Amanita calyptroderma G.F. Atk. & V.G. Ballen, A. calyptrata Peck, and A. lanei (Murrill) Sacc. & Trott.

These names have been variously applied to what appear (at least) to be two distinct populations in western North America. The first fruits in the Fall (Nov.-Dec. with occasional records from early January) and has an orange to brown pileus and usually grows with madrone, tan oak, and Douglas fir. The second is a Spring-fruiting (Feb.-Jun. with occasional records from early January) species with a lemon-yellow to pale yellow to cream pileus that grows with live oak in coastal California, pine in the Sierra Nevada and San Bernardino mountains, and occasionally under fir in the Pacific Northwest.

Other than these clearly distinguishing features, the two amanitas are fairly similar. They each have an unusually thick universal veil which leaves an ample volval sack and a thick white patch on the cap. Another interesting feature is that both species are edible. In my experience, when cooked, the Fall entity generally has a more strongly fishy, slightly metallic flavor. The Spring species has a milder flavor, and I’ve never noticed the metallic component in the latter.

Finally to add to the confusion in northern California I have collected what appeared to be a pure white version of the Fall fruiting entity. It was fruiting in the same habitat (mixed madrone, fir, and tanoak) and at the same time of year as the fall species. I consider this to be an albino variant of the fall organism.

At the current time (April 2007), no one is completely confident in how the names should be correctly applied. Most of what I know about the nomenclatural and taxonomic issues comes from personal communications with Rod Tulloss, Jan Lindgren and David Arora. I take full responsibility for any errors.

The problem started in 1900 when Peck applied the name Amanita calyptrata to an “Autumn” collection from Oregon. This collection generally matches both of the species described above, except that it was described as being greenish. H. D. Thiers suggests that this is a weather induced variation, but I have never seen an example of it. For the sake of completeness of our story, Peck’s name was recombined in the genus Venenarius by W. A. Murrill in 1912.

The name Amanita calyptrata has a problem with the rules of botanical nomenclature. It was previous applied by the French encyclopedist Lamarck¹ to a completely different European mushroom that is now in a completely different genus. At that time the name Amanita was applied to most gilled agarics by Lamarck and others who followed a biologist with the latinized name Dillenius.

A few years later the name Amanita was applied as it is today for the first time by C.H. Persoon², but in any case the point remains that Amanita calyptrata cannot be used for the species described by Peck. Peck’s name is invalid according to the currently accepted nomenclatural rules, so A. calyptrata Peck non Lam. is to be rejected.

Before this was an issue for the rest of the world, W. A. Murrill published a new name for the species in 1914 based on Peck’s type collection – Venenarius lanei. Murrill was a purist following the American nomenclatural rules of that time and had banished the name Amanita altogether. Saccardo & Trotter, followers of the International Code of Botanical Nomenclature, which we follow today in modified form, recombined Murrill’s new species epithet in the genus Amanita in 1925 – yielding A. lanei.

However, before any of Murrill’s renamings entered the picture, another name (possibly/probably) for the same fungus was published by Atkinson and Ballen in 1909 – Amanita calyptroderma. Atkinson’s name is based on a type collected in “December” in southern California and forwarded to Cornell by Ballen. Ballen also forwarded other material to Atkinson including material collected in the Spring. Together they published a description that included both the Fall and Spring entities under their single new name. The important point for the meaning of the name is that its type was collected in the late Fall or early Winter. If the Fall and Spring variants are to be separated taxonomically, the Spring variant still needs a name.

According to David Arora, the type of A. calyptroderma was collected in Ben Lomond, and “it is unquestionably the familiar fall form of the coccoli. The cap is described as ‘maize yellow’, which, if we remember the era in which it was published when most corn varieties were a dark, rich yellow-orange, is a very good description of the fall form (excluding its tendency to be brown in some localities). For those readers familiar with the color Maize Yellow (a somewhat tannish or dingy yellow of moderate intensity) as defined in Ridgway’s classic book of colors (1912), Atkinson and Ballen were using their own personal terminology because Ridgway’s book was still three or four years from publication.

Spore size and shape may not be a help in differentiating the two seasonal variants. Using his preferred spore measurement variables, Rod Tulloss reports data from spores of the Fall variant as follows: [160 spores from 8 specimens from 6 collections] (8.6) 9.0 – 12.0 (24) × (5.5) 5.8 – 7.0 (8.5) µm, (L = 9.9 – 10.9 (11.1) µm; L’ = 10.4 µm; W = 6.3 – 6.6 µm; W’ = 6.5 µm; Q = (1.29) 1.43 – 1.81 (3.0); Q = 1.55 – 1.68 (1.69); Q’ = 1.62).

