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Provisional World Key to Species Closely Related to Amanita hemibapha with Notes on the “Slender Caesar’s Mushrooms” of Eastern North America

Rodham E. Tulloss, P. O. Box 57, Roosevelt, NJ 08555-0057, USA

Revision of (Tulloss.
1998).
Most recent change: April 22, 2007

Background tile: Amanita garabitoana on world map showing approximate distribution of Amanita stirps Hemibapha as currently known.


Amanita sp. CR6
Photo: R. E. Tulloss
  In 1986 [Boston Mycol. Club Bull. 41(3): 10-13], I wrote a little summary of what was known about Amanita hemibapha (Berk. & Broome) Sacc. and its subspecies. I reported then that the North American Caesar's mushroom should probably be called A. hemibapha. I have begun to study this group of amanitas in greater detail, and thought it was time for an update.

A More Conservative Approach

It seems that a more conservative attitude may be appropriate. Until a better understanding of all the involved taxa has been obtained, I now think it is best to treat our red “Caesar's mushroom” as a distinct species and to use the existing name for that, North American entity—Amanita jacksonii Pomerleau.

To give the reader a feeling for the number of North American taxa in the group of species similar to A. hemibapha (let's call it “stirps Hemibapha”), a provisional key to the group has been prepared. It is nearly certain that there are more taxa in the world than are accounted for in this key; for apparently new species of “stirps Hemibapha” seem to be reported on a regular basis by my correspondents and in the literature.

This key is restricted (as much as possible given current knowledge) to taxa apparently more related to A. hemibapha (described from Sri Lanka) than to the true A. caesarea (Scop.:Fr.) Pers. of Europe. The latter taxon has a subhymenium up to five inflated cells thick—as opposed to the one to three cell subhymenium thickness that, at present, I take to define “stirps Hemibapha.” A number of taxa included in this key have not yet been reviewed by me; and, undoubtedly there are errors. In addition to A. caesarea and its white variety, at least four additional, Mexican taxa with the thicker type of subhymenium have been omitted from the key in keeping with the delimitation of its scope. The omitted Mexican species include (1) an orange-capped species common in markets of the central volcanic region, Amanita basii Guzmán & Ramírez-Guillén; (2) A. laurae Guzmán & Ramírez-Guillén; (3) A. yema Guzmán & Ramírez-Guillén; and (4) a white species eaten by the Mountain Pima in the region of Nabogame, Chihuahua [Laferrière, 1991; Laferrière and Gilbertson, 1992]. The overall number of taxa with the thicker subhymenium similar to that of A. caesarea is not known.

There is a third group of robust, often colorful taxa of Amanita section Vaginatae that often have an annulate stipe and share many characters with the taxa in “stirps Hemibapha” and “stirps Caesarea.” These are taxa with subhymenia not dominated by inflated cells -- tentatively called stirps Calyptroderma. Amanita calyptroderma G. F. Atk. & Ballen [=A. lanei (Murrill) Sacc. & Trotter =A. calyptrata Peck non Lam.) is one species of this stirps that is common in the western U.S.A.; another is one of the undescribed entities known as “A. caesarea” in the SW states of the U.S.A. The number of taxa in this third group is not known.

The “slender Caesar's mushrooms” of the New World

How many members of “stirps Hemibapha” do we have in North and Central America? What are their ranges? We may not have the plenitude of taxa in this group that can be boasted by southeast Asia; but, nevertheless, the New World members of “stirps Hemibapha” form a strikingly beautiful part of our mycological patrimony. Let's indulge in some mycological “hemisphere chauvinism” for a few moments.

We are all familiar with the beautiful red Caesar's mushroom depicted in North American mushroom field guides. Mushroom book authors covering the fungi of eastern Canada, the eastern U.S.A., and eastern and central Mexico often include a striking illustration of A. jacksonii in their books. It is less well known that in this same region (and extending southward at least to Costa Rica) there occur a number of closely related species of “stirps Hemibapha” ranging from drab to showy in coloration and, in size, from the length of my index finger to two-thirds the length of my forearm. Among the largest of the species are A. arkansana Rosen (known from the southeastern U.S.A.) and Amanita sp. CR6, a beautiful species with an orange-brown cap and hints of olive in the stipe decorations, that will be described in the near future based on material from Honduras and Costa Rica.

