pavlick1983fescue by R Wheeler

The taxonomy and distribution of Festuca idahoensis in British Columbia and northwestern Washington LEONE. PAVLICK Botany Division, British Columbia Provincial Museum, Victoria, B.C., Canada V8V 1x4 Received August 10, 1981

PAVLICK, L. E. 1983. The taxonomy and distribution of Festuca idahoensis in British Columbia and northwestern Washington. Can. J. Bot. 61: 345-353. Plants of Festuca idahoensis from west of the Cascade Mountains in British Columbia and northwestern Washington differ in their leaf morphology and panicle size from those east of the Cascade Mountains. The western plants are recognized as F. idahoensis var. roemeri. The geographic and ecological distribution of the species and a key for distinguishing the two varieties are given.

PAVLICK, L. E. 1983. The taxonomy and distribution of Festuca idahoensis in British Columbia and northwestern Washington. Can. J. Bot. 61: 345-353. Des plants de Festuca idahoensis provenant de l'ouest des Cascade Mountains en Colombie-Britannique et du nord-ouest de 1'Ctat de Washington diffkrent de ceux i I'est des Cascade Mountains par la morphologie de leurs feuilles et la grandeur de leurs panicules. On appkle les plants de la rigion ouest les Festuca idahoensis var. roemeri. La distribution gCographique et Ccologique de l'espkce est donnCe ainsi qu'une clef pour en distinguer les deux variCtCs. [Traduit par le journal]

Introduction Festuca idahoensis has long been known to occur east of the cascades in ~ ~ icolumbia, ~ i ~ but h its presence in the province west of the Cascades is not well docuknown from the Olympic PeninIt is' "la (Jones 1936; KuramotO and 1970), the Puget 1961) and the Sari Juan Islands (R. W. Fonda and J. A. Bernardi, unpublished data). It has been reported as occumng on Vancouver Island by St.-Yves (1925) and on the Gulf Islands by Carter and Newcombe (1921) and Eastham (1947), but none of these authors cited specimens to document the reports. ~~~l~~and M ~ ( 1 9 ~ and ~ ) scoggan B (1978) ~ also ~ state that F . idahoensis CIccurs on Vancouver Idand and the nearby Islands, but they9 to cite any specimens. Some time ago I noticed that a grass similar to F. idahoensis grew on summer-dry grass balds and other open sites at low elevations in western British Columbia. Although the plants were similar to F . idahoensis in many respects, they differed in some of their floral and vegetative characteristics, as well as ecological preferences, from specimens of that species growing east of the Cascades. A survey of various herbaria (V, UBC, DAO, CAN) showed that the western taxon was poorly understood, specimens of it sometimes being identified as F. idahoensis but some often as F. rubra. I decided, therefore, to examine the western taxon and compare it with the eastern taxon to clarify its taxonomy and distribution. 7

Materials and methods During 1977- 1979 I observed specimens of F. idahoensis in the field both along the Pacific Coast and east of the Cascade Mountains in British Columbia. In this period I collected F. idahoensis from five sites on southeastern Vancouver Island and the Gulf Islands and from four widely separated areas east the Cascade Mountains: South Thompson River Valley; Okanagan River drainage; Similkameen River drainage; east Kootenay River Valley. Voucher specimens of these are in V. These and additional specimens from the following herbaria were examined: ACK, CAN, DAO, LEA, UAC, V, WTU, and WWB. In all, about 150 specimens of F. idahoensis,

TABLE ~ 1. Geographical areas and numbers of F. idahoensis specimens studied. ( a ) Specimens identified and label data for 21 characters; ( c ) specirecorded; ( b ) specimens mens scored for leaf sheath diameter; ( d )specimens scored for leaf cross-section characters Geographic area British Columbia Vancouver Island, Gulf Islands East of Cascade Mts. Washington P"getLOwlandsy SanJuanlslands Olympic Mts. East of Cascade Mts. Alberta Saskatchewan Oregon california

(a)

(b)

(c)

27 62

10 10

18 19

20 14

11 2 42 1 9 2

6 1 2 1 5 2

4 10 2 2 1 4 1

0008-4026/83/010345-09$01 .OO/O 01983 National Research Council of Canada /Conseil national de recherches du Canada

CAN. 1. BOT. VOL. 61, 1983

TABLE2. Comparison of F . idahoensis specimens from seven areas in Canada and West of Cascade Mts. Vancouver Island, Gulf Islands Culm height, cm Basal tuft length, cm Basal leaf sheath diameter, mm Basal leaf sheath vestiture

36-85 14-33 (0.6)l.O-1.4(2.0) Glabrous, scabrous or pubescent

Basal leaf vestiture, abaxial

Puget Lowlands, San Juan Islands

Olympic Mts.

