papaver at high altitudes in the rocky mountains

10
PAPAVER AT HIGH ALTITUDES IN THE ROCKY MOUNTAINS DORIS LOVE 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. INTRODUCTION 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). 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). PAPAVER PYGMAEUM Rydb. The name Papaver pygmaeum was given by Rydberg (1902) to specimens taken 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. It resembles still more the European P. pyrenaicum which has larger petals, 1-2 cm. long, stamens much exceeding the ovary and sphaerical flowerbuds." 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. 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, BRITTONIA 21: 1--10. Jam-Mar. 1969.

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Page 1: Papaver at High Altitudes in the Rocky Mountains

PAPAVER AT HIGH ALTITUDES IN THE ROCKY MOUNTAINS

DORIS LOVE

LSve, Doris (Inst i tute of Arctic and Alpine Research, Universi ty of Colorado, Boulder). Papaver at high alti tudes in the Rocky Mountains . Bri t tonia 2 1 : 1-10. 1969. In the Rocky Mounta ins south of the Yukon Terri tory there are two, possibly three, Papaver species a t alpine elevations. The diploid P. pygmaeum Rydb. is found only in and around the Glacier-Waterton Internat ional 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 f rom Alaska and Yukon south to New Mexico. In the Uintah Mounta ins of Utah is a small population of Papaver whose affiliation and rank within the section Scapi/lora is not yet established. I t seems to be related mos t closely to P. pulvinatum A. Tolm. and P. alaskanum Hult~n. A key to the taxa of this region is presented.

INTRODUCTION

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. I t 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).

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).

PAPAVER PYGMAEUM Rydb. The name Papaver pygmaeum was given by Rydberg (1902) to specimens taken

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."

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.

I t 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,

BRITTONIA 2 1 : 1--10. J a m - M a r . 1969.

Page 2: Papaver at High Altitudes in the Rocky Mountains

2 BRITTONIA [VOL 21

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Fro. 1. Distribution of Papaver kluanensis D. L Sve (dots) south of the Yukon Territory, and of Papaver sp. from Utah (triangles).

FIG. 2. Distribution of Papaver pygmaeum Rydb.

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.

I t 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.

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

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1969] LSVE: PAPAVER 3

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.

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4 BRITTONIA [VOL. 21

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.

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 & Freedman, 1956; Asahina et al., 1957; Knaben, 1959a,b; Sokolovskaja & Strelkova, 1960; L/Sve & L/~ve, 1961; Kawatani & Ohno, 1965; Zhukowa, 1965, 1968; Hedberg, 1967; Kiipfer & Favarger, 1967; Mosquin & Hayley, 1966; Sugnik, 1967; Packer, 1968).

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 & L6ve, 1961; Kiipfer & Favarger, 1967; Sugnik, 1967). I t 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.

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.

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(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.

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 & 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. 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.

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.

PAPAVER KLUANENSIS D. L6ve.

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,

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6 BRITTONIA [VOL. 21

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.

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 specimens at US had been studied by Fedde before 1902, when Rydberg's paper was published, but Fedde (1909a,b) does not mention Rydberg's work. The specimens are, however, clearly those of P. pygmaeum Rydb. and were actually cited by Rydberg in his original description. Fedde (1909b) remarks on this variety: "Eine durchaus selbst~indige Form mit den Bl~ittern des P. nudicaule ssp. rubroauriantiacum var. corydalifolium, aber in den sonstigen Merkmalen zur subsp, radicatum gehSrig." Above I have suggested that P. rubroaurantiacum should belong to the alpinum complex, which seems to be supported by Fedde's note.

The other two of Fedde's Rocky Mountain taxa, var. columbianum and var. coloradense, are certainly allied to the radicatum complex. Fedde (1909a,b) states that var. columbianum, which he knew from a single specimen only (Macoun in 1890 on the Mountains of the Kicking Horse, 8000 ft, US), seems to form a link between the arctic var. radicatum and his other two taxa, var. pseudocorydali]olium and vat. coloradense, 375 and 1500 km further south, respectively, but he expects that in time more collections will fill these gaps. As var. pseudocorydali]olium now is known to be a synonym of P. pygmaeum of the alpinum group, the gap between the arctic and alpine representatives of radicatum is even larger, though more collections have been added later (see map, Fig. 1).

Fedde's var. columbianum and var. coloradense are very similar and both definitely belong to the radicatum complex, characterized by low growth, hairy leaves and scapes, ovate to conic capsules covered with dense brownish to black stiff hairs, and with a rather flat or only vaulted but not peaked stigma. Actually, the two varieties differ only slightly in characters of minor importance in this group so that it can be assumed that they are only color variants of the same genetical taxon.

Further studies in herbaria and in nature have led me to the conclusion that Fedde's two varieties are identical with P. kluanensis D. LSve which I described from Quill Creek Camp, 5577 ft altitude, near Kluane Lake in SW Yukon (LSve & Freedman, 1956). I t is apparently a fairly widespread taxon, even if its area is disjunct with very wide gaps (Fig. 1).

