mo’ bettah than google!
a term that I had never heard before, either.
Perhaps Kiki will enlighten us?
The point of the clamps is that with Amanita muscaria you occasionally see clamps at the base of the basidia. In my estimate they are between 3-15% of the cases, depending on the specimen. This is a pretty good feature to tell what is closer to A. muscaria vs. A. gemmata/pantherina.
The clamps of Amanita are not particularly easy to see and some patient searching might be necessary. One has to do a diligent work before declaring that there are no clamps.
At the same time the presence absence of clamp collections and the frequency of that occurring is a topic where the mycologists have not been particularly good to develop solid defintions.
In many genera and infrageneric clades the clamps present a useful tool for taxonomic analysis.
There are also many microscopic features, which are way overrated by taxonomists of the golden era of microscopy (1920-1990). One has to develop a balanced approach of what is a good character and what not and how they can be used in combination.
Hmmmmm…..did you rule out proliferated clamps???
so the assignment is complete, but the observations will likely continue to grow whenever I’m afforded the time to return to them. I’ve got a list of characteristics that need reviewing for one or more of the six (hypodermia, subhymenia, more stipe characters, etc.). the question is when they’ll see the light of day again, as three months of Bolivian collections are about to jump to the front of the queue.
i’d like to point out something that so far seems to have gone unnoticed. Dimitar’s thread at mushroom talk mentions that one of the key characters of the not-breckonii A. breckonii is clamped basidia. that character was absent in this material, as the notes and photos illustrate, unless somewhere in those images are clamps in disguise.
BTW Danny, nice job in documenting your find. I see that you used Rod’s method of spore-graphing, and borrowed Dimitar’s penchant for Congo Red stains!
All that was lacking in your micrographs was veil tissue; the membranous UV is particularly interesting in this species, in comparison with the more friable veil of a typical gemmatoid species.
Excellent observation, I’m in awe of the work you’ve done thus far and how you’ve learned. I love the observations here that are detailed and carry a large discussion along.
It continues to blow me away how many jewels we have here, brainy jewels..
Ok, Debbie, Rod, I didn’t realize that you own all discourse on genus Amanita and moderate this site too. I thought we were having a very pleasant conversation here, appropriately positioned below a very nice observation. I just wanted to open up on some of the challenging aspects of molecular work with genus Amanita, which manifest themselves most acutely when working with this particular group. And then I see this unprovoked assault on my persona… Well, I do believe that I know a few things that you don’t and the audience would benefit from what I have to say. But if you find my contribution unworthy of this site then I shall withdraw and continue at another forum, most likely MushroomTalk.
Quietly, with love, so long.
The full type study of Amanita breckonii is on-line, with additional data from a Washington collection made by Jan Lindgren.
It has been available for some time at
We were having a conversation on this observation. Now we have a loud voice with name calling, disdain, and bullying.
I like the Amanita augusta name, that sounds really good. Its also simple and concise. I think I also like Amanita kashaya, but I’ll have to think about it a bit more before I am sure. How would that be pronounced?
Ok, the history from where I entered in the picture is approximately correct – many observations of this species were called A. breckonii and at every iteration Rod would in a low humming voice dispute that id on account of micro evidence. But our local heroes in California continued to use that name over and over. One day I grew tired of that senseless repetition and being a man of action went straight to the SFSU herbarium to check on the A. breckonii type collection. Indeed, the spores Q-ratio was as reported by Rod who had a 9 page type study included. So this species here is not the true A. breckonii. The beautiful observation here gives exactly my interpretation of Q-ratio for the species spores: ~1.25. And the A. breckionii was much, much higher ~1.7. Imagine that the confusion started with one irresponsibly selected photograph of this species that was labeled A. breckonii.
But what is A. breckonii? I agree that it appears to be a muscarioid taxa based on the evidence… Unless… ..and here I have a hypothesis that I will keep for myself for now. And while being orthogonal to our work on the California gemmatoid Amanitas we tried very hard to sequence the A. breckonii type. So far we have been unable to accomplish that and it has caused me some aggravation. But we plan to employ more powerful means on the next try and even offer an authorship to whoever succeeds.
So, the long story short – we have made significant progress in the study of this group via modern means and some interesting results will come. We are not there yet, but at some point, not far distant from now we will have to name this species, unless someone beats us to it. So, if it is us doing it, here is your chance to suggest names. I will listen very carefully and take all considerations – let’s skip people’s names on this try, can we, unless very major? My thought was Amanita kashaya – to honor the indigenous tribe from the local area. The name has a nice feminine ring to it too that I am fond of… But I am ready to listen if the majority of people find this name undesirable. I want a name most will like to pronounce and relate to others.