Scabrous or pubescent Scabrous

Basal leaf shape

Glabrous to scabrous (infrequently pubescent) Short to long scabrous (infrequently short pubescent) Obovate-pyriform

Glabrous to scabrous Scabrous or pubescent Scabrous to short pubescent Obovate-pyriform

Basal leaf diameter, mm Basal leaf nerve number Basal leaf margin disposition Basal leaf sclerenchyma patternt Panicle length, cm Spikelet length, mm Lower glume length, mm Upper glume length, mm Lowermost lemma length, mm Lemma awn maximum~length,mm Anther length

(0.5)0.6-O.g(l.2) 5*-7 Tending to open a, b, c, d 9.5-16.0 9.0-13.5 2.5-5.0 4.0-6.2 6.2-8.0 3.2-5.0 2.9-3.6

0.6-0.8 5-7 Tending to open a, b, d, e 12.0-15.0 9.5-10.5 3.2-3.5 4.4-4.8 6.2-6.8 3.5-5.0 (2.6)2.8-3.5

(0.4)0.6-0.8(1.2) 5-7 Tending to open a, b, d 8.0-14.0 10.0-12.0 2.5-3.9 4.2-5.0(5.5) 5.5-6.5(7.0) (2.5)3.0-4.5 3.0-3.6

Basal leaf vestiture, adaxial

Scabrous to short pubescent Obovate-pyriform

*Five nerves rather than seven were recorded in about 10% of Vancouver Island specimens. +Letters refer to leaf cross sections in Fig. I , which illustrates approximating sclerenchyma patterns

mainly from Canada but with lesser numbers from throughout the range of the species (Table l), were examined using a maximum 8 0 x dissecting microscope. All specimens used in the study were identified and annotated and their distribution (Fig. 2) and habitat, as reported on the labels, recorded. Forty F. idahoensis plants from widely separated areas of its range were scored for 21 characters; 58 plants were scored for 6 characters as seen in leaf cross section; 37 plants were scored for leaf sheath diameter (Table 1). A synthesis of characters which varied, expressed for each geographic area as a range of variation between individual specimens, is presented in Table 2. Within specimens, three-five basal leaves and (or) threefive spikelets, which I judged to express the range of variation for at least 80% of the leaves and (or) spikelets, were measured and the median value was recorded. For some, extreme values were also recorded. Within one of the median-value spikelets, glume lengths, lowermost lemma length, and maximum awn length were scored. Anther length was obtained from any available anther in the panicle. Within one of the median-value leaves, cross sections were cut from between one-quarter and one-half way up the length of the leaf blades and were examined at 80x with a dissecting microscope; shape, diameter, nerve number, vestiture, margin disposition, and sclerenchyma pattern were recorded. For basal leaf sheath diameter five leaf sheaths per specimen were measured about 7mm below sheath summits and the range of variation was recorded. The mean range of variation and extreme values for each geographic area are given in Table 2.

The following type specimens were examined. Festuca idahoensis Elmer, Abrams 688, Smith's Meadow, Shoshone Co., Idaho (DS); F. ovina var. ingrata Hack. ex Beal, Howell 23, Oregon (US); F . ovina var. oregona Hack. ex Beal, Cusick 753, Oregon (US); F . ovina var. colutnbiana Beal, Lake 2184, Blue Mts. E. Washington (WS). The latter three varieties are usually included in F . idahoensis. A type to represent the western F . idahoensis material was selected and examined.

