Historic, archived document
Do not assume content reflects current scientific knowledge, policies, or practices.
Bulletin No. 11. Agros
U. S. DEPARTMENT OF AGRICULTURE.
DIVISION OF AGROSTOLOGY. [Grata and Forage Plant Investigations.]
STUDIES
ON
AMERICAN CRASSE
I. A REVISION OF THE NORTH AMERICAN SPECIES OF CALAMAGROSTIS.
By Thomas H. Keakney, Jr.
II. DESCRIPTIONS OF NEW OR LITTLE-KNOWN GRASSES.
By F. Lajison-Sckibneb.
ISSTTED JTJLY SO, 1898.
WASHINGTON:
GOVERNMENT PRINTING OFFICE.
1898.
Bulletin No. 11. Agros.
U. S. DEPARTMENT OF AGRICULTURE.
DIVISION OF AGROSTOLOGY.
[CJrass antl Forage Plant Investigations.]
STUDIES
ON
AMERICAN GRASSES
I. A REVISION or THE NORTH AMERICAN SPECIES OF CALAMAGROSTIS.
By Thomas H. Keabxev, Jr.
II. DESCRIPTIONS OF NEW OR LITTLE-KNOWN GRASSES.
By F. Lamson-Scribnek.
ISSUED JULY SO, 1898.
WASHINGTON:
GOVERNMENT PRINTING OFFICE. 1898.
LETTER OF TRANSMITTAL
U. S. Department of Agriculture,
Division of Agrostology, Washington, D. C, May 19, 1898.
Sir: I have the honor to transmit for your approval the manuscript of a continuation of Studies on American Grasses, embracing (1) "A revision of the North American species of Calamagrostis,11 by Mr. Thomas H. Kearney, jr., assistant agrostologist, and (2) "Descriptions of new or little-known grasses," by the Agrostologist, and respectfully recommend its publication as Bulletin No. 11 of this division. In the paper on the revision of the genus Calamagrostis only those species are included which occur in North America north of Mexico. All enumerated are natives, and of the thirty-eigbt species described thirty-three are believed to be endemic. Twenty-three species and varieties are described as new.
In the Rocky Mountain region there are twelve species, and while none are found in the Gulf States excepting in northern Georgia, they extend southward through Mexico, and the species multiply along the Andes of South America. In his enumeration of the species of Calamagrostis of the higher Andes Mr. H. L. Weddell characterizes sixty species. In North America the species are most abundant on the Pacific slope, where twenty-five species are known to occur.
The new species are fully described, and the synonymy and distribu- tion of all the species included is fully presented. The keys of analysis have been made with much care, and can not fail to be found helpful in determining the species. Some of the species present so many forms that their limitation is difficult. In several cases it is impossible to draw sharp lines of separation, and this is especially true of Calama- grostis canadensis and Calamagrostis langsdorffii, also Calamagrostis hyperborea and Calamagrostis inexpansa. The descriptions and the data furnished relative to the distribution of the species are based upon specimens contained in the United States National Herbarium, and also those in the leading herbaria of the country.
That it has been possible to examine so large a series of specimens is due to the courtesy of those in charge of the several collections sub- mitted for the purpose. Expression of thanks is here tendered to Mr. James Macoun, of the Geological and Natural History Survey of
3
4
Canada; Dr. B. L. Eobinson, Gray Herbarium, Cambridge, Mass.; Dr. John K. Small, Columbia University; Mr. Stewardson Brown, of the Academy of Kataral Sciences of Philadelphia; Dr. E. L. Greene, Catholic University; Mr. Theo. Holm, Washington, D. C; Miss Alice Eastwood, California Academy of Sciences. Thanks are also due Dr. C. H. Merriam and Mr. Vernon Bailey for their kind assistance in determining the zonal limits of the species. Kespectfully,
F. LAMSON SCEIBNER,
Agrostologist.
Hon. James Wilson,
Secretary of Agriculture.
CONTENTS.
Page.
I. A revision of the North American species of Calamagrostis. By Thomas H.
Kearney, jr 7
Introduction 7
Classification 7
Geographical distribution 9
Ecology 10
Teratology 12
Hybridism 13
Species excluded 13
North American species of Calamagrostis 13
Analytical key to the species 13
II. Descriptions of new or little-known grasses. By F. Lamson-Scribner 42
5
ILLUSTRATIONS.
PLATES.
Page.
Plate I. Panicum linearifolium 60
II. Panicum equilaterale 60
III. Chfetochloa latifolia 60
IV. Stipa williainsii r 60
V. Sporobolus palnieri 60
VI. Zeugites pringlei 60
VII. Eragrostis viscosa 60
VIII. Poa juncifolia 60
IX. Poahanseni 60
X. Poa atropurpurea 60
XI. Poa longepedunculata 60
XII. Agropyron elmeri ,. 60
XIII. Agropyron brevifolium 60
XIV. Elymus capitatns 60
XV. Elynras saxioolus 60
XVI. Elymus ciliatus 60
XVII. Elymus simplex.. 60
FIGURES.