One more piece of Amanitology news – we have submitted for a review a short taxonomic paper naming the commonly known Amanita franchetii in California as Amanita augusta. This species has been long overdue for proper placement. Of course, good illustrations, phylogenetic analysis, etc. are all parts of our standard delivery package that we never fall short of. I hope you like that name augusta – it means – “venerable, majestic, etc.” – I think the name is short, simple, feminine and meaningful in the context of this beautiful species.
Best to all,
you can see lots of these all over MO by searching for breckonii and gemmata exannulata.
when I first collected and documented this species in November of 2000 (I certainly wasn’t the first to have seen it, but it was the first time that I had seen it), I sent a copy of my illustration to Rod Tulloss. He called it at the time, and only with an illustration, Amanita gemmata var. exannulata. And that indeed is what I called it for many years.
By 2005, Darvin DeShazer had convinced me that this species was actually called breckonii, and showed me photos of this species, published in Thiers’ Agaricales of California to prove it. I traveled up to Darvin DeShazer’s house in Sebastopol to sketch this example from Humboldt:
Fast forward several years ago, and Dimitar, after goading me to “show him the money” on this species, went to SFSU and photographed a drawer full of the type specimen of breckonii (collected on the SFSU campus rather than up in Humboldt Co.)…that resembled a muscaria-like amanita.
Existing names for this distinctive fungus were thrown into a tizzy.
WE all know it, but we haven’t quite decided what to call it yet. Name pending.
In regards to pileipellis, it isn’t always useful to give a label to what you have seen. Sometimes the mushroom doesn’t have anything that quite fits into the categories commonly used (which many times are used incorrectly…YIKES!), such as cutis, ixocutis, ixomixocutis, etc etc etc etc etc etc. So what to do in that case? Leave the full description, mentioning the relative disposition of the kinds of cells/hyphae. For the reader this is more informative.
As a suggestion, if you are patient enough, and before putting into words what you observe microscopically, try and draw what you see (at different depths..moving the fine adjustment knob back and forth). Drawing a section can teach you how to see subtle details, observation alone seems to miss at times. I find it useful (when terminology is confusing) to use my own words to best describe what I have just seen/drawn. Later on, I find selecting the proper terms to use (let’s say like that of types of pileipellis) is less daunting when you can compare it to what you have. Hopefully you can compare with terminology particular to the genus in question. This will save you a lot of headaches.
Good luck and don’t despair!
thanks for your comments. this being more of an educational exercise, I never expected to achieve anything near the level of comprehensiveness to satisfy a holotype collection. just reading through the rather head-spinning degree of detail in pileipellis characteristics is humbling enough. as the expectations for the assignment don’t seem to require detail of such meticulous proportions, my “write-up” (essentially the notes from this observation printed in 8 pt. font and placed in the specimen box) will be more general than would be ideal. without question, I aim to up my taxonomy game as time goes on. it’s good to know that there are experts who are willing to take some time out for those of us still getting our legs.
Nice observation Myxomop. We in Nor. Cal. are very familiar with this Amanita in the /gemmata clade. It tends to have a softer cottony UV than the other gemmatoid Amanita we see. We have some good molecular data already, but the group is complex and needs very careful study. I will let everyone know as we progress in our studies. The idea is to set some order in this clade and name the species properly. So far the two gemmatoid Amanitas that we commonly see are not closely related, but still within the same clade. More to come.
That’s a cool idea.
I’m sure it exposes the students to a lot of interesting stuff, not to mention all the technical terms that can vary from one genus to the next. With a large number of visible characters, Amanita can give a lot of “payback” for the work expended with the microscope; however, checking all the characters for every taxon is not a small task. Once a person asked me to identify material for a study in the eastern U.S. Many of the dried specimens were accompanied by no notes and had no illustration; but I was expected to work through them all. Because of the limited literature and my limited experience at the time, it took me nearly a week to recognize one of the most common autumn species in the eastern states. Did that make me a believer in photography and detailed annotation of fresh material? Yup. Do I do what I should every time no matter how many collections come in and no matter how long it takes. No. It is not humanly possible; however, it is scientifically necessary to have everything done several times in order to describe a new species.
You’ve probably noticed that, even restricting one’s self to agarics, different notes, different chemical tests, etc. are required for different genera. When I was helping with the protocols for a Costa Rican biodiversity study of fungi, I realized that to get the most out of the material it would be best if a Russulales expert designed the field note form for russulas, an Amanitaceae expert designed the field not form for amanitas, etc.
Of course that is not practical. Practicality is a mother of entropy…, but so are we all.
Whatever the cause, you have collections that don’t quite satisfy the requirements for a well-documented holotype. Then you still may have to ask the expert to come down and go into the field with the parataxonomists and try to find the particular mushroom again.