Results and conclusions As viewed in the field the western F. idahoensis taxon from Vancouver Island typically has larger tussocks and coarser leaves than F. idahoensis specimens from east of the Cascade Mountains. A comparison between the two is given in Table 2 and Fig. 1. The western taxon typically has wider diameter leaf sheaths, wider diameter basal leaves, different leaf shape and disposition of leaf margins, seven rather than five nerves, different vestiture on adaxial leaf ribs, and different basal leaf sclerenchyma patterns (size, shape, and distribution of sclerenchyma strands in the leaf). The specimens from the Puget Lowlands and Olympic Mountains of western Washington tend to be shorter of culm, panicle, and lemma lengths than specimens from Vancouver Island, but they are similar in basal leaf

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Washington. Values in parentheses are the extremes recorded for individual specimens

East of Cascade Mts . British Columbia

Washington

38-75 16-32 (0.6)O.g-1.2(1.6)

42 20

47-57 12-15

62 16

Scabrous

Glabrous to puberulent Scabrous

Glabrous to scabrous Scabrous

Scabrous or pubescent Scabrous

Scabrous

Alberta

Saskatchewan

Densely long pubescent Plicate, hexagonal

(0.3)0.5-0.7 5

0.5-0.6 5

(0.3)0.4-0.5 5

0.7 5

Involute

10.0 11.8 4.3 5.7 8.5 4.0 2.4

7.0 8.0-8.8(9.3) 3.0(3.3) 4.1-4.3(4.4) (5.4)5.5-5.7(6.0) (3.2)3.8-6.0(7.0) (2.5)2.8(3.1)

diameter, shape, nerve number, margin disposition, adaxial vestiture, and sclerenchyma pattern (Table 2, Fig. 1). The western F. idahoensis taxon might be the F. ovina L. referred to by Macoun (1888) as occurring in the vicinity of Victoria (Macoun's voucher was not seen). It is not F. ovina L.; it has a very different leaf sclerenchyma pattern (cf. Fig. 1 with that of the European F. ovina L. sensu stricto in Holmen (1964) and Frederiksen (1981)) and distinctly longer panicles, spikelets, lemmas, awns, and anthers. The western F. idahoensis taxon is the plant occurring in Quercus garryana communities near Victoria which Roemer (1972) found to be similar to the European F. amethystina L. (H. L. Roemer, personal communication). It is likely the plant which St.-Yves (1925) reported (without specimen citation) from Vancouver Island as F. amethystina L. var. asperrima Hack. in Beal subv. idahoensis (Elm.) St.-Yves. It is not F. amethystina L.; F . amethystina L. var. asperrima, recognized by Hackel in Beal(1896) and St.-Yves (1925) for western North America, was treated as a synonym of F. idahoensis by Hitchcock (1951) and Hitchcock et al. (1969); it is different from the description of F. amethystina sensu stricto in Markgraf-

7.5 10.2 3.4 4.8 6.0 3.5 3.1

Dannenberg (1980) in that its lemmas range longer, its awns are much longer (usually 3-6 mm (cf. 1-2 mm for F. amethystina)),and its rachillas are visible at anthesis. It is not F. rubra; its leaf sclerenchyma pattern is very different (cf. Fig. 1 with that of F. rubra in St.-Yves (1925), Hubbard (1968), and Auquier (197 1a , 197 16)). It is a densely tufted bunchgrass with intravaginal innovations, whereas F. rubra is typically loosely or not at all tufted, with long rhizomes and extravaginal innovations; the scarious basal parts of its leaf sheaths do tend to split but do not shred into fibres in the manner of F. rubra. Specimens from Vancouver Island and western Washington are similar to those from east of the Cascade Mountains in inflorescence characters of spikelet, lower glume, upper glume, lemma, awn, and anther lengths (Table 2). In some of these characters the eastern specimens range to smaller lengths than the western specimens; but overall the spikelets of the two are very similar in appearance and overlap in size considerably so that they cannot be separated on this basis. On the basis of their similarity in inflorescence characters and overlap in morphometric range of vegetative characters, e.g., culm height, basal tuft height, leaf diameter (Table 2), 1 take them both to belong to F. idahoensis.

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FIG. 1. Leaf cross sections of F . idahoensis. (a-e) Variety roemeri. (a) Grand Mound Prairie (Puget Lowland). (b) Mount Angeles, Olympic Mountains. (c) Mount Finlayson, Vancouver Island. (d) Bear Hill, Vancouver Island. (e) Shelton (Puget Lowland). (f-h) Variety idahoensis. lllustrations show that little variation was found in var. idahoensis. (f) Paul Creek (Similkameen Drainage). (g) St. Mary's Prairie (east Kootenay Valley). (h) Chase (South Thompson Valley).