Fig. 1. Calamagrostis fasciculata 23
2. Panicum implicatum 43
3. Panicum baldwinii 44
4. Panicum wrightianum 45
5. Sporobolus thurberi 48
6. Sporobolus simplex 49
7. Agrostis paludosa 50
8. Trisetum argenteum 51
9. Trisetum ■wolfii 52
10. Trisetum muticum 53
11. Poa capillaris 54
12. Elymus hanseni 56
6
STUDIES ON AMERICAN GRASSES.
I. A REVISION OF THE NORTH AMERICAN SPECIES OF CALAMAGROSTIS.
By Thomas H. Kearney, Jr.
INTRODUCTION.
In North America, north of Mexico, thirty-eight species of Calama- grostis are known to occur, eleven of which are here published for the first time. All of these are native, no introduced species having as yet been reported. All belong to the section Deyeuxia Hack. (Clarion, as a genus), which is characterized by the usually hairy prolongation of the rachilla behind the palea. This prolongation is villous along its whole length, to just below the apex,1 except in 0. cinnoides, which has the extension of the rachilla naked to just below the- apex, where it bears a circle of long hairs, not unlike the pappus of some Cichoriacece. C. cinnoides is also unique among North American species in its pubes- cent caryopsis. All of our species, excepting C. breweri, haye well- developed creeping rootstocks and at least some of the innovations extravaginal. The culms, usually simple, are sometimes branched in G. langsdorffii, G. canadensis, and G. macouniana.
CLASSIFICATION.
A satisfactory segregation of the North American species of Calama- grostis is rendered difficult by their great variability and the existence of a series of intergrading forms between many that are none the less too distinct in the typical form to admit of their being united. Thus from 0. canadensis to G. langsdorffii a perfect gradation can be traced. From G. hyperborea to G. neglecta, on the one hand, and to C. inexpansa on the other, the transition is equally uninterrupted. The consequent impossibility of sharply defining some of the' species has made it expedient to regard several of these intermediate forms as varieties of one of the two connected species, although rather arbitrary characters must be used for distinguishing them. This frequency of intergrada- tion has made the construction of a serviceable key a task of more than ordinary difficulty.
1 Quite naked abnormally in one or two species.
7
8
As the sequence of species adopted iu this revision is somewhat artificial, a brief discussion of their apparent natural relationships is here inserted. Most of the North American species can be arranged into five rather well-defined groups, represented, respectively, by G. neglecta, G. canadensis, G. purpurascens, C. aleutica, and G. deschamp- sioides.
The first group is the most strongly European in its affinities. The following diagram indicates the relationships and possible genealogy of its members, the length of the connecting lines measuring to some extent the closeness of the relationship :
micrantha laxiflora \ I
alaskana neglecta
i
crassiglumis h yperborea
I I labradorica
breviseta
The second group is possibly derived from the neglecta type, although the probable line of descent is obscure. This group, like the first, is closely related to European and northern Asiatic types.
macouniana I
nemoralis — canadensis — langsdorffii
I I N
porteri blanda scribneri
The third group is perhaps derived from the neglecta type through ■G. hyperborea. It is significant that both C. purpurascens and G. hyper- borea are found in North America and, elsewhere, only iu Greenland, and are apparently the only species with such distribution. The pur- purascens allies are related to the European G. sylvatica, but are other- wise more nearly an endemic type than are the two preceding groups.
montatiensia I
tweedyi PURruRASCENS — vaseyi
I
howellii— foliosa
The fourth group contains two well-defined series of species, the first nearly related to G. aleutica, the second to G. suhsdorfii. Through the second series this group is connected with C. purmirascens and its allies. Both types are purely endemic.
scopn lorum I
cumckii anqusta koelerioides
I I I
suksdorfii aleutica — vilfa'formis
I I
ru bescen s 8« bflexuosa
fascieitlata
californica I
inexpansa
9
The fifth group is of entirely problematical relationship. With the exception of G. deschampsioides, which ranges from the Pribilof Islands to Kamchatka, and is therefore more Asiatic than American, the spe- cies are endemic and are found only in the mountains of California.
breweri — lemmoni deschampsioides lolanderi.
Finally, 0. cinnoides stands alone, an extremely isolated type without near relations.
GEOGRAPHICAL DISTRIBUTION.
The genus Calamagrostis contains approximately one hundred and fifty species, widely distributed over the globe, but in such a manner as to point strongly to a boreal origin, according to the generally accepted theory of migration during the glacial epoch, for outside of cold and cool temperate regions the species are found only on the higher mountains.
The development of numerous endemic forms in Australia and New Zealand is at first sight an obstacle to this theory of the origin of the genus, but the occurrence of scattered species on the mountains of Iudia and the Malayan region deprives that objection of its force. The great number of species along tlie South American Cordilleras is per- fectly in harmony with the known laws of plant distribution. The genus is at present most strongly developed in northern Europe and Siberia (twenty-five species), western Forth America (thirty-two spe- cies), the Andes region (seventy species), and Australia, including New Zealand and Tasmania (twenty species). There are eleven species in Atlantic North America, fifteen in Mexico and Central America, fifteen in the Himalayan-Malayan region, ten in eastern Asia, and fifteen in central and western Asia. With the exception of about ten species of the section Upigeos Koch, inhabiting the northern temperate regions of the Old World, all belong to the section Deyeuxia, which is often regarded as a separate genus, but is at best an artificial and unsatis- factory one.
