Papaver at High Altitudes in the Rocky Mountains

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<ul><li><p>PAPAVER AT HIGH ALTITUDES IN THE ROCKY MOUNTAINS </p><p>DORIS LOVE </p><p>LSve, Doris (Institute of Arctic and Alpine Research, University of Colorado, Boulder). Papaver at high altitudes in the Rocky Mountains. Brittonia 21: 1-10. 1969. In the Rocky Mountains south of the Yukon Territory there are two, possibly three, Papaver species at alpine elevations. The diploid P. pygmaeum Rydb. is found only in and around the Glacier-Waterton International Park. It is a member of the alpinum complex of sect. Scapiflora and linked to the European P. alpinum L. by a chain of taxa through inner Asia. The hexaploid P. kluanense D. LSve is an alpine representative of the circumpolar radicatum complex of the same section and occurs disjunct from Alaska and Yukon south to New Mexico. In the Uintah Mountains of Utah is a small population of Papaver whose affiliation and rank within the section Scapi/lora is not yet established. It seems to be related most closely to P. pulvinatum A. Tolm. and P. alaskanum Hult~n. A key to the taxa of this region is presented. </p><p>INTRODUCTION </p><p>In contrast to the richness and variety of Papaver sect. Scapi]lora in the mountains of Eurasia and in the circumpolar Arctic, this genus and section is represented by only a few species in the alpine areas of the North American Rocky Mountains. It seems that south of the Yukon Territory only two, or possibly three, high altitude species of poppies occur. The first belongs to a very rare, pink-flowered species, Papaver pygmaeum Rydb.; the second to a yellow-flowered and not so rare taxon, P. kluanensis D. LSve; whereas the third is a population of yellow-flowered poppies in the Uintah Mountains of Utah of doubtful relationship (Fig. 1). </p><p>The areas of the first two species consist of widely separated populations at high altitude, always above timberline. P. pygmaeum is practically confined to the Glacier- Waterton International Park and its immediate surroundings on the border between southern British Columbia and Alberta in Canada and Montana in the United States (Fig. 2). P. kluanensis, on the other hand, extends as isolated populations along the lofty peaks of the Rocky Mountains from Alaska and Yukon south to the Taos Mountains in the Sangre de Cristo Range in northern New Mexico (Fig. 1). </p><p>PAPAVER PYGMAEUM Rydb. The name Papaver pygmaeum was given by Rydberg (1902) to specimens taken </p><p>by R. S. Williams (no. 992) on a mountain above Stanton Lake in Montana on August 1, 1894. In his description of the new species, Rydberg states: "This species is nearly related to P. radicatum Rottb., but is a still smaller plant with shorter, broader, less divided, less bristly leaf blades and smaller flowers. I t resembles still more the European P. pyrenaicum which has larger petals, 1-2 cm. long, stamens much exceeding the ovary and sphaerical flowerbuds." </p><p>Williams' specimens are, however, not the oldest known of this kind of poppy, nor had the resemblance to P. pyrenaicum passed unobserved earlier. Studying what is preserved in herbaria of P. pygmaeum and correspondence relating to the collections, I have been able to trace its interesting history. </p><p>It seems that the first specimens of this taxon were collected in the Canadian Rocky Mountains as early as July 31, 1881, by J. M. Macoun of the Canadian Geological Survey, on Sheep Mountain near Waterton Lake at an altitude of 7,500 ft. The latitude is given as 49o05 ' N. About two weeks later, on August 15, 1881, </p><p>BRITTONIA 21 : 1--10. Jam-Mar . 1969. </p></li><li><p>2 BRITTONIA [VOL 21 </p><p>" ..... . -O'''NT aTON LAKES l _ _ { . NAT IONAL PARK </p><p>ALTA. e*" *2,.j - - . \\x I </p><p>~'. ~. \ </p><p>~ ....