I can’t quite picture where these cells fit into the lamellar trama. I need some orientation.
You don’t need the word “elongating” … that’s always the business of hyphae.
My suggestion is that you have terms that separate “normal” hyphae from vascular hyphae and talk about “orientation.”
1) describe the thickness and layering first…these are variables that can be segregated from the cell descriptions.
2) describe what’s going on with filamentous [undifferentiated] hyphae second.
3) describe any vascular hyphae that may be present third.
4) describe presence or absence of clamps last.
5) if there are true inflated segments in the hyphae of the subpellis (very rare in an Amanita pellis), treat them as “intercalary cells” under the filamentous hyphae.
In my descriptions I tend to shy away from terms such as “ixocutis” in favor of describing in detail what I see. An ixocutis could be gelatinizing only in the uppermost one or two hyphae or gelatinization would take up half or more of the thickness of the pellis. In the Vaginatae I’ve seen several case in which there is no true suprapellis of any thickness; and I just said “gelatinizing only at surface” in my description. Placement and degree and depth of gelatinization is related to the ontogeny (basidiome development) and is definitely species-dependent. If you’re describing a species, I think you should describe everything that you think MIGHT BE species dependent. In the end, I’ve found that a great number of the characters I described do indeed appear to be species dependent. You only find this out if you describe everything you can for every species.
Did I screw up along the way and miss some stuff. Probably.
Be sure to segregate septate hyphae with yellowish refractive walls from vascular hyphae (which arise from a septate hyphae…if you can find that, but lack septa and contain insoluble stuff that leaks out in “bumps” when you (unavoidably) break a vascular hypha).
Hence, a description in which there is a distinct upper gelatinized layer to the pileipellis would look like this:
PILEIPELLIS: ? – ? um thick, comprising two layers; suprapellis ? – ? um thick, entirely gelatinized, colorless; subpellis ? – ? um thick, ungelatinized, [color, e.g., “yellow-orange”] in exsiccata; filamentous undifferentiated hyphae ? – ? um thick, branching, subradially arranged, densely interwoven vertically, etc.; vascular hyphae ? – ? um wide, infrequent, sinuous, infrequently branching, sometimes crossing between supra- and subpellis, apparently ungelatinized when in suprapellis, sometimes penetrating pileus context; clamps [give qualitative term for frequency, e.g., “rather common”].
A lot of the adjectival phrases are intended to stand as examples. Don’t assume that the above “model” will always apply. There are cases in which the pileipellis is not at all well-developed…just a sort of transitional zone between the pileus context and the volva. Cristina has just found a species in which we had to create a phrase (“subvelar layer”) for a pileipellis that has an ungelatinized thin layer on top (connected to the volva), THEN a suprapellis, and THEN a subpellis.
So there can be unusual variations.
The “model” (above) applies in a very large number of cases.
I’ve done as you’ve advised and reexamined the pileipellis. when seen from above, there’s nothing erect about it. i’m calling it an ixocutis of radially elongated hyphae. i’ve updated the notes to match this correction and replaced the old pileipellis photomicrographs with a new scalp section. would you agree with this description?
been learning boatloads from these six specimens (http://mushroomobserver.org/species_list/show_species_list/260). thanks for your continued help. I’ll do my damnedest to get this material in the mail to you before taking off for the holidays. otherwise, expect it in early january.
I think your thoughts are solid. To get a feel for the real range of size of the basidia, check them out in material of different ages and at different distances from edge of the gill. Generally the basidia are smaller near the edge of the gill and smaller in older parts of the gill (nearest the stem). The largest basidia often include some of the earliest to form. The subhymenium deepens a bit with age and basidia become shorter as they arise farther from the central stratum. It is as if there were an invisible wall (instead of a glass ceiling) to keep the tips of basidia from opposite gill faces from getting too close to each other. After all an ejected spore is not supposed to hit an opposing basidium.
I think that the illustrations title as the pileipellis may show something else. I suggest trying to view the pileipellis in a scalp mount (look down on the top of a very thin slice of the surface of the cap that you have arranged so that you known which edge of the scalp was nearest the margin. You should get a clue that the hyphe of the pileipellis are pretty much radially arranged and follow the curve of the surface. It looks like there might be a lot of partially gelatinized clutter from the universal veil on the surface in one of your photos labeled the pileipellis. In your scalp mount you should make every effort to have as little detritus as possible. Getting the impression that the hyphae of the pileipellis are vertically oriented is very likely to be a mistake. (E.g., the supposed trichoderm reported for A. magniverrucata was actually part of the volva…see my paper (from 2009, I think) on a revision o that species. Also see the description of Amanita basiana, which is one of the species in which the hyphae that connect the universal veil to the pileipellis in the button stage persist in mature material for quite a while before breaking down. The eastern farinosa has this sort of vertical element connecting the pileipellis to the volva; and the western one may have the same thing going on…in case you want to check out an example of this phenomenon. I really recommend drawing as a way of seeing when working with the scope.