Two of the thirty-eight North American species (G. langsdorffii and C. neglecta) occur also in northern Europe and Asia, two more (C. pur- purascens and C. hyperborea) are found outside this continent only in Greenland, while a fifth (C. deseliampsioides) is common to the Pribilof Islands and northeastern Asia. The remaining sj)ecies are believed to be endemic. Considering the distribution of the genus in North America from a purely geographical standpoint we find ten species in the Atlantic- Appalachian region. Of these, three extend into the Southern States, one occurring in the undulating "middle" country and in the foothills, the other two only upon the highest mountains (above 1,800 meters). The genus is not represented in Florida, Mississippi, Loui- siana, Arkansas, the Indian Territory, and Texas. Some half dozen species occur in the prairie districts and the region about the Great
10
Lakes. Four or live are found on the Great Plains east of the Eocky Mountains. In the Eocky Mountains proper there are twelve species, of which three extend as far south as New Mexico and Arizona. On the Pacific slope from southern Alaska to California there are twenty- five species in the mountains and along the coast. On the Alaskan peninsula and islands the genus is represented by seven species.
Turning to the biologically more important distribution by life zones, whose limits are fixed by sums total of effective temperature during a definite period, we find the species pretty equally divided between the Boreal and Transition zones — a distribution to be expected in a genus of northern origin. Only a single species (G. purpuraseens) passes timber line northward and on the higher Eockies, and becomes truly Arctic- Alpine. Several, notably G. breweri in the Sierra Nevada, and C. neglecta, G. langsdorffii, and G. hyperborea northward, extend into the upper, or Hudsonian, belt of the boreal zone, in some cases probably nearly reaching timber line. Other species, for example, C. cinnoides in the Atlantic States, and G. canadensis and C. hyperborea elongata in the central prairie region, occur in country which is, broadly speaking, upper Sonoran. It is probable, however, that the modifying influence of a moist habitat accounts for such extensions, and that they are to be regarded as outposts of the Transition. None of the species can be regarded as of definitely Sonoran distribution.
The following is an approximate classification of the species by zones. It must be noted, however, that the most careful and thorough field work is necessary to make possible a complete and accurate defini- tion of the zonal limits.
1. Arctic-Alpine and Boreal. — C. purpurasoens.
2. Boreal. — C. deschampsioides, lemmoni, breweri, foliosa (?), vaseyi, tweedyi, aleuUca,
riibe&cens, breviseta, langsdorffii, canadensis acuminata, scribneri, alaskana, laxiflora, neglecta, micrautha, labradorica, hyperborea, crassiglumis.
3. Boreal and Transition. — C. suksdorfii, canadensis, hyperborea elongata, hyper-
borea americana.
4. Transition. — C. bolanderi, howellii, inontanensis, vilfaeformis, koelerioides, angusta,
subflexuosa, fasciculata, suksdorfii luxuriates, porteri, nemoralis, blanda (?) macouni- ana, ensickii, scopulorum, cinnoides, inexpansa, Californica(f).
ECOLOGY.
Plants are classed in respect to habitat and the resulting adaptations as hydrophytes (water plants), xerophytes (dry-soil plants), mesophytes (intermediate as to moisture requirement), and halophytes (salt-loving plants), the last class, however, being of decidedly less importance than the others. Nearly all the North American species of Galamagrostis belong to the first two classes, and the majority are hydrophytes. It is noteworthy, however, that nearly all the species, even those growing in the wettest soil, have characters that are generally supposed to belong rather to xerophile plants, such as narrow, erect, strongly involute leaf blades, short hairs or papillae on the upper leaf surface, a thin coating
11
of wax giving the plant a glaucous appearance, etc. This apparent anomaly is sometimes ascribed to the coldness of a wet soil, which diminishes the water-absorbing capacity of the root system and thus renders necessary the same contrivances to prevent too rapid transpi- ration that we see in plants inhabiting very dry soil. The smallest amount of xerophile habit is exhibited by 0. canadensis and its neai^est allies, especially when growing in shaded ground. On the other hand, the purpurascens group, with the exception of C. tweedyi, are u bunch grasses" with marked xerophile adaptation, conspicuously glaucous, having low, strict culms with tunicated bases in strong clumps from short, strong rootstoeks, thickish, strongly involute, narrow leaf blades, etc.
An absolute ecological classification of the species is impracticable, for the same species may be under different conditions either hydro- phile, niesophile, or xerophile. As has already been noted, this is fre- quently the case with species which toward the north inhabit dry, sunny situations, but further south take refuge from the increasing heat in moist woods or cold swamps. The following classification is based partly upon that of Warming.1 A number of species are entirely omitted, no data as to their habit being available. Others are enu- merated under more than one association class.
A. Hydrophytes.
1. Open marshes and wet meadows. — C. canadensis, langsdorffii, scribneri,
macouniaua, neglecta, micrantha, laxiflora, hyperborea (especially var. elongata), inexpansa, crassiglnmis, cinnoides, breviseta, bolanderi, deschampsioides, aleut- ica (possibly halophile).
2. Sphagnum bogs. — C. cinnoides (sometimes), alasl'ana (?).
3. Low, moist woods and thickets (almost a mesophile association). —
C. canadensis (sometimes), langsdorffii (sometimes), blanda, neglecta borealis, suksdorfii (normally xerophile).
4. Wet cliffs. — C. hyperborea americana (sometimes), memoralis (sometimes).
B. Xerophytes.
1. On or among rocks. — C. purpurascens, vaseyi, howellii, scopulornm, labradorica,
hyperborea (sometimes).