FIG. 2. Distribution of F . idahoensis in Canada and Olympic Peninsula area of Washington. Distribution in Prairie Provinces is consistent with Moss (1959) and Looman and Best (1979). Plotted from herbarium specimen data. A , var. roemeri, a, var. idahoensis.

The range of F. idahoensis in Canada and western Washington, as plotted from herbarium specimens as well as the collections made in 1978-1979, is given in Fig. 2. Both varieties occur in open situations unshaded by forest trees. Variety idahoensis occurs in grasslands and in open areas in the montane and subalpine zones. In the

grasslands it occupies relatively mesic situations; in the dry zone of southern British Columbia it occupies the more mesic midvalley slopes. In the montane and subalpine zones it occurs on the open, south-facing slopes, in open forests, and in grass balds. It has previously been recorded as an important component of the grasslands and dry forest vegetation of southern British Columbia, east of the Cascade Mountains, by Spilsbury and Tisdale (1944), Tisdale (1947), Eastham (1947,1948), Tisdale and McLean (1957), McLean and Holland (1958), Brayshaw (1965, 1970), and McLean ( 1970). The habitat of var. roemeri on Vancouver Island is influenced by being in the rain shadow of the Olympic and Vancouver Island mountains and in having mild winters in which most precipitation is rain. Leaf growth occurs in var. roemeri during the rainy winter months as well as the spring. On Vancouver Island var. roemeri occurs on south-facing, grass balds up to about 500 m elevation and at lower elevations in open, often rock, outcrop areas in the Pseudotsuga menziesii or Quercus garryana forests. It is recorded from grasslands on south-facing slopes in the San Juan Islands (R. W. Fonda and J. A. Bernardi, unpublished data) and

FIGS.3-8. Comparisons of form, leaf sheaths, leaves, panicle, etc. of F. idahoensis type specimens, as discussed in text. Specimens are mounted on standard herbarium sheets. In Figs. 3 , 5 , 7 , and 8 the tussocks have been more split up than in Figs. 4 and 6. Fig. 3. Festuca idahoensis Elmer. Typus, Abrams 688. Fig. 4. Festuca idahoensis var. roemeri Pavlick. Typus, Pavlick 78-233.Fig. 5. Festuca ovina var. oregona Hack. ex Beal. Typus, Cusick 753. Fig. 6. Festuca idahoensis var. roemeri Pavlick. Isotype, Pavlick 78-221. Fig. 7. Festuca ovina var. ingrata Hack. ex Beal. Typus, Howell 23. Fig. 8. Festuca ovina var. columbiana Beal. Typus, Lake 2184.

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prairies of the Puget Lowland (Lang 1961). In the subalpine zone of the Olympic Mountains it occurs on south- to west-facing slopes on the dry, northeastern rain shadow side of the mountains (Kuramoto and Bliss 1970; Buckingham and Tisch 1979). The F. idahoensis s~ecimens from west of the Cascade Mountains have a distinct range; they differ from the eastern specimens as noted above and in the key; and their ecological situations are different as noted above. These western specimens are proposed to be a new variety, var. roemeri as diagnosed below. Plantae dense caespitosae caespitibus plemmque illis varietatis idahoensis amplioribus. Vaginae exteriores basales (0.6)l .O- 1.4(2.0) mm latae, diametro intra 7 mm ab apice (vaginarum) metientes, habitum grossum caespitum formantes. Panicula 9.5- 16 cm longa, illa varietatis idahoensis (7-1 1 cm) longior. Laminae foliomm conduplicatae, sectione transversali obovatopyriformes, diametro (longae axi pertinenti) (0.5)0.60.9(1.2) mm, plemmque 7 nerves, costis sparsim breviter vel longiuscule scabrosis (rarius breviter pubescentibus). TYPUS:Pavlick 78-233. Mt. Finlayson, Vancouver Island. 28 June 1978 (V) (Fig. 4). C. L. Hitchcock in Hitchcock et al. (1969) applied the name var. oregona to alpine and subalpine plants in western Washington, which presumably covered the Olympic Mountain specimens examined in this study. However, for this taxon his key suggests a plant with short blades mostly less than 7(10) cm long and panicle usually spreading. In the present study the Olympic Mountain specimens had a basal tuft length of 10-21 cm and panicle branches which varied from spreading, ascending to erect-appressed. The type specimen of var. oregona has lower panicle branches ascending and leaf blades 10 cm long thus differing from Hitchcock's key for var. oregona. Buckingham and Tisch (1979) recognized both var. oregona and var. idahoensis for the subalpine of the northern Olympic Mountains, presumably based on the key provided in Hitchcock et al. (1969). The specimens from this area examined by the author show a large range in morphological variation but are similar in most leaf and spikelet characters so as to be considered part of the same variety (Table 2). Both varieties of F . idahoensis vary in degree of anthocyanic coloring, in vestiture of vegetative parts, and in the angle of their panicle branches. High-altitude plants, e.g. those from the Olympic Mountains, have intensely anthocyanic spikelets, rather than green, and the tips of the lemmas have a gold border, which is characteristic of the species. Anthocyanic culms are often present in subalpine and some montane plants, although green culms are more usual for the species.