Going back to your pileipellis, in addition to the scalp view to determine that whether or not you can see subradial orientation in the hyphae of the pileipellis, a cross section is critical to understanding the vertical structure. If you’d like to discuss the further, let me know.
is there any sense in measuring basidioles? my guess would be no, since immature structures, by definition, can come in a range of sizes up to (but perhaps not including) their mature counterparts. strangely, not one of three gill mounts is showing more than three or four true basidia, which is making their measurement rather difficult.
ps: Rod, once I finish my write-up, I’ll be sure to send some material your way.
I’d like to take a look at a mature specimen that has the interesting spores.
Please contact me so that I can get your email address in order to send you a few things that might be useful.
The Amanita spore wall is very, very thin (perhaps a couple tenths of a micrometer under normal conditions…there’s more about this in the Tulloss and Halling paper related to crassospores). When a spore is damaged and collapses, it is usually something you might call a “catastrophic failure.” There is a significant change in the shape of the spore, not simply a distribution of small impressions in the spore’s outline.
Crassospores are apparently the result of disease or infection. They have been seen in a number of species of sections Amanita and Caesareae. I listed a few species in which I had seen such spores in the note that Debbie(?) posted on BAMS. So, no, presence of crassospores is not diagnostic of a taxon or group of taxa. Appearance of crassospores seems to be linked (in some cases) to a geographic area that is (hypothetically) somehow infected. There is a lake in the Argentine Andes by which several collections of two species of Amanita have been collected that had crassospores (A. morenoi and A. pseudospreta).
It would really be worthwhile (I think) for you to see the Tulloss and Halling paper that discusses crassospores in the Argentine species. The paper is listed on this page:
Control-F and search for “crassospores”…you’ll also find at least one other (older paper) that discusses crassospores. The bibliography of the Tulloss and Halling paper will also give you leads to follow.
I hope that you get a chance to check this out. My secondary hope (of course) is that I am not pointing you down a blind alley. :(
crassospores are or are not diagnostic of a particular Amanita sp. or lineage? is it possible to determine from these pictures whether or not the indentations are this species’ spores in their natural state or a deformation due to age?
there is definitely enough dessicata to go around. I’d be happy to mail something your way if you’d like to take a closer look.
another possibility…namely, that you are seeing large granules or small guttules filling the spores. However, this would not be a possible cause of the apparent indenting of the spore wall (noted around the outline of the spores), which is the appearance that one would expect of crassospores.
to also have plenty of spores that are “normal.”
First of all… I agree. This is a good observation with plenty of detail and interesting photographs at the macro and micro levels. Congratulations.
I have never seen this sort of “apparent” patterning of the spore wall happen due to collapse of the wall. The spore wall is normally a few tenths of a micrometer thick at best in Amanita. Spore collapse in Amanita [edited] tends to be more of a “catastrophic failure.”
This kind of patterning (real patterning) occurs in crassospores. Crassospores are apparently the result of an “infection” of a fruiting body, possibly by a virus. We’ve had the good fortune to see crassospores earlier this year in a specimen of Amanita multisquamosa. There was a discussion of the topic on the BAMS yahoo site. The image is posted on the techtab of this page:
The three dimensional “golf ball” or “cratered moon” effect is clearly visible when using a scope equipped with Nomarski Differential Interference Contrast capability.
The BAMS posting includes input from me about a paper that I wrote with Dr. Roy Halling concerning crassospores as observed in some material from Argentina. If material of the present collection with the apparently patterned spores is available for review, it would be interesting to try to photograph the patterning with Nomarski DIC. I can do this; or, perhaps, you’d rather keep the material in the western states and locate a lab that can help you there.
Now this is a top notch observation. Nice work.
More solid work on this not-breckonii taxon.
Both my knowledge of microcharacters and my skill with a microscope are in their baby stages. That said, taking my cues from the MushroomTalk discussion on the as of yet unnamed A. breckonii look-alike, I looked specifically for clamps at the base of basidia and narrow, cylindrical spores. A discrepancy in spore shape was noticeable right away. Basidia clamps resisted my every painstaking attempt to locate them, if they are there at all, but I’m just as inclined to blame it on inexperience.
The material is being dried for deposit in the SFSU herbarium. It will constitute one of six requisite descriptions/depositions for Dr. Desjardin’s Mushroom Taxonomy course. Whether or not an ID is not readily discernible from the data presented thus far, I’m eager to continue working with the material with some guidance from whatever gracious Amanitologists would care to point me in the right direction.
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