Notes: Growing in a group of two, from soil, at the base of a Pinus virginiana, in mixed woods.
7.2-9.8cm broad, margin striations (.8-1cm), convex to nearly plane, smooth with white patches or warts
Orange to peach in the center fading to yellow to pale yellow towards the margin
Free, white to cream, crowded
12.7-13.2cm long, Apex: 1.2-1.4cm wide, Base: 1.5-2.4cm wide, superior yellow-ish membranous skirt-like partial veil, some veil remnants on the margin.
I don’t know how to describe the texture of the stem (any help?)
smelled just like syrup
Those damn white larvae ate these guys, but I got a spore print out of them!
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you have there the business ends of the basidia. Moving into the gill perpendicular to the surface of the gill, you next come to the bases of basidia. Basidia are of different lengths. The longest are oldest and are the first to produce spores. After spores are produced on a basidium it collapses and disappears to make room for more basidia. Below the the basidia is a distinctive layer that takes different forms in differents species and at different times in development. We have to pick a standard point for description of the subhymenium, the subhymenial base (also called the lateral stratum) and the central stratum. Characters of the tissues are most differentiated at maturity; so this is a good time to pick. For the sake of argument, let’s say that maturity is reached when a good spore print from whole gills (stipe to cap margin) becomes possible.
The subhymenium can be cellular (composed of approximately globose cells tightly packed…like plant tissue), but it can also take other forms showing distinct branching of partially inflated hyphal segments and other variation.
Still moving toward the center of the gill along the same path perpendicular to the gills surface, we pass through the subhymenium and enter the subhymenial base or lateral stratum. This is the part of the gill dominated by hyphae and groups of cells that, in combination, provide the divergent character of the Amanita gill. If you look at a gill in cross-section, you will see that the combination of the central stratum and the lateral strata give the impression of a stack of “Y’s” (open end down) running from the cap’s flesh to the gill’s margin. The surfaces of the gill slant slightly inward from the cap’s flesh to the gill edge (that is, the gill narrows as it goes from cap to gill edge). This makes it easier for spore dropped from basidia near the cap flesh to make it all the way out of the hymenium and into whatever micro air currents there may be.
I make all my measurements at about the midpoint between the cap flesh and the gill’s edge.
These sections are not easy to make.
Rehydration of tissues is tricky.
Well-dried tissues are necessary to good rehydration.
I always record my estimate of the quality of rehydration when I make measurements of the thickness of the gill trama.
You will end up with a range even if you use only one fruiting body on which to collect data because of the variable length of basidia (remember they get shorter with fruiting body age).
So the depth of the subhymenium is something shallower than the distance from the base of a basidium (a position somewhere in the subhymenium) to the central stratum because most of the distance you measure (in most species) is the thickness of the subhymenial base (lateral stratum).
I don’t have enough data on the yellow variant (I don’t think that “race” works here…as it would in birds). At any rate no one has enough information to know why the pigments vary so far as I know. At the moment, it seems likely to be the case that some genetically determined process is being turned off or on or turned partially off or partially on. This has been studied in plants that have multiple pigment patterns for their flowers.
As I said before, I only have good data regarding gill thickness on the red-capped taxa. I’ve just never had time to study the yellow-capped variants in sufficient depth.
I hope this helps.
Does that include both the eastern and western race? How does the “breadth of the central stratum and thickness of the tissue between the central stratum and the base of the nearest basidia” differ from the depth of the subhymenium? Sorry if I’m being a pest, but, I don’t think anyone else is going to answer these questions for me here. I’m looking forward to examining some cross-sections this year.
The breadth of the central stratum and thickness of the tissue between the central stratum and the base of the nearest basidia works to differentiate the European muscaria and the dNAm.
Would you say the depth of the subhymenium is not a valid way to delineate the eurasian clade from the dNAm clade? Should we consider the eastern race “blondes” with big “teeth”?
I have put all my effort into the description of the red variant. The data on the var. guessowii page is based on very limited samples. The character is variable.
The depth of the subhymenium is (according to you) a reliable way to separate (morphologically) the eurasian from dNAm clade. However the dNAm eastern race has a subhymenium depth that is closer to the eurasian clade (according to info at your website). How can this subhymenium info be used to separate the eurasian from the dNAm clade when the depth of the subhymenium of the dNAm clade is different between the east and west?
This is not a case in which the phrase “having it both ways” is appropriate.
One is trying to have things both ways when one maintains the traffic light is both red and green at the same time or when one runs a race with a broken leg.