~,. .... .O ,, </p><p>GLACIER </p><p>NAT I ON~L </p><p>PARK 9 ~ I ) , </p><p>J </p><p>"'~/ t ( </p><p>MONTANA </p><p>I </p><p>' , , . , ,~ I x \ </p><p>0 , . . , , , \ , , , , ,"! \ </p><p>,.. \ \ </p><p>9 \, "~, ":.": / </p><p>I " l </p><p>1 2 </p><p>Fro. 1. Distribution of Papaver kluanensis D. L Sve (dots) south of the Yukon Territory, and of Papaver sp. from Utah (triangles). </p><p>FIG. 2. Distribution of Papaver pygmaeum Rydb. </p><p>a second collection was made at S. Kootanic Pass, Rocky Mountains, by G. M. Dawson, also of the Canadian Geological Survey. Dawson's specimen is labelled Papaver nudicaule L., but the word var.? has been added to it in a different handwriting, at a later date. The label on Macoun's sheet is more interesting, since there appears in Macoun's hand the name Papaver dawsonii J. M. Macoun, but the epithet dawsonii has been crossed out and under it, in a different hand, stand the words p ygmaeum Rydb. Both of these collections are in the herbarium of the National Museum of Canada. </p><p>It is apparent from some letters, labels, and notes which are affixed to the old specimens in the herbarium of the New York Botanical Garden, that J. M. Macoun, N. L. Britton, P. A. Rydberg, and T. Holm had some differences of opinion as to the identity of these plants. Holm identified them with P. pyrenaicum, whereas Rydberg felt it safer to describe them as the new species P. pygmaeum. P. pyre- naicum (L.) A. Kerner is a Pyrenean and Alpique representative of the P. alpinum complex of Central and South' European mountains, as defined by Fedde (1936) and Faberg(~ (1943). As all other diploid taxa of this group from that region, it is presently regarded as a subspecies only: ssp. rhaeticum (Let.) Mgf., of P. alpinum L. (Markgraf, 1958a,b, 1963), since Fabergfi (1943, 1944) demonstrated that the morphologically and geographically distinct taxa of this group lack all reproductive barriers. </p><p>When comparing specimens of the American P. pygrnaeum with representatives of the European P. alpinum complex, it is evident that these are closely related and that the American taxon belongs to this group, which' is best characterized by rigid, more or less appressed, ivory-colored, bulbous-based bristles on the capsules (Fig. 3). This character shows no variation and is always observable, even on very young and immature capsules. In the radicatum complex the capsules are hairy, but </p></li><li><p>1969] LSVE: PAPAVER 3 </p><p>FIG. 3. Capsules and hairiness: A. Papaver pygmaeum Rydb.; B. P. microcarpum DC.; C .P . alboroseum Hult~n; D. bristle typical of the alpinum group; E. hair typical of the radicatum and nudica~le groups; F. P. kluanensis D. LSve; G. Papaver sp. from Utah; H. P. radicatum Rottb. ssp. occidentale Lundstr.; I .P. keelei Porsild. A-C, F- I X 3; D, E X 10. </p></li><li><p>4 BRITTONIA [VOL. 21 </p><p>although some species have hairs which are stiff with broad bases, they are certainly not bulbous-based bristles, nor do they have the characteristic pale color. Instead they usually vary from pale brown to an ebony black (Fig. 3). In the nudicaule complex the capsules vary from glabrous to very hairy, and the hairs are sometimes paler brown than in the radicatum complex, but though they may be quite stiff and have broad bases, they are never bristle-like nor do they have bulbous bases (Fig. 3). Thus, bulbous-based, ivory-colored bristles on the capsules can be considered a key character for the alpinum complex of the Scapi]lora section. It should be remarked, however, that bulbous-based bristles are also known in species from other sections of Papaver. Other characters which unite the alpinum complex are more variable and harder to use as distinctive from similar characters of the other complexes. </p><p>As far as is known, the chromosome number typical of the alpinum complex is 2n = 14, with the possible exception of P. albo-roseum (2n = 28) if it belongs here (cf. below). The nudicaule complex is known to have both diploid and polyploid numbers (2n = 14, 28, and 42), while the radicatum complex is highly polyploid (2n = 28, 42, 56, 70, and 84). It must be admitted, however, that with the exception of the radicatum complex relatively few Scapi]lora species are cytologically known (cf. Ljungdahl, 1922, 1924; Sugiura, 1936a,b, 1937, 1940; Horn, 1938; Faberg6, 1942, 1943, 1944; *. L6ve, 1955, 1962a,b; Li~ve &amp; Freedman, 1956; Asahina et al., 1957; Knaben, 1959a,b; Sokolovskaja &amp; Strelkova, 1960; L/Sve &amp; L/~ve, 1961; Kawatani &amp; Ohno, 1965; Zhukowa, 1965, 1968; Hedberg, 1967; Kiipfer &amp; Favarger, 1967; Mosquin &amp; Hayley, 1966; Sugnik, 1967; Packer, 1968). </p><p>The chromosome number of P. pygmaeum from the Piegan Pass of the Glacier National Park in Montana (collected by Dr. M. Bowers, COLO.) is 2n -= 14, or the same as reported by Packer (1968) from Waterton National Park in Alberta. This agrees with what is known from the alpinum complex in Europe (cf. LiSve &amp; L6ve, 1961; Kiipfer &amp; Favarger, 1967; Sugnik, 1967). It is evident, therefore, that P. pygmaeum is not only morphologically but also cytologically very similar to the geographically remote European P. alpinum group, as originally claimed by Holm. </p><p>When looking through collections of Papaver in American herbaria, I have found a number of specimens from North America and Asia that are given various names but fit very well into the general type of the alpinum complex. The taxa in question apparently form a chain of disjunct links from the European P. alpinum aggregate to the North American P. pygmaeum through the mountains of inner Asia. Specimens which I have seen and found to be characterized by bulbous-based bristles on the capsules and a general resemblance to the P. alpinum complex, carry several species names: P. walpolei Porsild, from Seward Peninsula in Alaska and Mys Deshevna in Siberia; P. microcarpum DC., from Seward and Kenai Peninsulas in Alaska, the Aleutian Islands, and Kamtchatka; P. ]auriei Fedde, from Rishiri Island near Hokkaido; and P. nudicaule L. ssp. baicalense Tolm., from the Olchon Island in Lake Baical. DeCandolle (1824) noted the similarities of P. microcarpum to P. pyrenaicum and place the new species between this taxon and P. nudicaule with the remark that it has smaller capsules than any of these species. Hult6n (1945) draws attention to the likeness between P. walpolei on one hand and P. pyrenaicum and P. pygmaeum on the other, and also to the fact that P. walpolei is distinct from other poppies in Alaska which belong either to the radicatum or nudicaule complexes. From studies of descriptions and illustrations in the literature available (Popov, 1937; Tolmachev, 1931a,b, 1932, 1953, 1960) I have come to the conclusion that the follow- ing species may perhaps also be placed in the alpinum complex: P. rubroaurantiacum (DC.) Fisch. ex Lundstr. (Dauria, Mongolia), P. ledebourianum Lundstr. (Dauria, Mongolia), P. ajanense M. Pop. (Far East Siberia, Udsk), P. involucratum M. Pop. </p></li><li><p>1969] LSVE: PAPAVER </p><p>(Central Asia, Pamir-Altai), P. ti~nschanicum M. Pop. (Altai, Tian-Shan, Dzhunga- rian Alatau), P. canescens Tolm. (Dzhungaria, Tarbaratai Mts.), P. pseudocanescens M. Pop. (Altai, Angan-Sayan, Mongolia), which are all mentioned in the Flora SSSR, and P. nudicaule L. ssp. gracile Tolm. (Central Yakutsk, cf. Tolmachev, 1960). All of these taxa are characterized by having small flowers, broad-segmented leaves, and a very variable hairiness. They all have coarse, strong bristles on the small capsules. </p><p>It seems safe to conclude from the numerous genetical experiments which have been made with populations of Papaver sect. Scapiflora that speciation within these complexes has mainly been