There are several collections of a member of the stirps with a cap having a brownish disc and buffy cream margin. These mushrooms are referable to Amanita murrilliana Singer (see key, below). This entity gets the prize for the most retiring and unobtrusive coloring among known Western Hemisphere members of the stirps. It is also an oddity in that it is the only member of the stirps in which the base of the stipe is usually attached to the universal veil on its sides as well as at its very bottom.

Many mushroomers familiar with the oak-beech-hickory woods of the eastern half of the U.S. will have seen a small “caesarea” with the stature of A. jacksonii, but with a yellow to orange-brown cap instead of a red one. This is not simply a faded jacksonii, but a distinct species called Amanita sp. 16 in the key below. It is this species that I believe was illustrated by Peck's correspondent Mary Banning in one of her beautiful and idiosyncratic watercolors labeled “Agaricus caesareus.” It would be most appropriate to name the species for Miss Banning, and I hope to do so soon.

There is at least one more species in the stirps that I know only from collections brought to me from Florida by Aaron Norarevian (Glen Oaks, New York). This species has a yellow cap with a red umbo and is known only from sandy coastal plain habitat.

Will we find more of these interesting and colorful species in North and Central America? Quite possibly. There remains a considerable area that has not been explored by mycologists and ethnomycologists. If such species exist, they may be a part of the diet of indigenous peoples and could be first noted by scientists in markets. And how will we know if we have something new? We need a key to what is known. The following might serve as a rough start.

A key

The following key is very rough and is too frequently based on pileus coloration without regard to other macroscopic and microscopic characters. I hope this problem will be relieved in time.

Some more caveats: There are four groups of names or provisional names that may actually represent a single taxon each. These are to be found at key choices numbers 6, 7, 27, and 28. Hence, between 30 and 35 taxa are represented in this key. Additional material from India and Thailand (representing as many 5 additional taxa) has been seen by me, but not reviewed microscopically. In addition, there is work underway in several quarters that will undoubtedly clarify species concepts involved and the relations among the taxa. Z. L. Yang has just completed a thoroughly fascinating, detailed account of the amanitas of southwestern China in a Ph.D. thesis at Tübingen University. A number of relevant taxa are treated in this work (Yang, 1997). Dr. Alec Wood (Oatley, New South Wales, Australia) has recently published a paper describing several additional taxa close to A. egregia D. A. Reid (see below) from eastern Australia. In other words, this key will require revision soon.

Accurate knowledge of geographic ranges of the included taxa is not available; I have listed what information on range is known to me. A hollow bullet ( ° ) before a name indicates that I have personally reviewed material and that the spore measurements are mine.

In the sets of spore data, bold face Q is the average of the length/width ratios of all measured spores for a single specimen. When more than one specimen has been examined, a range of values of Q is presented. Q gives a feeling for the shape of the spores (Bas, 1969) observed as follows: 1.0 - 1.05, globose; 1.05 -1.15, subglobose; 1.15 - 1.30, broadly ellipsoid; 1.30 - 1.60, ellipsoid; 1.60 - 2.0, elongate; 2.0 - 3.0, cylindric; > 3.0, bacilliform.

1. Taxa having 95% of spores < or = 10 (-10.1) µm long.
  2. Taxa having Q < or = 1.40
    3.  Pileus dominated by fuscous, fuligineous, umbrinous, or yellowish brown (fading to shades of yellow) tones; known from eastern Asia.
      4.  Pileus dominated by fuscous, fuligineous, or umbrinous tones; described from Java (type); recorded from Malaya, Singapore, Borneo, at least sometimes with Quercus; Q = 1.15 - 1.2; spore length 7.0 - 9.9 µm.

1. Amanita similis Boedijn.
=Amanita hemibapha subsp. similis (Boedijn) Corner & Bas.

      4.  Pileus yellowish brown at first; disc fading to sordid yellow with margin orange-yellow; described from islands southwest of Japan; occurring with Quercus and Castanopsis; Q = 1.35; spore length 7.0 - 9.0 µm (but few spores measured for protologue).

2. Amanita rubromarginata Har. Takahashi.

    3. Pileus with red, orange, or yellow tones dominating.
      5. Pileus red to orangish red from center to margin.
        6. East Asian or Indian taxa with Q = 1.15 - 1.35; spore length (7.0-) 7.5 - 9.5 (-12.0) µm. Described from Russian far east, in Quercus forest. Known also from southwestern China, northern India, Japan, and Korea. Also occurring with conifers and other genera of the Fagaceae.