In the present case we have plentiful phylogenetic evidence that the color variants of the dominant North American muscarioid taxon are interbreeding. Hence, the most appropriate hypothesis is that the color variants belong to a single species.
We also have phylogenetic evidence that the apparently, nearly circumboreal Amanita muscaria is a distinct species from the dominant North American muscarioid (dNAm) taxon. No confllict so far.
We also have morphological evidence that Amanita muscaria is a distinct species from the dNAm taxon. Supportive of the genetic evidence. Still no conflict.
That’s the story. No logical conflicts. And no one “having things both ways.”
What could change the picture? Having the ability to compare a larger chunk of the genome of muscarioids might show the picture to be different and/0r more complex than the partial view we have from three or four genes at the moment.
Maybe we will have a surprise from another research direction.
Right now we have a consistent set of hypotheses:
1. Amanita muscaria and dNAm are distinct species.
2. Amanita muscaria exhibits natural color variation (incuding radial stirpes!).
3. The dNAm exhibits natural color variation.
That’s more than what we new a decade ago. Not too bad.
You can’t have it both ways. If you use the depth of the subhymenium and spores to differentiate the the eurasian and north american clade. Why ignore the same morphological differences within members of the american clade (eastern yellow and western red). If the yellow color of the eastern species is not genetic, what possible environmental factors would cause this? Also has it been suggested that different forms cannot interbreed? I see Rod does not like to use the terms var. or f., if not variety or form what possible designation do we give these different races.
I think these conversations were partially (at least) face-to-face.
Bas represented the rank decisions based on “differences” as a rule of thumb for a genus in which mating experiments had not yet been carried out successfully. I am quite sure his species concept would have been based on mating studies if these had been possible.
Things have changed even since the MO observation that you cited in your email.
On the observation you cited there are two points to make. (Maybe you could post this email on that observation as a comment?)
I. We no longer assume that a new name will be required for the red-white-yellow muscarioid that is the dominant species of this group in North America.
1. The extended studies with Dr. Geml over several additional years have STILL NOT produced evidence that differs from the viewpoing of the original Geml et al. publication: The red, yellow, and white capped collections that occur from southern Alaska to the eastern coast of Canada and throughout the northeastern quarter of the U.S. and down the West Coasts of Canada and the U.S. Costa Rica and further south (possibly having been introduced in the Pine plantations further south (where Pine is not native) is apparently one species with all the populations found to date apparently capable of interbreeding.
2. We have realized that if we cannot produce counter evidence using red-capped and yellow-capped collections of the most common American muscarioid, then the one place that the hypothesis may break down would involve purely white-capped taxa (no yellow showing in the pileipellis in cross-section throughtout the organism’s life).
3. Such white-capped collections are infrequently reported, and we have been able to collect very few of them.
4. We are going all out now asking for people to collect white-capped muscarioids from the region of interest, dry them carefully, but quickly, take pictures with rulers in them or (better) take detailed notes using my collecting form. Finally, send me the documented collections.
5. We particularly want to get material from Michigan (from which A. chrysoblema was originally reported) and from New York (from which A. muscaria var. alba was originally reported).
6. If we don’t see two different taxa in specimens we received, we will conclude that the name for the North American species is required to be Amanita chrysoblema based on the existing evidence.
7. If we find two, distinct white muscarioids in eastern North America, we will have to get detailed morphological data on both (THIS IS WHY IT IS SO IMPORTANT THAT COLLECTORS DESCRIBE THEIR COLLECTIONS IN DETAIL.) Amanita chrysoblema was reported to stain yellow when cut. Amanita muscaria var. alba was described without mention of staining. The yellow staining could be of genetic origin in the white muscarioid material; but it could also be due to the yellowing syndrome, which I have reported in normally unstaining species of Amanita.
II. Separation of taxa with the “rule of thumb for Amanita” that was described by Bas.
1. The rule was at least sometimes too conservative and rapidly becoming outdate. (The changes I have seen in my life time are quite astounding.)
2. After very thorough study of the European Amanita muscaria and the taxon discussed above (“most plentiful North American muscarioid”), it is clear that they do not currently interbreed.
3. Both these taxa can have caps that are red or yellow or white. Both can be found with yellow or whitish volvas, etc. After extensive morphological study, I found two characters that can differentiate populations (maybe not always individual fruiting bodies) of the two taxa: size/shape of spores and thickness of the lamella trama (gill flesh). That’s basically two characters, not three. Bas did not use the thickness of the lamella trama; so in his work these good species would have been found to differ by one character that he could measure. Of course, genetic studies provide other distinctions that (in this case) have nothing to do with morphology so far as we know. So there are lots of characters that Dr. Bas couldn’t detect and, hence, couldn’t consider. We have really entered a world in which so many new characters can be measured through chemistry that the old “count the differences rule of thumb” has become outdated.
