ART KRUCKEBERG DOES A TURKEY TROT

Our month in Turkey this past summer, courtesy the National Science Foundation focussed on the many serpentine outcrops there. We were especially keen on finding serpentine plants that accumulate high levels of nickel (tissue with >1000 ppm Ni, called hyperaccumulators). Serpentine exposures near Ankara and in the lofty Taurus Mountains to the south of Ankara are rich in narrow endemics; most species of Alyssum (Brassicaceae) proved to be hyperaccumulators, via our simple field test for nickel [see BEN # 143]. Our field party, Dr. Roger Reeves of Massey University, New Zealand, three Turkish botanists, and ARK, did find time to enjoy Turkish beer, their ever-present watermelons and Turkish tea and coffee. We took time off from plant hunting to visit the amazing tuff formations in Cappadocia and the ancient subterranean "cities" dug into the volcanic tuff. Turkish flora, geology, landscapes and the friendly people all were delightful.

Bill Weber, my friend, you would have revelled in the geobotanical richness of Turkey, especially the subalpine/alpine flora of the Taurus Mountains...the Rockies of Turkey. I'll bet Turkey has even more locoweeds than Colorado!

SOME NOTES ON PAPAVER

I have just returned from Oslo where I spent five weeks at the Center for Advanced Study of the Norwegian Academy. Norwegian colleagues Inger Nordal and Reidar Elven, University of Oslo, had agreed to take on leadership of the Panarctic Flora Project, and they obtained funds to support a Panarctic Year at the Center. The year was kicked off in late September with a small symposium to discuss the species concept. Since our proximate goal is to complete a checklist of arctic vascular plants, it is imperative we reconcile the different taxonomic traditions of participating countries and create a panarctic synthesis.

After the symposium and throughout the rest of the year, working groups will be assembled in Oslo to work intensively on certain genera. Papaver is one of those. Following the symposium we looked at the scapose taxa of what most people still call the Scapiflorae (section Meconella). To accomplish the panarctic view, we looked together at specimens, including types, with key references close at hand. Important questions were: are we recognizing the same taxa; are we applying the same names, and if not why not?

The "we" of the poppy working group consisted of the following. Inger Nordal and Reidar Elven, and Heidi Solstad who had just completed an M.Sc. with them, brought to the table a lot of experience and recent research on breeding systems and isozyme variation for the North Atlantic region. Vladik Petrovsky had worked with I. A. Tolmachev on the treatment of poppies in Arctic Flora USSR and had himself named new taxa from northeastern Asia. Moreover, with his wife, Paulina Zhukova, he had reported chromosome numbers, important reference points, for many of those taxa. Orjan Nilsson, who has completed a treatment of poppies for Flora Nordica, provided another Scandinavian perspective. I was there to represent the North American experience gained during my struggle to produce the FNA treatment. What I am reporting here draws from our discussions.

To be certain we were all on the same page, as the saying goes, we first developed a character list, which was an interesting exercise. After some testing and refinements, this will be come the basis for a morphological reassessment of taxa with our individual biases removed.

In my contribution to the treatment of Papaver with Bob Kiger in Flora of North America, some points were clearly left unresolved. One was the question of what to call the plant of the high arctic latitudes that I had named P. radicatum subsp. polare. The similarity of our plants, and those of arctic Russia, to the Svalbard P. dahlianum is clear. What was ambiguous was the relationship between the plants on Svalbard and the type specimen for name P. dahlianum from Finmark, northern Norway. Due to my very limited experience with the Norwegian plants, I was unsure the Finmark and Svalbard plants were really the same taxon. Now, it is clear from what I saw in Oslo they are indeed the same and P. dahlianum is the name we should apply to our plants. Having answered that question, another was raised. Does P. gorodkovii belong with P. dahlianum as well?

There is still the question of what is P. radicatum? The technical problems of typification will be "solved" by others shortly. The view from the North Atlantic has been rather narrow, reflecting the legacy of Gunvor Knaben, who applied the name only to plants of Iceland, Norway, and Svalbard with a chromosome number of 2n=70. All plants having 2n=56 she called P. lapponicum. Yet, it has been clear to us in the North American that all our taxa simply cannot not be lumped in that species. There is P. lapponicum and another complex of forms that I placed in P. radicatum. It is obvious that species of Papaver cannot be defined by one chromosome number; indeed even two or three ploidy levels are reported for some morphological geographical entities. So, either we expand our concept of P. radicatum to include the 2n=42 and 56 chromosome taxa or we have to find another name for our material.