3. Amanita caesareoides Lyu. N. Vassilieva
=Amanita hemibapha subsp. hemibapha sensu auct. japon.
=Amanita caesarea sensu
A. Kumar et al.

        6. Pileus entirely red or orangish red at least at first fading throughout with age; described from Québec, Canada, recorded from throughout SE Canada and NE U.S.A., in deciduous forest including Quercus or in mixed forest (most often then with Pinus); Q = 1.25 - 1.37 (-1.40); spore length (7.0-) 7.8 - 9.8 (-12.1) µm.

4. ° Amanita jacksonii.
=Amanita umbonata Pomerleau non (Sumst.) Sartori & L. Maire.
=Amanita tullossii Guzmán & Ramírez-Guillén.

      5. Pileus orange or red over disc and otherwise yellow. [Possibly a single taxon.]
        7. Recorded from SW China, in forest dominated by Quercus and Rhododendron; Q = 1.20; spore length (6.0-) 7.0 - 9.0 (-9.5) µm.

5. Amanita hemibapha sensu Zhu L. Yang.

        7. Recorded from Himachal Pradesh, India, in mixed forest with Quercus incana and, sometimes, Rhododendron arboreum and Pinus wallichiana; Q = 1.14 - 1.25; spore length (7.2-) 8.0 - 9.5 (-11.5) µm.

6. ° Amanita sp. IHJ6.

  2. Taxa having Q > or = 1.4. [See also A. jacksonii, above.]
    8. Pileus nearly uniformly orange-tan to brownish orange, lacking red areas; known from NE Queensland, Australia; spores with Q = 1.53 - 1.54; spore length 8.5 - 10.2 µm.

7. ° Amanita sp. AUS3.

    8. Pileus yellow for outer half of radius, “beautiful scarlet” for inner half; described from Sri Lanka, with unknown symbiont(s); Q = 1.40 - 1.54; spore length (7.5-) 7.9 - 10.1 (-10.2) µm.

8. Amanita hemibapha subsp. hemibapha.

1. Taxa not having 95% of spores < or = 10 µm long.
  9. Taxa having 95% of spores < or = 12 µm long.
    10. Taxa having Q usually < 1.35
      11. Pileus not dominated by shades of yellow, orange or red.
        12. Pileus brown to grayish brown; Q = 1.04 ± 0.04; spore length (8.0-) 8.8 - 10.8 (-12.0); described from Japan in forests with Quercus and members of the Pinaceae.

9. Amanita imazekii T. Oda, C. Tanaka & Tsuda.

        12. Pileus white or largely white or whitish with disc taking on brown tones at maturity or with brown disc and pallid margin from the first.
          13. Pileus white at first, with disc taking on yellowish or brownish tones at maturity; described from Queensland, Australia, in open eucalypt forest; Q = 1.14 - 1.17; spore length (7.0-) 9.1 - 11.2 (-13.0) µm.

10. ° Amanita egregia D. A. Reid.

          13. Pileus brown over disc from the first, virgate between disc and very pallid to white region over striations; recorded from Thailand, in mixed dipterocarp forest; Q = 1.11 - 1.17; spore length (8.2-) 9.2 - 11.0 (-13.0) µm.

11. ° Amanita sp. Thai3.

      11. Pileus dominated by shades of yellow, orange, or red.
        15. Pileus with brownish and/or olivaceous tones.
          16. Pileus brownish yellow to brownish orange, darkest over disc; stipe with patches concolorous at first, becoming more orange when handled; described from Arkansas, U.S.A., re-corded throughout SE U.S.A., in frondose woods; Q = (1.22-) 1.24 - 1.38; spore length (7.0-) 7.7 - 10.5 (-15.0) µm.

12. ° Amanita arkansana.

          16. Pileus with olivaceous tones at least at first, with umbo reddish brown and margin brownish yellow; stipe having sordid yellow patches; recorded from Honduras and Costa Rica in association with Quercus; Q = 1.25 - 1.28; spore length (7.5-) 7.8 - 11.0 (-12.5) µm.

13. ° "Amanita garabitoana" Tulloss, Halling & G. M. Muell. nom. prov..