2. Sandy or gravelly shores of lakes and rivers.— C. rubescens, hyperborea
(sometimes), hyperborea americana (sometimes), hyperborea elongata (some- times), crassiglnmis (sometimes), breriseta lacuslris (probably often hyclro- phile). It is doubtful whether this association is properly xerophile.
3. Sand hills. — C. hyperborea americana (sometimes).
4. Prairies and "bench lands."— C. montanensis.
5. Upland woods. — C. porieri, memoralis, suksdorfii (usually ), rubescens, purpu-
rascens (sometimes), vilfaeformis, koelerioides.
C. Halophytes.
Coast marshes. — C. aleutica (f).
In regard to abundance of individuals, the xerophile species are either copious or sparse, but usually scattered. (C. suksdorfii is an exception, for in the Cascade Mountains of Oregon and Washington it grows socially, sometimes covering the ground in the pine woods to
1 Lehrb. der oekolog. Pilanzengeogr. Deutsche Ausgabe, 1896.
12
the exclusion of other herbaceous vegetation.) Species of hydrophile habit, ou the other hand, are often gregarious or even social. G. cana- densis, for example, occasionally grows in nearly pure formations, cover- ing open swamps and meadows with an almost unmixed growth.
TERATOLOGY.
Abnormal variation of different organs is not infrequent in Calama- grostis, especially in specimens growing at high altitudes and latitudes. Proliferation of the spikelets, the presence of a second (staminate) flower or even a third in the spikelets, and increase in the number of nerves in the glumes, are the most common deviations from normal characters. Such deviations are sometimes, but by no means always, due to disease. The following abnormalities in various organs were noted in the material examined in the preparation of this paper:
1. Three well-developed leafy branches from one of the cnlm nodes (C. canadensis acuminata).
2. Rudimentary sheath (?) at first node of rachis of panicle extended into a several- nerved bract about 5 mm. long, having the appearance and texture of a ligule <G. langsdorffii): into a bract about 4 mm. long, flat, ovate, 3-nerved (C. canadensis): into a bract 3 mm. long, somewhat spreading, sulbulate, chartaceous, possibly repre- senting a blade (C. macouniana).
3. Panicle deeply lobed, with the appearance of a cluster of panicles (C.neglecta horealis).
4. Most of the spikelets 2-flowered, one 3-flowered. In the latter case the first flowering glume subtending an undeveloped caryopsis, the secoud a mature cary- opsis, and the third empty, the rachilla being villous between the glumes and extending beyond the uppermost one (0. aleuUea). Many of the spikelets in some panicles 2-flowered, the second flower staminate, with well-developed but some- what smaller anthers, the rachilla bearded between the two flowering glumes and with the usual bearded prolongation (C. subflexuosa). A number of spikelets in one panicle 2-flowered, the second flower staminate, with a smaller but well-developed flowering glume and paloa (C.breviseta). A second (staminate) flower with well- developed anthers, the rachilla villous between the flowering glumes and with the usual villous prolongation (C. purpurascens). In one (diseased) panicle each spikelet with a second (5-nerved) flowering glume slightly raised above the first (7-nerved), both glumes having dorsal awns of about equal length and neither having palese, the Tachilla not extended beyond the second (C. purpurascens). A second* (staminate) flower with well-developed anthers subtended by a narrow, short-awned glume (C.cusickii). A second flowering glume, bearing a long, nearly apical awn (C. howellii). A slender awn, equaling the flowering glume, borne at the summit of the prolongation of the rachilla (C. bolanderi).
5. First empty glume sometimes with a faint lateral nerve (C. purpurascens, etc.). In an abnormal, 2-flowored spikelet, the first empty glume cleft nearly to the middle, the lobe nearly as wide as the glume itself and with a second (lateral) nerve extend- ing from the base of the glume to the apex of the lobe ; the second glume cleft to the very base, the halves each 3-nerved and each nearly as large as the normal empty glume (C. subflexuosa). First empty glume frequently 3-nerved and the second 4-nerved, the fourth nerve faint and disappearing a short distance above the base (C. aleutica).
6. Flowering glume in one spikelet 6-nerved, the sixth nerve fainter than the rest and extending into a minute lateral tooth some distance below the apex of the glume (C. purpurascens) . Teeth of the flowering-glume sometimes 0.5 mm. long(C purpurascens).
13
7. Palea distinctly 3-norved in one spikelet, the mid-nerve as prominent as the keels but not reaching the apex; in this case the prolongation of the rachilla unusu- ally well developed (C. breviseta). Palea o nerved in one specimen, the nerves nearly equidistant and equally prominent to near the apex, but only the normal keels quite reaching the apex, the palea being strongly grooved only near the base and the strongly developed prolongation of the rachilla slightly lateral ; in other spikelets the two outermost nerves anastomosing with the normal keel nerves and the central one disappearing below the middle of the palea (C. aleatica).
The frequency of two-flowered spikelets in this genus, often when the plant is apparently quite healthy, shows how slight the dividing line between the tribes Agrostidew and Avenem may become.
HYBRIDISM.