Knaben's work at the Botanical Garden in Oslo demonstrated that species of poppy could be crossed, despite big differences in ploidy level, and we must conclude that hybridization plays a role in creating the myriad forms one sees in a large series of specimens. On the other hand, Inger Nordal and her students have now shown that poppies are not only self-compatible but also have flowers structured in such ways as to make them essentially autogamous, at least early in the season, before the flowers open. She has, therefore, supplied an explanation for the pattern of variability we see--more variation among populations than within them. Occasional hybridization produces new, local variation , and these variants are maintained by selfing. Therefore our species concept and circumscriptions must take the breeding system into account.

We are establishing a living collection of poppies at the University of Oslo. Reidar Elven will be participating in the Swedish arctic expedition --Tundra Northwest 1999--this summer and will collect material from the Canadian Arctic to supplement what they already have from Svalbard and Finmark and what I have sent from Alaska.

BRACHYTHECIUM CALCAREUM KINDB. IN THE INTERIOR PACIFIC NORTHWEST

In the past several years I have been collecting and comparing Brachythecium samples wherever I go in the hopes of gaining a picture of this difficult genus. Of all of the mosses, the genus Brachythecium may be the most commonly encountered in the interior. In almost every forest stand, every streambed and seep, every roadside and every alpine meadow, one encounters Brachythecium, and oftentimes two or three species. In doing forest vegetation plots, I am constantly faced with the question of what species I have, and in some cases I find six species in a single 100 m square plot. Nonetheless, the experience has been enlightening, and morphological and autecological patterns have become quite clear.

One of the more significant discoveries in the course of the Brachythecium study centres around a julaceous, straw-coloured species which grows in ditches along country roads and highways, on disturbed soil in forests, and in old cutblocks. Its occurrence is almost predictable in these habitats and once known is easy to recognize. My early attempts at keying the moss put it in Brachythecium salebrosum, though when sterile, it seemed to key to B. albicans, and this is the name that the material seems to carry in herbaria. Two summers ago a German phytosociologist I was working with, Hans-Georg Stroh, pointed out to me that this moss, which I had come to call B. albicans, did not at all look like B. albicans as he knew it in Europe. This sparked my interest, and I compared the moss closely with descriptions of B. albicans in European floras. It became difficult to fit our plants into the concept of B. albicans. In particular, our material is autoicous, and fruits often, B. albicans is dioicous, and fruits rarely. Our material has weakly decurrent leaf margins, B. albicans has strongly decurrent leaf margins. In these and other finer points, our plants were something quite different than B. albicans, and this was reinforced when I found material in dry forests which actually fit the description of B. albicans very nicely.

Perusal of Robinson's (1962) key to North American Brachythecium placed the material very tidily in Brachythecium calcareum, a species described by in 1895 by Kindberg from Ontario, and subsequently reported from Newfoundland and Alaska (Crum & Anderson 1981) and Colorado (Weber & Wittmann 1992). Unsure, I sent a piece to Robinson, who confirmed my suspicion. New to Montana and the Pacific Northwest, I was subsequently able to find the species in its predictable roadside habitat in Idaho, Washington and British Columbia. It has become evident that Brachythecium calcareum is one of the most common ruderal species of the genus in the interior Pacific Northwest, and may, in fact, have its primary distribution here.

Selected specimens (collection numbers are those of the author and are housed in the Forest Service Fortine District Herbarium in Fortine, Montana, unless otherwise indicated): CANADA, BRITISH COLUMBIA. Southern interior, Creston Valley, West Creston Road, 7714; U.S.A., IDAHO. Bonner Co., Priest River region, Hager Lake, 6668; MONTANA. Flathead Co., Columbia Falls, North Fork Road, 1608 (US, !Robinson); Lake Co., Swan Valley, near Point Pleasant Campground, 6625; Lincoln Co., Murphy Lake, 6583; Sanders Co., Vermilion River at Bear Creek, 7516-A; WASHINGTON. Pend Oreille Co., Frater Lake, 7680 (WTU).

References:

Crum, H.A. & L.E. Anderson 1981.

Mosses of Eastern North America. Columbia University Press, New York.

Robinson, H. 1962.

Generic revisions of North American Brachytheciaceae. Bryologist 65(1): 73-145.

Weber, W.A. & R.C. Wittmann. 1992.

Catalog of the Colorado Flora: a biodiversity baseline. University Press of Colorado Press, Boulder.