4. I suspect that the rank of form may rarely be used from this point forward. It was extensively overused in the past for cases of minor variations in pigment that have nothing to do with defining a biological species. This was fairly obvious before molecular studies became possible.
5. I have not yet seen a case where the term “variety” is (in Amanita) sustained after genetic studies although I can imagine such a case.
6. On the other hand, we are finding examples of “almost species”…for example taxa that retain a low probability of successful mating despite millions of years of geographical separation. But why should we expect “clean boundaries”? Evolution is a process that is dependent on random change. Who would think that this would generate “clean boundaries” on all occasions?
7. On a third hand, we are finding examples of what appear to be the results of incomplete segregation of taxa due to geographic separation followed by successful interbreeding of the offspring once the populations are no longer geographically separated. This situation can sometimes create a highly complex “swarm” of interbreeding hybrids WITHIN A SINGLE BIOLOGICAL SPECIES. One example of this is present in Amanita in eastern North America…a citrina-like, purple-staining mushroom that is NOT Amanita lavendula.
The hybrid taxon is presently called “Amanita sp-lavendula01.”
Amanita citrina [form or variety] lavendula is a good species and is expected to become Amanita lavendula.
Amanita brunnescens [form or variety] straminea is a good species and is expected to become Amanita americitrina.
The WAO taxon pages for Amanita sp-lavendula02 and Amanita sp-lavendula03 appear to assignable to the two names just given (one each, of course). So these temporary codes will be merged into morphological concepts for lavendula (sp-lavendula02) and americitrina (sp-lavendula03). This is our hypothesis at present. Sporograph comparisons for the three temporary code names are available on one or more of their taxon pages.
This is a fast summary. You might want to watch the changes in the above listed group of citrina-like amanitas’ taxon pages on the www.amanitaceae.org (WAO) web site. Most of the critical data is on the technical tabs.
Dr. R. E. Tulloss
firstname.lastname@example.org [<= Share DropBox folders to this email only, please]
Herbarium Amanitarum Rooseveltensis
P. O. Box 57, Roosevelt, NJ 08555-0057, USA
On 2/15/2014 10:19 AM, email@example.com wrote:
Question from Another User
lightworkerpeace asked the Mushroom Observer website
to forward the following email to you:
I was partially reviewing your notes from http://mushroomobserver.org/68146?q=1kPiY
and I was wondering if you have the original transcript(s) of your communication >with Dr. Bas regarding the three differences required to describe a new species.
Thank you Rod for taking the time to talk about this. I guess we will just have to wait and see.
Britney: Thanks! I’m sure they’ll start popping up soon!
Dr. Tulloss: Thank you for explaining all that you did. It was very helpful!
In the present case, if you trace back through the tree from the leaves representing samples from yellow-capped specimens to find a point at at which all paths down the tree from these leaves meet, you will find that the branch of the tree you have defined includes leaves representing red-capped species. That is the case with the tree published by Geml et al. I am the second author of the paper. Dr. Geml and I discussed this point in particular.
There is no justification…in that tree…for designation of a subspecific taxon consisting of only yellow-capped, northeastern North American muscarioid mushrooms.
Because we were concerned that there was more to be said on the matter, I have been collecting dried, yellow-capped specimens hoping that we might find that there were a pretty big, distinct clade of yellow-capped material if we just had a large enough sample from a large enough geographic area. In obtaining the new samples we have significantly expanded the geographic area from which yellow-capped material has been collected. I assure you I wouldn’t have bothered with this if there was already good evidence that there was a yellow-capped clade that could be said to represent a phylogenetic infraspecific group (like a variety or a subspecies).
You are quite correct that from a morphological view the conservative thing to do is first define the red taxon that Singer distinguished as A. muscaria subsp. flavivolvata and then progress to demonstrating exactly how similar or dissimilar the yellow populations are with respect to a clearly defined North American red-capped species. That is the approach that I am indeed taking.
I think I am following a logical, conservative approach, taking one step at a time through a logical process.
Just a day or two ago we were talking about hypotheses. Testing these hypotheses is also part of the ball game. The current hypothesis is based on the paper of Geml et al. I am indeed involved in getting material together to test that hypothesis (see above). I do not have absolute certainty about the current hypothesis. Nor do I have the hope of attaining an absolute truth of the situation.