Festuca idahoensis usually has glabrous or scabrous leaf sheaths and culms and scabrous leaves (abaxial surface). However, this is variable and a few specimens examined were either pubescent, pilose, or villose on one or more of these organs, e.g. Harrison 61 . I , Mt. Douglas, Vancouver Isl. (V); Pavlick 79-370, Bear Hill, Vancouver Isl.; Pavlick 79-43, 79-41, Buck Hill, Vancouver Isl.; Pavlick & Sax 78-150,Tobacco Plains (Kootenay R.) (V); Pavlick & Sax 78-5,Surveyors Lake (Kootenay R.) (V). St.-Yves (1925) described F. amethystina L. var. asperrima Hack. ex Beal subv. idahoensis (Elmer) St.-Yves f. dasyphylla from Oregon having pilose leaves and villose sheaths. The type specimen of var. ingrata, although distinct in leaf morphology from the Vancouver Island specimens, has pilose sheaths, leaves, and culms. The panicle in F. idahoensis varies from the usual condition of having its branches ascending to erect-appressed to one of having branches spreading and open. This latter condition was notably present in the east Kootenay Valley and in the Olympic Mountains. The material on the herbarium sheet of the type specimen of F. idahoensis (Abrams 688) might be heterogeneous. A culm with a panicle, a second panicle in a paper packet, and a separate basal shoot is present (Fig. 3). The culm with the panicle is here selected as the type specimen. It is similar to and well represents the Canadian F. idahoensis specimens I examined. It is also similar to, and I regard as conspecific with, the type specimens of F . ovina varieties ingrata, oregona, and columbiana. However, the leaves of the basal shoot on the Abrams 688 sheet are different from those of any of the other F . idahoensis specimens I have examined. The leaves are uniformly minutely scabrid, the second nerve from the leaf margin is unusually large, and the leaf sclerenchyma is continuous around the entire leaf. More studies on material from the western United States is needed to determine whether this basal shoot belongs to F. idahoensis. The type specimens of F . ovina L. varieties ingrata, oregona, and columbiana are all very similar in morphology (Table 3, Figs. 3, 5, 7 , 8). Variety ingrata is notably different from the others in having pilose leaf sheaths, leaves, and culm. The type of variety oregona differs from the other types in having a shorter culm, some spikelets with fewer florets, and a panicle with only one branch per node. In view of the shared similarities with the other types and with the specimens from east of the Cascade Mountains which I examined these differences are not considered significant. A nomenclatural summary for F . idahoensis can be found in Hitchcock et al. (1969). Recently Boivin (1967, 1979) included F. idahoensis in F. occidentalis; Pavlick (1982) has presented an argument for recognizing each of these as distinct species.