To what extent the species of Calamagrostis hybridize with One another, if at all, can only be determined by actual experiment or the closest kind of field observation. It is not improbable that natural hybrids occur in the genus. Apparent crosses of 0. hyperborea with G. purpurascens, C. hyperborea with G. langsdorffii and G. canadensis with C. sulesdorjii, are represented in the United States National Herb- arium.
SPECIES EXCLUDED.
Galamagrostis gigantea Nutt., Trans. Am. Phil. Soc. (II) 5: 143 (1837 ) — GalamovilJ 'a longifolia (Hook.) Scribn.
Galamagrostis andina Butt. PI. Gamb., Journ. Acad. Phila. (II) 1: 187 (1848), not identifiable from the description, but certainly not a
Calamagrostis.
Agrostis wquivalvis Trim, referred to Deyeuxia by Beutham on account of the minute usually hairy prolongation of its rachilla, is in all other respects an Agrostis and should be retained in that genus.
NORTH AMERICAN SPECIES OF CALAMAGROSTIS.
ANALYTICAL KEY TO THE SPECIES.
1. Awn strongly geniculate, exserted, callus hairs shorter than the flowering glume
2
1. Awn straight or nearly so, included, callus hairs not much shorter than the
flowering glume. 24
2. Awn much exceeding empty glumes 3
2. Awn shorter than or not much exceeding empty glumes 11
3. Panicle loosely flowered, rays spreading 4
3. Panicle densely flowered, rays appressed 8
4. Plants not csespitose 5
4. Plants strongly Ciespitose 6
5. Tall (o to 14 dm. high), flowering glume strongly granular-scabrous
1. C. bolanderi.
5. Low (1.5 to 3 dm. high), flowering glume scabrous, but not granular
2. C. deschampsioides.
6. Leaves of innovations short, not nearly equaling the dark purple panicle, plant
soft - 7
14
6. Leaves of innovations long, often equaling or surpassing the whitish panicle,
plant rather rigid 5. C. howellii.
7. Innovations extravaginal, rootstock strong, creeping, spikelets 4 to 5 mm. long
3. C. lemmoni.
7. Innovations intravaginal, rootstock none, spikelets 3.5 to 4 mm. long
4. C. breweri.
8. Strongly csespitose, rather hard in texture, leaf-blades strongly involute 9
8. Not cjespitose, soft in texture, leaf-blades flat 9. C. tweedyi.
9. Innovation leaves very long (often surpassing the culm), filiform, not rigid,
spikelets 8 to 10 mm. long 6. C. foliosa.
9. Innovation leaves short, not filiform, somewhat rigid, spikelets 4 to 8 mm. long. 10 10. Marcescent basal sheaths closely investing culm for J to J its length, panicle very dense, empty glumes more or less scabrous all over. . 7. C. purpurascens.
10. Marcescent basal sheaths much shorter, loose, panicle somewhat interrupted,
empty glumes nearly glabrous except on keel 8. C. vaseyi.
11. Panicle open, loosely flowered, the rays wide-spreading 12
11. Panicle contracted, usually densely flowered, the rays erect or nearly so 13
12. Plant low (1.5 to 3 dm.), soft, panicle dark purple, few-flowered, empty glumes
not sharply keeled 2. C. deschampsioides.
12. Planttall, ratberhard, paniclepale, empty glumessharply keeled. 11. C.aleutica.
13. Spikelets strongly compressed, empty glumes sharply keeled 14
13. Spikelets not strongly compressed, empty glumes not strongly keeled 19
14. Plant low (not exceeding 5 dm.), leaves short, almost filiform, rigid, erect.
Rocky Mountain species 10. C. montaneusie.
14. Tall, leaves longer and wider. Pacific coast species 15
15. Panicle narrow and dense, spike-like, leaves all strongly involute 16
15. Panicle wider, not spike-like, usually loosely flowered, leaves often flat
11. C. aleutica.
16. Plant yellowish, sheaths not bearded at summit 17
16. Plant not yellowish, sheaths bearded at summit 18
17. Culms and sheaths strongly twisted toward base, panicle oblong-lanceolate, not
interrupted, spikelets 5 to 5.5 mm. loug 12. C. vilfaeformis.
17. Culms and sheaths not or but slightly twisted, panicle oblong, much interrupted
toward base, spikelets 4.5 to 5 mm. loug 13. C. koelerioicles.
18. Panicle strict, spikelets about 6 mm. long 14. C. angusta.
18. Panicle usually somewhat flexuous, spikelets 4 to 5mm. long. 15. C. subflexuosa.
19. More or less ca;spitose, culms usually rather rigid, leaf-blades usually involute.
Rocky Mountains and westward 20
19. Not csespitose, culms not rigid, leaf-blades usually flat. Eastern species 22
20. Lower leaves in a dense tuft, short, rather rigid, strongly involute, spikelets
about 4 mm. long 16. C. fasciculata.
20. Lower leaves rarely forming a dense tuft, usually elongated, not rigid 21
21. Panicle narrow, spiciform, usually red-purple 17. C. rubescens.
21. Panicle wider, rarely spiciform, usually pale-green 18. C. suksdorfii.
22. Sheaths rarely bearded at summit, panicle more or less tinged with lurid purple,
empty glumes rather thick 19. C. breviseta.