        15. Pileus with strong red and yellow tones, lacking brown tones.
          17. Pileus reddish with orange disc; described from Himachal Pradesh, India, in association with Cedrus deodara; Q = 1.25; spore length 7.0 - 11.5 µm.

14. Amanita simlensis R. P. Bhatt et al. in A. Kumar et al.

          17. Pileus yellow with brilliant reddish orange disc; recorded from Florida, U.S.A., in Pinus-Quercus forest; Q = 1.26; spore length 9.1 - 10.8 (-12.6) µm.

15. ° Amanita sp. F10.

    10. Taxa often having Q > 1.35. (See also, A. arkansana, above.)
      18. Pileus fuligineous to umbraceous to fuscous; recorded from Japan, with symbiont(s) unknown; Q = 1.4; spore length 7.5 - 10.5 µm. (See also, A. rubromarginata, above.)

16. Amanita hemibapha subsp. similis sensu auct. japon.

      18. Pileus with yellow, yellow-orange, orange, yellow-brown, or orange-brown, or warm brown tones.
        19. Pileus yellow-orange, darker and redder over disc; described from Tanzania, from miombo woodland; Q = 1.36; spore length 8.5 - 12 µm.

17. Amanita masasiensis Härk. & Saarim. in Härk., Saarim. & Mwasumbi.

        19. Pileus yellow to orange-yellow or ocher yellow to yellow developing brown tinted disc or warm brown at first becoming yellow-brown to orangish tan with age.
          20. Pileus yellow at first and becoming yellow-brown to orange-brown at least in part or warm brown at first and becoming paler orangish brown at maturity.
            21. Pileus yellow at first, becoming yellow-brown to orange-brown over disc; recorded from E U.S.A., in forests dominated by Fagus and Quercus or Pinus and Quercus; Q = 1.39 -1.60 (-1.69); spore length (7.5-) 8.4 - 11.9 (-15.0) µm.

18. ° Amanita sp. 16.

            21. Pileus warm brown at first, becoming paler orangish brown overall at maturity; recorded from NW Pakistan in mixed forest with Abies pindrow and Taxus wallichiana and in Nepalese markets; Q = 1.35 - 1.47; spore length (7.9-) 8.8 - 11.0 (-14.0) µm.

19. ° Amanita sp. PAK2.

          20. Pileus not entirely brown at first and disc not becoming brownish yellow or brownish orange with maturity.
            22. Pileus with red disc, otherwise yellow; recorded from Himachal Pradesh, India, in Shorea robusta forest; Q = 1.53; spore length (7.8-) 8.0 - 10.8 µm.

20. ° Amanita sp. IHJ4.

            22. Pileus lacking red disc.
              23. Pileus with brownish orange disc and yellow margin; known from NE Queensland, Australia; spores with Q = 1.44 - 1.49; spore length 8.6 - 11.2 µm.

21. ° Amanita sp. AUS2.

              23. Pileus lacking brownish orange disc.
                24. Pileus orange-yellow to ocher yellow, with yellow margin; described from Java, with symbiont(s) not reported; Q = 1.45; spore length 8 - 12 µm. (See also, Amanita hemibapha subsp. javanica sensu auct. japon., below.)

22. Amanita javanica (Corner & Bas) T. Oda, C. Tanaka & Tsuda.

                24. Pileus light cadmium to lemon chrome; recorded from China and Tibet, with symbiont(s) unknown; Q = 1.6; spore length 10 - 12 µm. (See also, Amanita hemibapha subsp. javanica sensu auct. japon., below.)

23. Amanita caesarea sensu Teng/Mao.

  9. Taxa not having 95% of spores < or = 12 µm long.
    25. Pileus entirely white (even at maturity) or with yellowish disc at maturity, described from Nepal in association with Shorea robusta, recorded from SW China with unknown symbiont(s); Q = 1.14-1.15; spore length (6.5-) 9.2 - 12.5 (-14.8) µm.

24. ° Amanita chepangiana Tulloss & Bhandary.

    25. Taxa usually having Q > or = 1.2.
      26. Pileus brown with white or pallid (not yellow) region over marginal striations or white to cream, sometimes with olivaceous brown to tan to buff to straw-colored disc.
        27. Pileus brown with paler region over marginal striations (0.3R - 0.5R); lamellae with pink tint, subdistant; described from Japan, with symbiont(s) unknown to me; Q = 1.22 ± 0.06 (std. dev.); spore length (9.5-) 10.0 - 12.5 (-13.5) µ. (Yang, 1997).