abrupt, caused by differences in chromosome number, whereas differentiation within each ploidy group has presumably been subspecific and only rarely has involved gradual differentiation of the chromosomes sufficient to create reproductive barriers. This is especially evident from the genetical investi- gations by Faberg6 (1943, 1944) on the diploid alpinurn group, but also by the extensive studies by Knaben (1959a,b) on the octoploids and decaploids of the radicatum group. Therefore, a classification of each ploidy group into subspecific rather than specific units seems to be well conceived (L6ve &amp; LSve, 1961; A. LSve, 1962a,b). Because of our knowledge of the lack of reproductive barriers within each ploidy group so far studied, and because of the close morphological resemblance of the Asiatic and American taxa to the European diploids, it is tempting to regard them all as subspecies only of P. alpinum, equivalent to its several races so named from the European mountains by Markgraf (1958a,b, 19'63). In the case of the American taxon, all available facts support such a treatment. Such a conclusion ought not to be made, however, before closer studies have revealed their morphological re- lationships and the cytological characteristics of their hybrids. Thus, I prefer to delay the transfer of the American taxon and the Asiatic taxa to subspecific status of the European plant until hybridization experiments have demonstrated a lack of reproduc- tive isolation between them, since it is "a lesser evil to keep forms separated that are identical than to identify such' as are distinct" (Halle, 1913). </p><p>P. alboroseum Hult6n from Kamtchatka (Hult6n, 1928) also seems to belong to the alpinum group from a purely morphological point of view. Specimens in the herbarium of the Canada Department of Agriculture in Ottawa, collected on the Kenai Peninsula in Alaska, have the characteristic ivory-colored, bulbous-based bristles on the capsules, although the capsules are bigger but of the same shape (Fig. 3). Since it has 2n = 28 chromosomes (Knaben, 1959a), it cannot be con- specific with alpinum. If it really belongs to this complex, it would constitute the first known case of polyploidy within that group. </p><p>It is very likely that a more detailed study of the Asiatic ScapiJlora section of Papaver, with the characteristics of the alpinum complex in mind, would lead to a distinction between a nudicaule, an alpinum, and a radicatum complex also in this area, where until now it has been assumed that only the nudicaule and the more arctic radicatum complexes existed. If this can be shown to be correct, then the occurrence of P. pygmaeum, distinctly of the alpinum complex, in the Rocky Mountains, would no longer seem surprising since it would merely be an outpost for an alpine chain of the P. alpinum aggregate, a phenomenon which is known also from several other species that range from southern European mountains to the Rockies. </p><p>PAPAVER KLUANENSIS D. L6ve. </p><p>Fedde (1909b), in his monograph of Papaveraceae, included a number of taxa of Papaver sect. ScapiJlora from the Rocky Mountains, i.e., P. nudicaule L. ssp. radicatum (Rottb.) Fedde var. columbianum Fedde, var. pseudocorydali/olium Fedde, </p></li><li><p>6 BRITTONIA [VOL. 21 </p><p>and vat. coloradense Fedde. These varieties were originally described elsewhere (Fedde, 1909a, pp. 255-256), and type specimens are deposited in the Herbarium of the Smithsonian Institution in Washington, from where I have had them on loan. </p><p>I t was a surprise to discover that the holotype for Fedde's vat. pseudocorydali]olium is none other than Macoun's sheet of 1895 from Sheep Mountain near Waterton Lake in southern Alberta, with the name P. pyrenaicum on the original label and P. pygmaeum written in Rydberg's hand above it. I t is not known if the...</p></li></ul>