22. Sheaths usually bearded, panicle not lurid purple, empty glumes thin 23
23. Spikelets 4 to 6 mm. long, callus hairs very sparse, the longer J to f as long as
glume, palea equaling or very nearly equaling glume 20. C. porteri.
23. Spikelets 3.5 to 4 mm. long, callus hairs rather copious, the longer about J as
long as glume, palea considerably shorter than glume 21. C. nemoralis.
24. Panicle open, the lowerrays wide-spreading, leaf-blades flat, callus hairs copious,
nearly or quite equaling glume 25
24. Panicle more or less contracted 28
25. Spikelets 4 to 6 mm. long, empty glumes narrow, sharp-acuminate, awn stout,
attached below middle, considerably exceeding glume.. .. 22. C. langsdorffii.
15
25. Spikelets 4 mm. long or less 26
26. Spikelets 3 to 4 mm. long, panicle usually loosely flowered. 27
26. Spikelets 2 to 2.5 mm. long, panicle rather densely flowered. 25. C. macouniana.
27. Panicle branches conspicuously flexuous, awn attached near apex of glume and
considerably exceeding it 23. C. blanda.
27. Panicle branches not conspicuously flexuous, awn attached usually near middle,
equaling or slightly exceeding it 24. C. canadensis.
28. Leaf-blades flat or nearly so, panicle not spiciform 29
28. Leaf-blades strongly involute 33
29. Empty glumes not setaceous-pointed, prolongation of rachilla bearded its whole
length, caryopsis glabrous 30
29. Empty glumes setaceous-pointed, prolongation of rachilla bearded only near
summit, caryopsis pubescent especially at summit 30. C. ciunoides.
30. Callus hairs copious, f as long to longer than glume 31
30. Empty glumes narrow lanceolate, callus hairs sparse, | as long as glume or less
32
31. Not csespitose, longer callus hairs not equaling glume 26. C. scribneri.
31. Strongly csespitose, longer callus hairs considerably exceeding glume
27. C. alaskana.
32. Tall (9 to 12 dm.), sheaths bearded at summit 28. C. cusickii.
32, Usually low (not exceeding 8 dm.), sheaths not bearded 29. C. scopulorum.
33. Culms and usually almost filiform leaf-blades soft, not rigid, plant not csespitose
or not strongly so 34
33. Culms and (usually) wider leaf-blades hard, more or less rigid 36
34. Panicle dense, narrow, spike-like 35
34. Panicle loosely flowered, open, flexuous 31. C. laxiflora.
35. Spikelets 2.5 to 4 mm. long, empty glumes thin, acute sharp-acuminate
32. C. neglecta.
35. Spikelets 2 mm. long, empty glumes thickish, barely acute . . 33. C. micrantha.
36. Panicle elongated, not spike-like, rather loosely flowered, plant tall, not cajspi-
tose or but slightly so 37
36. Panicle usually short, dense, spike-like, plant usually low, strongly csespitose. 38
37. Ligule 4 to 6 mm. long, hairs of callus about equaling glume. 34. C. inexpansa.
37. Ligule 2 to 3 mm. long, hairs of callus i to f as long as glume. 35. C. californica.
38. Not exceeding 5 dm. in height, panicle Blender, much interrupted toward base,
awn not nearly equaling glume 36. C. labradorica.
38. Panicle thick, not conspicuously interrupted, awn nearly equaling to slightly
exceeding the floweriug glume 39
39. Taller (4 to 12 dm.), panicle larger (5 to 20 cm. long), empty glumes merely firm-
membranous, oblong-lanceolate to ovate-lanceolate, short-acuminate
37. C. hyperborea.
39. Low (not exceeding 5 dm. ), panicle-small (4 to 6 cm. long), empty glumes almost coriaceous, broad ovate, acute 38. C. crassiglumis.
A. Awn strongly geniculate, conspicuously exserted, callus hairs usually much shorter than
the flowering glume. a. Awn greatly exceeding the empty glumes. * Panicle loosely flowered, branches spreading. t Not caispitose, panicle dark-broivn purple. 1. Calamagrostis bolanderi Thurb.; S. Wats. Bot. Calif. 2: 280 (1880). C. varia. Boland. in Thurb. 1. c, not D. C. Deyeuxia bolanderi Scribn./Bull. Torr. Club. 10 : 8 (1883). Northern California.
Type specimen collected in swamps, Mendocino County, by H. N. Bolander (No. 6471 in part).
16
Specimens examined. — California: (Bolander 6471 in part) ; Mendocino (Pringle),
1892; (Kellogg and Harford 1092), 1868-69. G. bolanderi is unique in the peculiar granular roughening of its flowering glumes.
It has no near relative among the North American species.
2. Calamagrostis deschampsioides Trin. Ic. Gram. 3 : t. 354 (1836). C. obtusata
Turcz., Bull. Soc. Nat. Mosc. 29 : Pt. 1,26(1856). Deyeuxiadeschampsioides Scribn , Bull. Torr. Club 10: 8 (1883). Bering Sea region.
Type specimen collected in Kamchatka.
Specimens examined. — Pribilof Islands: St. Paul Island (Merriam), 1891; St. Paul Island (Macoun 16224 G. S. C), 1897. Kamchatka: Petropaulovski (C. Wright), 1853-56.
ft Strongly cajsjtitose.
t Plant soft, leaves of innovations short, not nearly equaling the dark purple panicle.