He reports the data from spores of the Spring variant as follows: [138 spores from 7 specimens from 6 collections] (9.0) 9.5 – 12.4 (17.2) × (5.5) 6.0 – 7.8 (9.8) µm, (L= 10.6 – 11.2 µm;L’= 10.9 µm; W= 6.6 – 7.3 µm; W’ = 6.9 µm; Q = (1.35) 1.40 – 1.77 (2.18); Q = 1.53 – 1.67 ; Q’ = 1.57).

The differences (to date) seem not to be very great; however, the majority of required microscopic analyses have not been carried out on either variant. [Stability in the ranges of spore size and shape (based on Q) is not achieved until nearly 500 spores have been measured.]

Interestingly, Dr. Anne Pringle mentioned that she sequenced DNA from the two variants and found that they are phylogenetically extremely close. She did not have sufficient data to draw conclusions or to publish her findings.

Based on all of the above, I use the name Amanita calyptroderma for both variants. I then indicate the season of collecting: Amanita calyptroderma “Fall variant” or “Spring variant”.

My reasoning is as follows: Since A. calyptroderma is the oldest validly published name for the species combined in Amanita, it must apply to the seasonal variant which is represented by its type collection (almost certainly the Fall variant). Amanita lanei is also a valid name, but is only usable if the original greenish type material turns out to belong to a species distinct from A. calyptroderma (which Thiers, for example, believed to be false). If it is distinct then it might refer to an out of season fruiting of the Spring variant or potentially a completely separate taxon which is quite rarely found, characterized by the greenish color body and Fall fruiting season. Given all this confusion, I find it easier to simply avoid the name A. lanei until such a time that the open questions have been resolved by comparing the type collections to the more carefully and completely described collections of the Fall and Spring variants.

Notes:

¹ This is the same Lamarck who was perhaps unjustly (see Steven Jay Gould) criticized for believing that giraffe’s necks were stretched and the stretched neck lengths were inherited.

² C.H. Persoon was a Dutch author who was widely known as a botanical illustrator at the time, but is now known for being the author of the formally conserved name Amanita and being the author of the fundamental work on gasteromycetes.)

References:

Arora, D. 1986. Mushrooms Demystified, 2nd Edition. Ten Speed Press, Berkeley, California. 959 pp. Discusses both forms as Amanita calyptrata. Includes color pictures of both.

Jenkins, D.T. 1986. Amanita of North America. Mad River Press, Eureka, CA. This text uses Amanita calyptrata Peck and mentions Amanita calyptroderma as a synonym. He mentions a pale yellow to white variety and a greenish variety. Interestingly he says both of these are ‘regularly collected’. Dr. Jenkins did not publish a study of the type of A. calyptrata in his article on Peck’s type collections. He did publish a brief account of the type of A. calyptroderma. The spore sizes provided are over 2 um higher on average than the size range resulting from Tulloss’ research. This is a relatively large difference. Both of the cited types will have to be reviewed.

Lincoff, G.H. 1981. The Audubon Society Field Guid to North American Mushrooms. Alfred A. Knopf, Inc., New York. Primarily discusses the orange-brown fall form as Amanita calyptroderma Atk. & Bal., but mentions two “color variants”, the Green-capped Coccora (A. calyptrata) and a white variant. The photograph is of the orange-brown “variant”.

Phillips, R. 1991. Mushrooms of North America. Little, Brown and Company, Bostan, MA. This text uses Amanita calyptrata Pk. The text says “color varies from whitish yellow to greenish to orange-brown and yellowish on the margin”. The included photograph is of the yellow spring variant. In the photo you can see dried live oak leaves.

Thiers, H.D. 1982. The Agaricales (Gilled Fungi) of California. I. Amanitaceae. Mad River Press. Eureka, CA. This text uses Amanita calyptrata Peck and lists Amanita calyptroderma Atk. & Ballen. as well as Amanita calyptratoides Peck (a distinct, smaller brown species with a collapsing annulus) as synonyms. Again the discussion mentions both the ‘commonly encountered form in which the color of the pileus is yellow’ and ‘the green or grayish state often seen in old basidiocarps or after exposure to low temperatures.’ Photographs are given of the fall variant and a ‘white form’.

Web references:

The CABI biosciences Index Fungorum (http://www.indexfungorum.org/Names/Names.asp, 4/21/2007) lists Amanita calyptroderma G.F. Atk. & V.G. Ballen as the only accepted name (green). It also lists Amanita calyptrata Peck, Amanita calyptrata Lam., Venenarius lanei Murrill and Amanita lanei (Murrill) Sacc. & Trotter all as questionable names (gray), but without clear synonymy to Amanita calyptroderma.

MykoWeb (http://www.mykoweb.com, 4/21/2007) lists Amanita lanei (Murr.) Sacc. & Trott. and gives Amanita calyptroderma and Amanita calyptrata as synonyms.

Rod Tulloss’ Amanita Site (4/21/2007) lists Amatnia lanei (Murrill) Sacc. & Trotter and gives Amanita calyptrata Peck non Lam. and Amanita calyptroderma G. F. Atk. & Ballen as synonyms.