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VOL. 61, 1983

KEYTO Festuca idahoensis IN CANADA Basal sheaths 0.8-1.2 mm diameter about 7 mm below summit; panicle 7-1 1 cm long; basal leaf diameter (long axis) (0.3-)0.5-0.7 mm diameter, plant thus fine in aspect; leaves plicate, hexagonal, involute (near midleaf), usually with 5 nerves, the ribs (adaxial side) densely long pubescent; range in Canada, east of the Cascade Mountains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . var. idahoensis Basal sheaths 1 .O- 1.4 mm diameter about 7 mm below summit; panicle 9.5- 16.0 cm long; basal leaf diameter (long axis) (0.5-)0.6-0.9(-1.2) mm, plant thus coarse in aspect; leaves obovate-pyriform in cross section (near midleaf) with margins tending to open, usually with 7 nerves, the ribs (adaxial side) lightly scabrous to sparsely short pubescent; range in Canada, west of the Cascade Mountains (southeastern Vancouver Island and Gulf Islands). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . var. roemeri

Acknowledgements

I wish to acknowledge the assistance of Dr. H. L. Roemer, British Columbia Ecological Reserves, Victoria, B.C., for his interest in and his views o n coastal F. idahoensis. Dr. Roemer discovered this variety o n several new sites o n southeastern Vancouver Island. I acknowledge and thank Dr. V. J. Krajina, University of British Columbia, and Dr. R. T. Ogilvie and Dr. T. C. Brayshaw, both at British Columbia Provincial Museum, for reviewing and commenting on the manuscript, and Dr. V. J. Krajina for translating into Latin the diagnosis of var. roemeri. AUQUIER,P. 1971a. Festuca rubra L. subsp. pruinosa (Hack.) Piper: morphologie, ecologie, taxonomie. Lejeunia, 56: 1-16. 1971b. Le probleme de Festuca rubra L. subsp. arenaria (Osb.) Richt. et ses relations avec F . juncifolia St. Amans. Lejeunia, 57: 1-24. BEAL,W. J. 1896. Grasses of North America for farmers and students. Vol. 2. Henry Holt & Co., New York. BOIVIN,B. 1967. Enumeration des plants du Canada. VI. Monopsides (2e partie). Nat. Can. (Que.), 94: 471-528. 1979. Flora of the prairie provinces. Part 4. Cyperaceae. Provancheria 5. Phytologia, 43(1): 1-106. BRAYSHAW, T. C. 1965. The dry forest of southern British Columbia. Ecol. W. N. Am. 1: 65-75. 1970. The dry forests of British Columbia. Syesis, 3: 17-43. BUCKINGHAM, N. M., and E. L. TISCH.1979. Vascular plants of the Olympic Peninsula, Washington. National Park Service and University of Washington. University of Washington/Cooperative Park Studies Unit Report B-79-2. CARTER,W. R., and C. F. NEWCOMBE. 1921. A preliminary catalogue of the flora of Vancouver and Queen Charlotte Islands. British Columbia Provincial Museum of Natural History, Victoria. EASTHAM,J. W. 1947. Supplement to "flora of British Columbia." (J. K. Henry). British Columbia Provincial Museum, Victoria. Spec. Publ. No. 1. 1948. Notes on plants collected in 1947 chiefly in the Rocky Mountain Trench, between the Rocky and Selkirk Mountains of British Columbia. Rep. Prov. Mus. Nat. Hist. Anthrop. 1947. FREDERIKSEN, S. 1981. Festuca vivipara (Poaceae) in the North Atlantic area. Nord. J. Bot. 1: 277-292.