It is certainly possible that if we were to have a larger sample of muscarioid populations across northern North America we might find that there were one or more clades of yellow-capped samples suggesting one or more infraspecific taxa could be justified. The opposite is also possible.
I’m not sure what you think is wrong with what I’ve said or am I’m saying. I’m really trying to do my best and lay the whole thing out with all the uncertainties and all the warts. I’m not rooting for one outcome or another.
I can’t spend any more time on this subject for awhile.
When I look at Geml’s cladogram I see the eastern taxon grouped together and since no red “flavivolvata” occurs in the east I think it would be reasonable to assume that it is different on some level, although still part of the flavivolvata clade.
Besides a difference in color from the western taxon there is also a difference in spore size(slightly smaller for the eastern yellow taxon).
From what I can tell the yellow taxon that occurs in the PNW is actually part of the muscaria clade not the flavivolvata clade and is represented by clade II/A in Geml’s cladogram.
Some of the clades are polymorphic while other are monomorphic, Geml’s paper clearly states this.
If your convinced they are exactly they same, why haven’t you been using any specimens from the east in the technical description of your provisional name?
I hope this works. I’ll be back to edit it later…I’m sure.
Matt’s pictures are pictures of members of one of many local yellow populations in northeastern North America. This group of populations was (in recent times…and still on WAO) treated under this name: Amanita muscaria var. guessowii.
Geml et la. (phylogenetically) and RET (morphologically) produced a bunch of data that said, among other things, approximately this: “All the northeastern yellow-capped muscarioid populations [in the group defined above] belong in the same species as all the red-capped populations that we call Amanita_muscaria subsp. flavivolvata.”
I think we have all been over this many, many times. I’d like this to be the last time that I do it for awhile. :-)
Now, Herb raises an issue that requires a definition (actually a couple of them). We need to work our way toward the issue of what a subspecies is.
Before Bas (and for a lot of authors who ignored Bas in later years), there was no rule of thumb (or any other kind of systematic way) of deciding when to use words like “variety” or “form” or “subspecies” in the genus Amanita (or in a lot of [all of the?} genera. Bas is an orderly, careful thinker, who took his time thinking about this problem. And he decided to use this rule: [I’m putting words in his mouth that I can’t guarantee are the same words that he said to me several decades back.] “If I can find three unrelated characters that differ between two taxa, then I will feel comfortable saying that they are different species. [Remember, he greatly increased the number of characters that were important in describing an amanita.] If I can’t find three, but only two characters separate something that I believe to be new from an existing species, then I will call the new taxon a variety of the older one. [This turned out to be quite conservative compared to phylogenetic trees.] Finally, if I find only one character separates a new taxon from an older one, I will make the new one a form of the older one.”
It is important to understand that to this day, only a handful of people describing amanitas have caught on to the brilliance of Bas’ thesis and this rule of thumb in particular. In many works “form” and “variety” are used with no obvious reason for picking one word or the other. The good thing is that, outside of certain authors in southern Europe, the use of ranks of variety and form have not been very frequently used in Amanita in recent years. Where they have been used, they have often been misused. But we’re not going to go into that here.
OK. Now, what do we do about “subspecies.” Bas’ choice is in line with taxonomic tradition. He said (again paraphrasing), “If a newly discovered taxon differs from an established species by two unrelated characters AND the range of the new taxon is disjunct from the range of the older one, then I will call the new taxon a subspecies of the older one.”
Now we have to take genetics into account.
After collaborating with several phylogenetic labs looking into the phylogeny of Amanita, I’ve noticed that morphological taxonomists working in Amanita did a pretty good job. We are not finding that we “over split.” If anything we are finding that differences that were represented as being at the level of variety of subspecies have proven to be differences at the level of species.
What would a phylogenetic concept of a variety or subspecies be like? I am a student, not a professor in phylogenetics; but let me see if I can explain what I understand. Probably most of the readers of this have seen a phylogenetic tree. For our purposes, we can say that the absolute tips of the branches (the “leaves”) correspond to samples of a particular gene or part of a gene that was taken from a particular specimen. The tree is constructed by computer analysis from the information about these genetic samples. The analysis produces a branching structure in the tree. A separate analysis can also evaluate (separately) if genetic information is actively being exchanged between the organisms from which samples have been taken. A phylogenetic branch is a model indicating that all the sampled organisms represented by the leaves on the branch had a supposed common ancestor. These branches are also called clades. Certain clades represent phylogenetic species. One of the properties of a phylogenetic species is that the appropriate computer analysis can detect that all the sampled organisms represented by the leaves o eaves of the branch have been recently exchanging genetic information. They all belong to the “gene pool” of on species.