P. kluanensis belongs to, the radicatum complex, but, in addition to its morphological distinction, it is cytologically clearly separated from P. radicatum Rottb. s.str. (cf. ,~. LSve, 1962a) by having only 2n = 42 chromosomes instead of 2n = 56 as is typical of the latter species. The hexaploid number 2n = 42 + 0 to 2B chromosomes has been determined on plants from Colorado, Boulder County, Arapahoe Glacier, in screes, July, 1967, L6ve 11150 (COLO.).

The hexaploid P. kluanensis differs from the Alaska-Yukon populations of the octoploid P. radicatum ssp. occidentale Lundstr. by its overall smaller size, the scapes rarely exceeding 12-15 cm in height. The flowers and capsules are also smaller. The capsules are dark brown and covered by coarse brown hairs. The stigma is vaulted but not peaked, and its rays are connected by a yellowish membrane. P. radicatum ssp. occidentale is a much stouter and coarser plant in all respects and has taller and thicker scapes which often remain over winter and give the plants a ragged and untidy aspect. I ts capsules are about twice the size of those of P. kluanensis and have a broad, flat stigma with a yellow membrane between the rays.

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The stigma rays often appear to be more numerous than the four to six of P. kluanensis. In the herbarium of the University of Colorado Museum there are some specimens

of typical P. kluanensis which have previously been placed either under P. radicatum, P. nudicaule, or P. alpinum. In May, 1967, they were studied by G. Knaben, who made the remark on the labels: "Undescribed species, or if 2n = 42, P. hultenii Knaben." This must be based on some misunderstanding, since Knaben (1959a,b) referred to the paper by L6ve & Freedman (1956) which includes both descriptions and pictures which clearly show that this is a species of the radicatum group, whereas her P. hultenii obviously belongs to the nudicaule complex (cf. Knaben, 1959b, Figs. 6, 8). I t does not matter that both these taxa have the same chromosome number, since having the same number of chromosomes does not necessarily mean taxonomic identity. A morphological comparison of P. hultenii and other American species shows its clear relationship in all characters with P. macounii Greene and P. keelei Porsild, which obviously belong to the nudicaule complex (cf. L Sve & Freedman, 1956). P. macounii is a tetraploid with 2n = 28 chromosomes (Knaben, 1959a) and clearly distinct from P. hultenii the capsules of which are sturdier. P. keelei is rather similar to P. macounii but has a more easterly distribution, and Gj~erevoll (1963) has suggested that they may be conspecific subspecies. These three Alaskan taxa are characterized by tall growth, sometimes up to 50 cm in height, finely divided leaf blades on long petioles, relatively large flowers with numerous stamens, and very narrow, slender and long capsules with a peaked stigma. All close relationship between them and P. kluanensis can safely be ruled out.

THE UTAH POPULATIONS

P. kluanensis is in its southern range almost entirely situated on the high peaks along, or not too far from, the continental divide of the Rocky Mountains (Fig. 1). However, there are some localities in the Uintah Mountains of Utah (Fig. 1), in which Papaver has been found, and these poppies do not seem to belong near P. kluanensis. In the Summit County of Utah they have been collected on Gilbert and King's Peaks, and in the Duchesne County on Mt. Agassiz, everywhere at high altitude and in alpine ecological conditions and usually on west-facing talus slopes.

The most striking difference is the size and shape of the capsules (Fig. 3), which in the Utah specimens are larger, more obconical and with' coarser and somewhat darker hairs than those of P. kluanensis. Their leaves are longer, lax and weak, and have finely divided sections which end in an elongated point. The hairiness of the leaves varies from sparse to rather dense, and the hairs are mo~re or less similar to those of P. kluanensis. On the other hand, the sheaths are quite different from those of that species and resemble more the shiny, straw-colored and distinctive sheaths of P. alaskanum Hult6n. The manner of growth and the long cuffs, of sheaths also remind one of that species. In P. kluanensis the sheaths are dull and short, and the growth habit is more compact caespitose than in P. alaskanum, which has a loose, spreading rhizomatous root system. With the very few specimens at hand, five different collections only, and no information about their cytology, I feel it would be unwise to give the Utah specimens any taxonomic rank or name, though it may be suggested that they are related to P. alaskanum Hult6n and the Siberian P. pulvinatum Tolm. (Tolmachev, 1932), which are hard to fit into the radicatum or nudicaule complexes and are distinct from the alpinum group.

CONCLUSIONS

In the Rocky Mountains south of the Yukon Territory there are two, possibly three, distinct species that belong to the Papaver sect. Scapiflora. They are found exclusively at high altitudes above timberline and usually in screes or talus slopes.

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8 BRITTONIA [VOL. 21

The areas of these taxa do not overlap, and only one, P. kluanensis, has an extended but disjunct range.

P. pygmaeum and P. kluanensis belong to different groups within sect. Scapi/lora, the alpinum and the radicatum complexes respectively. The affiliation of the third taxon, here called the Utah population, is less clear. I t may be related to P. kluansis although I am inclined to. believe that it may represent a distinct complex to which P. ataskanum and P. pulvinatum may also belong. I t is not closely related to the alpinum complex.