HITCHCOCK, A. S. 1951. Manual of the grasses of the United States. U.S. Dep. Agric. Misc. Publ. 200: 1-1051. HITCHCOCK, C. L., A. CRONQUIST, and M. OWNBEY. 1969. Vascular plants of the Pacific northwest. Part I. Vascular cryptograms, gymnosperms and monocotyledons. University of Washington Press, Seattle. HOLMEN,K. 1964. Cytotaxonomic studies on the Arctic Alaskan flora. The genus Festuca. Bot. Not. 117: 109- 118. HUBBARD, C. E. 1968. Grasses. 2nd ed. Penguin Books Ltd., Harmondsworth, Middlesex, England. JONES,G. N. 1936. A botanical survey of the Olympic Peninsula, Washington. Univ. Wash. Publ. Biol. 5: 1-286. KURAMOTO, R. T., and L. C. BLISS. 1970. Ecology of subalpine meadows in the Olympic Mountains, Washington. Ecol. Monogr. 40: 317-347. LANG,F. A. 1961. A study of vegetation change in the gravelly prairies of Pierce and Thurston Counties, western Washington. M.Sc. thesis, University of Washington, Seattle. LOOMAN,J., and K. F. BEST. 1979. Budd's flora of the Canadian Prairie Provinces. Agriculture Canada, Ottawa. Publ. 1662. MACOUN,J. 1888. Catalogue of Canadian plants. Part 4. Endogens. Geological and Natural History Society of Canada, Ottawa. MARKGRAF-DANNENBERG, I. 1980. Festuca L. In Flora Europaea. Vol. 5. Edited by T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters, and D. A. Webb. Cambridge University Press, Cambridge. MCLEAN,A. 1970. Plant communities of Similkameen Valley, British Columbia and their relationships to soils. Ecol. Monogr. 40: 403-424. MCLEAN,A., and W. C. HOLLAND. 1958. Vegetation zones and their relationships to the soils and climate of the upper Columbia Valley. Can. J. Plant Sci. 38: 328-345. Moss, E. H. 959. Flora of Alberta. University of Toronto Press, Toronto. PAVLICK, L. E. 1982. Notes on the taxonomy and nomenclature of Festuca occidentalis and F . idahoensis. Can. J. Bot. 61. This issue. ROEMER, H. L. 1972. Forest vegetation and environments on the Saanich Peninsula, Vancouver Island. Ph.D. thesis, University of Victoria, Victoria. SCOGGAN, H. J. 1978. The flora of Canada. Part 2. National Museums of Canada, Otawa. pp. 93-545. SPILSBURY, R. H., and E. W. TISDALE.1944. Soil plant relationships and vertical zonation in the southern interior of British Columbia. Sci. Agric. 24: 395-436.

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ST.-YVES,A. 1925. Contribution a l'etude des Festuca (subgen. Eu-Festuca) de 1'Amerique du Nord et du Mexique. Candollea, 2: 229-316. TAYLOR, R. L., and B. MACBRYDE. 1977. Vascular plants of British Columbia. The Botanical Garden, The University of British Columbia. Tech. Bull. 4. The University of British Columbia Press, Vancouver.

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Appendix 1. Partial list of specimens examined East of Cascade Mountains in British Columbia and Alberta: Mile 14, Flathead Rd., Bell and Davidson 54 (V); Kikomun Creek Park, Pavlick & Sax 78-5 (V); Snehumption Creek, Brayshaw & Pavlick 77-348 (V); St. Mary's Band Reserve, Pavlick & Sax (V); Mt. Kobau, Brayshaw & Pavlick 77-484 (V); Pollock Mt., Flathead, Bell & Davidson 923 (V); Paul Creek Valley, Brayshaw & Pavlick 77-538 (V); S. ThompsonR. near Chase, Pavlick & Taylor 79-120 (V); Richter Pass, Tisdale, 1936 (ACK); Lytton, Malte, 1914 (CAN); Princeton, Krajina, 1965 (DAO); Invermere, McLean, 1954 (DAO); Nicola, Tisdale, 1940 (ACK); Cypress Hills Park, de Vries 2036 (UAC); Waterton Lakes Park, Malte & Watson 128 (UAC); Frank Slide, Sharp 15 (LEA). Vancouver Island and Gulf Islands: Newcastle Island, Pavlick 78-240 (V); Mt. Douglas, V.I., Harrison 61-1 (V); Mayne Island, Macoun, 1914 (V); Mt. Finlayson, Pavlick 78-233 & 79-18 (V); Buck Hill, V.I., Pavlick 79-41 & 79-43 (V); Mt. Bruce, Saltspring I., Shank 62 (V); Observatory Hill, V.I., Newcombe, 1921 (V); Bear Hill, V.I., Pavlick 79-371 (V). Puget Lowlands, Washington: Grand Mound Prairie, Thurston Co., Otis 1982 (WTU). Shelton, Mason Co., Jones 3821 (WTU). Olympic Mountains, Washington: Hurricane Ridge, Jones 3388 (WTU); Marmot Pass, Thompson 9957 (WTU); Mt. Angeles, Thompson 7530 (WTU); Deer Park, Denton 3355 (WTU); Deer Park, Belsky, 1978 (WTU). Oregon: Saddle Mt., Clatsop Co., Detling 6976 (WTU). Mt. Grayback, Josephine Co., Thompson 12998 (WTU); Umpqua Valley, Cusick 4056 (WTU). Siskiyou Mts., Thompson 12389 (WTU).