3. CALAMAGROSTIS LEMMONI Kearney, sp. n.-
Plants with slender, creeping rootstocks, numerous, short, erect, or ascending innova- tions, many slender erect culms 2.5 to 4.5 dm. high, and dark purple panicles. Culms often slightly geniculate, with a few short, loose, thin, mareescent sheaths at base ; interuodes 2, finally much exceeding their sheaths, the uppermost nearly twice as long as both sheath and blade. Sheaths loosely embracing the culm, thin, glabrous. Ligule 2 to 4 mm. long, thin, whitish, nearly glabrous, lower ones broadly truncate, upper ones narrowed from the base to the acute apex. Blades 2 to 20 cm. long, 1 mm. or less wide, strongly involute, especially toward the setaceous tip, somewhat spreading, flaccid, scabrous on the margins and upper surface. Panicle 4 to 8 cm. long, 1 to 2 cm. wide, oblong-lanceolate, acu- minate, contracted, loosely few-flowered, erect, often somewhat flexuous; rachis slender, somewhat flexuous, glabrous or very nearly so, dark purple, its lowest internode 1.5 to 2 cm. long ; branches spreading at a small angle (45 degrees or less) to nearly erect, slender, somewhat flexuous, dark purple, glabrous or the upper slightly hispidulous, secondary branches (usually) and shorter primary branches (often) 1-flowered, lower primary branches in 3's to 5's, the longest 2 to 3 cm. long. Spikelets 4 to 5 mm. long. Empty glumes lanceolate to ovate- lanceolate, acute or acutish, not strongly keeled, thin-membranous, dark red- purple, sparsely and minutely scabrous on the keel toward apex or entirely glabrous, equal or the first slightly longer. Flowering glume considerably (often nearly 1 mm.) shorter than the second empty glume (rarely nearly equaling it), oblong-ovate, broadly truncate, thin-membranous, with hyaline edges and tips, somewhat scabrous on the back, the nerves conspicuous, the lateral ones extended into slender, unequal terminal awns 0.3 mm. long or less ; awn attached one-sixth
1 Geological Survey of Canada.
2 In the descriptions of the species the following points are to be noted : Length of innovation refers to length of branch and sheathes, excluding the blades. Number of internodes of the culm refers to those of conspicuous length above the base and below the inflorescense. Width of leaf-blade and of the panicle denotes the greatest width. Shape of panicle refers to the outline in the dried plant, fresh material of none of the species having been available. Length of spikelets means length of the longest empty glume, and excludes the awn. The awn is described from dried mate- rial, and does not, of course, exhibit exactly the same characters as in the fresh or living state.
In giving the geographical range of the species the typical form alone is included, that of the several varieties being separately stated. Data as to localities, altitude, and latitude are taken for the most part from the collectors' labels, without attempt at verification.
17
above tlie base, much exceeding the glume, 5 to 6 mm. long, stout, minutely scabrous, strongly geniculate near the middle, the lower part somewhat twisted, the upper part divergent at an angle of 45 degrees, exserted, straight or slightly flexuous, dark purple. Palea about 1 mm. shorter than the flowering glume, ovate-oblong, truncate, entire or obscurely bidentate, glabrous. Anthers about 2 mm. long. Caryopsis nearly 2 mm. long. Callus hairs sparse, the dorsal ones very short, the lateral ones one-fourth to one-half as long as the flowering glume. Prolongation of the rachila copiously bearded, with its hairs exceeding the palea and sometimes equaling the flowering glume.
Type specimen in the United States National Herbarium, collected in California by J. G. Lemmon, in 1875.
Intermediate between C. deschampsioides and C. breweri. From the former it diifers in its more csespitose habit, culms taller; ligule usually longer ; panicle more contracted; spikelets more numerous and smaller; glumes narrower; flowering glume usually considerably shorter than the empty glumes and conspicuously awn-toothed; awn longer and attached nearer the base, palea shorter, anthers smaller, callus hairs usually shorter, and prolongation of the rachilla longer and with much longer hairs. From C. breweri it is distinguished by its creeping root- stock, innovations extravaginal, leaf-blades longer, panicle larger, contracted, with less divergent branches, spikelets larger and more numerous, flowering glume usually considerably shorter than the empty glumes, awn usually longer, and palea considerably shorter than the flowering glume.
4. Calamagrostis breweri Thurb.; S. Wats. Bot. Calif. 2: 280 (1880). Deyeuxia
breweri Vasey Descr. Cat. Grasses U. S. 50 (1885). Mountains of California.
Type specimen collected near Carsons Pass by W. H. Brewer (2128).
Specimens examined. — Sierra Nevada Mountains: Altitude 2,946 meters (Brewer 2128). Upper Tuolumne River: (Bolander 6098), 1867.
A peculiar species, with much the aspect of Festuca ovina L., distinguished from all other North American members of the genus by its strictly intravaginal inno- vations.
tt Plant rather stiff, leaves of innovations long, often equaling or surpassing the whitish
panicle.
5. Calamagrostis howellii Vasey ; Coult. Bot. Gaz. 6: 271 (1881). Deyeuxia how-
ellii Vasey Descr. Cat. Grasses U. S. 51 (1885). Washington and Oregon.