If we’ve got a branch as a model of a species, what would a phylogenetic variety or subspecies look like in terms of a part of that branch? Well, for there to be any taxonomic meaning, the variety or subspecies has to take up a whole sub-branch with all its twigs and all its leaves. In other words, all the sampled organisms that would be assigned to that variety or subspecies would have to have a presumed single common ancestor. Or to put it in the negative, a bunch of twigs from various parts of the species branch that don’t make up a whole sub-branch cannot a phylogenetic equivalent of a variety or subspecies be. Thank you, Yoda.
Now let us return to our sheep. [Big “no prize” for the person who can remind me of the source of the sentence.]
What did the big muscarioid species tree of Geml et al. look like with regard to the yellow-capped populations with which we started this excursion? Twigs. Twigs Twigs.
Sorry folks we have no evidence to suggest that the yellow-capped guys being treated as anything but the blondes in a family of redheads. All we can say right now (given the evidence we have) is that there are lots of not-too-closely-related yellow-capped populations of Amanita muxcaria subsp. flavivolvata (or use the provisional species name if you wish) in the northeastern part (and north central part) of North America.
After spending quite a lot of time on this observation today, I hope that we can let this topic rest for awhile. I think I did a reasonable job this time and went into more detail on a number of related subtopics than I’ve done on MO in the past.
So bookmark this explanation if you think you’re going to need it to pass on in a future discussion of this topic on another observation. You bet it’ll happen. Believe me. This topic is like the cat that kept coming back in the old folk song.
Now, before I quite, I want to thank Herb. Herb you have now got me completely convinced (and this exercise on this observation is the thing that did it) that there is no better way to lay this whole thing to rest than to use the provisional name now and to get it in print as soon as I can.
@ Matt, great pictures man~! I am not seeing any Amanita in my neck of the woods!
@Rod, thank you for always lending your info. You are a treasure here.
I like the idea of looking at the eastern taxon as a subspecies from the western species. It’s certainly not exactly the same as flavivolvata.
This is the name that has been used at this rank for the eastern taxon, in publications.
If you look you’ll see its placed in synonymy here, if your vote it down your voting down guessowii also.
When Lange first used the infraspecific name americana with regard to material of A. muscaria sensu lato that he had seen on his travels in the U.S., he said that the epithet might be applied to such material. From the words he used, he did not intend to publish the name. If he had intended to publish the name, he would have provided a description, which he did not. But whether or not there were a description, when an author does not definitively accept the name, the International Code of Botanical Nomenclature (ICBN) says the name is not valid. The name, Amanita muscaria var. americana in the combination attributed to Lange is not a valid name.
It is true that Singer could have picked up the name and validated it, but he did not. By the date of Singer’s publication, a Latin diagnosis was required for valid publication. Singer did not supply a Latin diagnosis. So the name mentioned by Singer is not valid either.
Moreover, I would argue that it is speculative to give the name a meaning. In the history Amanita names that I inherited from Dr. Bas and have tried to maintain as best I can since that time, the choice has been to list the invalid names of Lange and Singer that have just been discussed in a separate list of invalid names and nomina nuda (roughly “names published without being given a meaning”) and without guessing at possible synonyms or making any other speculation about them.
Now that we have evidence (so far not contradicted) that this taxon is a color variant of A. muscaria subsp. flavivolvata, the earliest valid (i.e., correct) name for the mushroom depicted on this page at the rank of subspecies is A. muscaria subsp. flavivolvata.
If you go for the idea that Geml et al. demonstrated phylogenetically and that RET demonstrated morphologically that the last mentioned taxon is actually a different species distinct from A. muscaria, then you don’t have a valid name to use. You can use a provisional name, which has been discussed (um)…let’s say “very extenstively” on MO. Or you can use one of the two valid names that are available for the mushroom.
There are three choices: var. guessowii, subsp. flavivolvata (“yellow variant”), and the provisional species name (“yellow variant”). So far as I know, there are no other available choices.
Under these circumstances, it seems to me that the provisional name is not so painful to use when you have no other available correct name at the rank (i.e., species) that is currently deemed correct for this mushroom.
Created: 2011-05-26 12:44:37 EDT (-0400)
Last modified: 2014-02-17 21:08:23 EST (-0500)
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