P. pygmaeum is clearly related to the P. alpinum aggregate of Europe and appar- ently connected to it by a chain of related, though disjunct, taxa across the high mountains of inner Asia. The complex is characterized by bulbous-based, ivory- colored bristles on the small, roundish' capsules. The chromosome number is 2n = 14, except in P. albo-roseum which possibly also belongs to this complex as a distinct species.

P. kluanensis is a 42-chromosome mainly alpine taxon that belongs to the high polyploid and mainly arctic radicatum complex. I t is characterized by compact, caespitose growth, low scapes with small flowers, coarse capsules with broad stigmas and few stamens, normally shorter than the mature capsules.

In the Rocky Mountains there is no native species, that belongs to the nudicaule complex, but garden varieties of P. nudicaule are commonly grown both' in Canada and the United States. I t frequently escapes into wild habitats, but is never found above timberline. The native species of the nudicaute complex, which are met with in Alaska and the Yukon Territory, are characterized by tall growth, narrow and tall capsules with peaked stigmas and rather large flowers with a very high number of stamens. Their chromosome numbers range from the diploid 2n = 14 to the hexaploid 2n = 42.

P. pygmaeum is geographically confined to a very limited area around the borders of British Columbia, Alberta, and Montana in and around the Waterton-Glacier International Park. P. kluanensis ranges disjunct from Alaska-Yukon, generally along the spine of the Rocky Mountains as far as the Taos Mountains in New Mexico. The so-called Utah population is confined to a few high peaks of the Uintah Mountains in Utah. I t is in need of closer studies before its rank and relationship can be determined with certainty.

The following key is intended as an aid to distinguish the taxa of native Papaver found at high altitudes in the Rocky Mountains south' of the Yukon Territory:

1. High-alpine, caespitose, scapose plants with small flowers (11~-3 cm diam), and yellow or salmon-pink petals.

2. Capsules with few bristles, these ivory-colored, rigid, sharp-pointed, gnd with a bulbous swelling at the base; low (5-10 cm tall) caespitose plants with very small (11/~-2 cm diam) flowers; petals salmon-pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P . p y g m a e u m

2. Capsules with numerous coarse hairs varying in color from light brown to black; low (7-15 cm tall) caespitose plants with small (2-3 cm diam) flowers; petals yellow.

3. Densely caespitose plants with light brown short hairs on leaves and peduncles, particularly dense at the top of the latter; peduncles firm and erect at anthesis; leaves densely hairy, with short petioles and a short blade divided into broad segments; capsules roundish, about as broad as long, covered with brown hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P . k l u a n e n s i s

3. Loosely caespitose plants with lax peduncles and scant brown woolly hairiness; leaves sparsely hairy with long woolly hairs; petioles long and lax, blade with oblong-lanceolate remote segments; capsule about twice as long as broad, obconic, covered with dark brown hairs ............................... P a p a v e r from Uintah Mts., Utah

1. Low-altitude, scapose, large-flowered (3-6 cm diam) plants with white, yellow, orange, or red petals. Cultivated or escaped from cultivation . . . . . . . . . . . . . . . . . . . . . P . n u d i c a u l e sens. Iat.

Page 9: Papaver at High Altitudes in the Rocky Mountains

1969] LOVE: PAPAVER 9

ACKNOWLEDGMENTS

I am indebted to the curators and staff of the various herbaria which I have visited or from which loans for this work have been received: the Smithsonian Institution, Washington, D. C.; New York Botanical Garden; National Museum, and Depart- ment of Agriculture, Ottawa; University of Alberta, Edmonton; University of Wyoming, Laramie; University of California, Berkeley; California Academy of Sciences, San Francisco; and University of Colorado Museum, Boulder. Valuable help and information has also been received from Professors Askell LSve and Brij M. Kapoor of the University of Colorado, Professor M. Bowers, University of North- ern Michigan, Professor J. G. Packer, University of Alberta, and Dr. O. R6nning, Museum of Natural History, Trondheim, the last-mentioned two at present visiting professors at the University of Colorado. Financial support has been received through NSF grants GB-3371 and GB-6299.

LITERATURE CITED

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Faberg6, A. C. 1942. Genetics of the Scapiflora section of Papaver. I. The garden ]cela.nd poppy. Jour. Genetics 44: 169-193.

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Hedberg, O. 1967. Chromosome numbers of vascular plants from arctic and sub-arctic North America. Arkiv fSr Bot. (Stockholm) II. 6: 309-326.

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Hult6n, E. 1928. Flora of Kamtcbatka and the Adjacent Islands. II. Kungl. Svenska Vet. Handl. III. 5(2) : 1-218. (Papaveraceae, pp. 138-142.)

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Kilpfer , P. & C. Favarger . 1967. Premieres prospections caryologiques dans la flore orophile des Pyr6n6es et la Sierra Nevada. Compt. Rend. Acad. Sci. Paris 264(Ser. D) : 2463-2465.

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