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A Preliminary Account of the North American Species of Rhizopogon

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A Preliminary Account of the North American Species of Rhizopogon MEMOIRS OF The New York Botanical Garden VOLUME 14, NUMBER 2 A Preliminary Account of the North American Species of Rhizopogon ALEXANDER H. SMITH AND S. M. ZELLER March, 1966 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 14(2):1-178 A PRELIMINARY ACCOUNT OF THE NORTH AMERICAN SPECIES OF RHIZOPOGONI ALEXANDER H. SMITH 2 AND S. M. ZELLER3 PREFACE BY ALEXANDER H. SMITH The genus Rhizopogon) a member of the Hymenogastraceae, is one of the largest and most diverse groups of the so-called hypogeous fungi. My own interest in it developed as its members were encountered in the conifer forests of our western states over the years since 1935, when I made my first trip to that area. However, it was not until many years after Dr. Zeller's death that I actually made an effort to collect and study the species critically, partly as a result of becoming interested at the research level in the problem of the origin of the Hymenomycetes with particular reference to secotiaceous fungi. The history of the study of Rhizopogon is as spotty as the collection of its species. As for all groups of higher fungi it dates roughly from the time of Fries (1817) . The treatment of Rhizopogon by the Tulasnes (1851) may be said to have clearly established the genus, but even to this day in Europe there has been no major revision of the genus along the lines of the modern approach to the taxonomy of the Hymenomycetes. In North America the monograph by Zeller and Dodge (1918) has been the outstanding contribution, although the treatment by Coker and Couch (1928) added much to clarity of species concepts regardless of the names used. A study of their approach furnished many of the ideas elab­ orated upon in the present work. In the European literature the works of Hawker (1954, 1955) and Morten Lange (1956, 1957) represent significant contributions. No systematic attempt has been made at this stage of my investigation to verify reports in the literature on Rhizopogon species as these have been used in mycorrhizal studies or reported in lists in North America. In one case, (Baxter, 1929) , the species involved turned out to be new though identified by Zeller as R. rubescens. The present approach to Rhizopogon is that currently in use for the Hymen­ omycetes. In other words I have tried to correlate the macroscopic features with the microscopic characters and have in addition included two of the more reali­ able chemical tests (contact with iron sulphate and with 2.5% potassium hy­ droxide, herein referred to as FeS04 and KOH respectively) along with the iodine reactions of the spores and tissues. After three years of studying fresh material in the Salmon River drainage of Idaho, such a large number of taxa had been encountered and studied that I felt rather hopelessly lost. It was seldom possible to make a satisfactory identification of any of this material in the existing literature. It was in this period that I was approached by The New York Botan- 'Publication of this monograph was financed under National Science Foundation Grant 6N-348. "University of Michigan, Ann Arbor, Michigan. "Deceased 4 November 1948. 2 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 14 ical Garden to finish the work started by Zeller on the Gastromycetes of North America. This seemed like a good opportunity to settle the questions in regard to Rhizopogon. However, things are not always what they seem, and after working through Zeller's collections I found relatively few of my Idaho species. In addi­ tion to this, other disconcerting problems raised their ugly heads. The first point to be clarified was that Zeller and Dodge (1918) and later Zeller (1939, 1941) had described many truly unusual species from the Pacific Northwest. These for the most part were easily recognizable once a careful study of the types had been made. However, when one approached the problem of the recognition of Euro­ pean species in the North American Rhizopogon flora chaos reigned supreme. American collections were frequently identified as European species even though such simple characters as spore size were completely out of line. In searching for an approach to more accurate and meaningful species concepts in this genus it at once became evident that if scientific accuracy was to have any priority at all it could be obtained only by describing the American species on the basis of their own features, and by not trying to "force" collections into old or indefinite con­ cepts originating in Europe and quite obviously not applying well to American material. A good house cannot be built with poor materials. Since European authors had not used the same characters r am using, meaningful comparisons for the most part are unobtainable at the present time. Rhizopogon species are generally so rare and difficult to find that one cannot as a rule go into an area and get a fair representation of the flora in a single season or even several. In a life-time one investigator cannot hope to more than collect the Rhizopogon flora of a single continent. The best procedure, in my estimation, is for investigators on different continents to work independently but use the same set of characters, so that their findings are directly comparable to those of other workers. It is only in this way that meaningful distribution pat­ terns can be worked out. As of the present writing I do not consider reports of the distribution of such common species as R. yubescens and R. roseolus to be at all reliable-true, some of them are correct-but this must be proven on the basis of more accurate species concepts than have existed to date. I am not being at all facetious in entitling this a preliminary contribution toward a monograph. It is an attempt to establish concepts upon which significant distribution data may be accumulated. This is a very important point because it now appears that many species of Rhizopogon are highly restricted in their distribution. In a sense this work is also preliminary because of the introduction of a different set of char­ acters than that used previously in the genus. The resulting concepts, like all new concepts, need testing by other workers than the one proposing them to see if they stand the test of objectivity. It is always easy to point out the short-comings of the classical characters used in a previous classification, but it is not always easy to introduce a better and different set. The shortest road to the evaluation of a new set of characters is to set up a classification based on them and then let your colleagues try to find fault with it. In short, this is what I am doing here. My outline (1964) of the infrageneric classification was published separately because Zeller had never used such features as the amyloid character for spores and tissues or the KOH and FeS04 reactions. I assume sale responsibility for the classifica­ tion, and any shortcomings which may come to light are to be laid at my door. As previously mentioned, the interesting feature to come out of this study so far is that the flora Zeller was working on in the coastal area is apparently quite distinct from that of the Salmon River drainage in Idaho. The two areas taken together, however, certainly represent the Rhizopogon center of the world. The number of previously undescribed species became particularly embarrassing, but 1966] NORTH AMERICAN SPECIES OF RHIZOPOGON 3 it was a choice between using the combinations of characters that were constant or simply ignoring them. The latter course has never contributed to a better understanding of species in any group. My greatest disappointment in this investigation was an inability to satis­ factorily identify dried specimens in herbaria where notes on the fresh characters were lacking. I know of many new species which are not included here simply because essential data are lacking. Actually dried specimens as found in herbaria present a most discouraging problem to an investigator. In the case of old coIlec­ tions (made more than 30 years ago) one does not know whether the specimens were first preserved in liquid and then dried (which makes color reactions ob­ tained from them by the application of chemicals subject to possible error) , or dried first and then soaked in some poison to keep out the insects. In either case the consistency of the specimen and reaction of chemicals on them could have been substantially altered. One does not know the drying techniques used-and this can make a tremendous difference in the consistency and appearance of the specimens. With these uncertainties in mind and a lack of notes on the features of the specimens when fresh, in most cases one simply cannot be sure as to the identity of a given coIlection. On the other hand if the color of young, mature, and old basidiocarps is recorded even in general terms, if color changes which take place when a speci­ men is injured are noted, along with distinctive odors, the color of the young, mature and old gleba, and the KOH as weIl as the FeS04 reactions of the peridium, then the specimens properly dried, such material is extremely valuable. The microscopic features can be obtained from dried material as can certain color reactions. The best drying technique is that of Hoseney (1963) because it is easy, safe and can be carried on in the home without the bother of a lot of debris. Specimens prepared by her technique are a valuable addition to any herbarium.
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