25. ° Amanita longistriata (S. Imai) E. J. Gilbert.

        27. Pileus white to cream, sometimes with olivaceous brown or tan to buff disc; with spores having Q in the range 1.24 - 1.41.
          28. African taxa; fruiting body robust; universal veil attached only at very base of stipe (like all other taxa treated in this key with the exceptions noted in the second half of this couplet). [Possibly a single taxon.]
            29. Pileus entirely white to pallid at first, then olivaceous brown over disc and progressively paler toward margin; described from Zambia, recorded from Zimbabwe and Tanzania, in miombo woodland and probably mycorrhizal with Brachystegia; Q = 1.24 - 1.41; spore length (10.0-) 10.5 - 13.8 (-21.0) µm.

26. ° Amanita zambiana Pegler & Piearce.

            29. Pileus bright white to cream, with fawn colored disc; Q = 1.3; spore length 13 - 14 µm; described from Congo, known from central Africa, with symbiont(s) not recorded.

27. Amanita loosii Beeli.

          28. Taxa of eastern North America and Asia; fruiting body rather gracile; pileus never developing olivaceous brown tint; universal veil attached 5 - 10 mm up the sides of the stipe, not only at very base of stipe.
            30. Asian taxon; pileus brown over disc, not virgate between disc and margin, gradually shading through pale tan to nearly white or cream at margin, with marginal striations 0.15R - 0.2R; recorded from Pakistan, associated with Abies pindrow; Q = 1.29 - 1.48; spore length (10.0-) 10.1 - 12.2 (-12.8) µm.

28. ° Amanita pakistanica Tulloss, S. H. Iqbal & A. N. Khalid.

            30. North American taxon; pileus with disc watery tan to straw color to pinkish buff, remainder cream to whitish or pale pinkish buff; described from Florida, U.S.A. with Quercus; also recorded from Michigan and North Carolina with Fagus and Quercus or isolated Quercus or with Tsuga, Betula, and Rhododenron; Q = 1.46 - 1.53; spore length (9.2-) 10.1 - 12.6 (-13.6) µm.

29. ° Amanita murrilliana

      26. Pileus orange-brown to red-brown or evenly yellow-orange or gray-brown or fuligineous to umbrinous or brown and (often) yellow toward margin.
        31. Pileus fuligineous to umbrinous or gray-brown or brown and (often) yellow toward margin.
          32. Pileus brown over disc and brown to yellow toward margin; recorded from China and Tibet, with Abies, Picea, Pinus, and Quercus; Q = 1.36 ±; spore length (8.0-) 9.0 - 12.5 (-17.0) µm.

30. Amanita hemibapha var. ochracea Zhu L. Yang.

          32. Pileus fuliginous to umbrinous to gray-brown.
            33. Pileus fuliginous to umbrinous; described from Japan, in Pinus forest; Q = 1.6; spore length 10.5 - 14 µm.

31. Amanita esculenta Hongo & I. Matsuda.

            33. Pileus dark gray-brown, paler toward margin; described from Hunan, China, in Pinus forest; Q = 1.47±0.01; spore length (8.0-) 9.5 - 12.5 (-15.5) µm per Yang and Zhang (2002: 718).

32. Amanita hunanensis Y. B. Peng & L. H. Liu.

        31. Pileus orange-brown to red-brown or evenly yellow-orange.
          34. Pileus orange-brown to red-brown, often with darker ring over inner ends of striations; described from Tanzania, in miombo woodland; Q = 1.44 - 1.66; spore length 9.5 - 13 µm.

33. Amanita mafingensis Härk. & Saarim. in Härk., Saarim. & Mwasumbi.

          34. Pileus evenly yellow-orange.
            35. Pileus lacking an umbo; described from Tanzania, in miombo woodland; Q = 1.58; spore length 8.5 - 12.5 µm.

34. Amanita tanzanica Härk. & Saarim. in Härk., Saarim. & Mwasumbi.

            35. Pileus with umbo; recorded from Japan, in forest with Pinus densiflora and Quercus serrata; Q = 1.47; spore length (9.2-) 10.0 - 13.0 (-16.0) µm.

37. ° Amanita hemibapha subsp. javanica sensu auct. japon.

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