Type specimen. collected in Oregon by T. J. Howell.
Specimens examined. — Washington: Larm Eiver, West Klickitat County (Suks- dorf 13), 1883. Oregon : Multonomah Falls (Bolander); Columbia Eiver below the Cascades (Howell, 356), 1880; Sandy River (Henderson, 13).
A specimen from Washington, collected by Suksdorf, has the contracted but loosely- flowered panicle of the extreme form of C. vaseyi. In habit and other characters, however, it is not distinguishable from G. howellii.
** Panicle densely flowered, branches oppressed. t Strongly cwspitose, rather hard in texture, leaf -blades strongly involute.
6. CALAMAGROSTIS FOLIOSA Kearney, sp. n. C. sylvatica longifolia Vasey
Monog. Grasses U. S., Contr. U. S. Herb. 3:83 (1892), not C. longifolia Hook. A somewhat glaucous species with slender rootstocks, numerous short, erect, or ascending innovations whose leaves often equal or surpass the culms, and dense epiciform panicles. Culms 2 to 4.5 dm. high, nearly erect, slender, somewhat flexuous; internodes usually 3, finally greatly exceeding their sheaths. Sheaths closely embracing the culm, slightly scabrous above. Ligule 4 to 5 mm. long, truncate rather firm, whitish, somewhat scabrous. Blades 1 to 3 dm. long, 4 20101— No. 11 2
18
mm. or usually much less in width, strongly involute and usually almost filiform, strongly scabrous on the margins and upper surface, glabrous beneath. Panicle 6 to 10 cm. long, 1 to 2 cm. wide, dense, spike-like, somewhat interrupted toward base, pale, purple, or greenish; rachis slender, somewhat scabrous above, its longest internode 1 to 2 cm. long; branches slender, hispidulous, erect and appressed, the lower primary branches mostly in 5's. Spikelets 8 to 10 mm. long. Empty glumes narrow-lanceolate, gradually attenuate from the base to the sharp- acuminate apex, thin, minutely scabrous, the first slightly longer. Flowering glume about 2 mm. shorter than the first empty one, lanceolate, acutish, minutely scabrous, of about the same texture as the empty glumes, the 4 lateral nerves extended into straight, erect, very slender awns of unequal length, the longest 1 to 2 mm. ; dorsal awn attached about one-fifth above the base (but sometimes as high as one-third), 10 to 15 mm. long, slender, bent near the middle, the upper part much exserted and spreading at au angle of about 45 degrees. Palea four- fifths to five-sixths as long as the flowering glume (or occasionally nearly equal- ing it) considerably narrower, acute, with nerves somewhat excurrent, nearly glabrous. Callus hairs rather sparse, very unequal, the longer ones one-third to one-half (usually one-half) as long as the flowering glume. Prolongation of the rachilla with its rather copious hairs about equaling the palea. California.
Type specimen in the United States National Herbarium, collected by H. N. Bolan- der (6470) in the Mattole district, Humboldt County.
C. foliosa is intermediate between C. howellii and C. purpurascens. From the former, its nearest ally, it differs in its contracted, dense panicle and usually larger spikelets. From C. purpurascens it is easily differentiated by its slender, some- what ftexuous culms, very long and almost filiform leaf-blades, considerably larger spikelets, much longer, more slender, less abruptly bent and nearly glab- rous awn, prominent aristate teeth to the flowering glume and proportionately longer callus hairs.
7. Calamagrostis purpurascens E. Br. ; Eichards, App. Frankl. Journ., 731 (1823). Deyeuxia purpurascens Kunth, Eev. Gram. 1: 77 (1835). Calamagrostis sylvatica A. Gray, Proc. Am. Acad. 6 : 80 (1866). Deyeuxia sylvatica Vasey, Descr. Cat. Grasses U. S., 51 (1885). Calamagrostis sylvatica americana Vasey, Monog. Grasses U. S., Contr. Nat. Herb. 3: 83 (1892). C. sylvatica purpurascens Thurb. ; Vasey, 1. c.
East Greenland to Alaska, and southward along the higher mountains to South Dakota, Colorado, and California.
Type specimen collected iu the Barren Lands between Point Lake and the Arctic Sea by Dr. Bickardson.
Specimens examined. — Greenland: Disco (Warming, Th. Fries, Holm); (Weth- erill 4), 1894. Arctic coast: (Eichardson, 59, 62, 67, Herb. Hook.). Rocky Moun- tains: Latitude 39 to 41 degrees (Hall and Harbour 624), 1862. South Dakota: Black Hills, altitude 1,670 to 1,824 meters (Eydberg 1130), 1892. Assiniboia: (Douglas). Montana: White Sulphur Springs, Belt Mountains, altitude 2,070 to 2,736 meters (Scribner 362), 1883; Spanish Creek, Gallatin County (Tweedy 1022); Baldy Peak near Bozeman (Eydberg 2224; Shear 468), 1895; Belt Pass (Eydberg 3313J), 1896; Barker, altitude 2,128 meters (Eydberg 3373), 1896; Black Hawk (Eydberg 3296), 1896; Spanish Peaks, altitude 2,432 meters (Eydberg 3074), 1896; Belt Mountains (E. S. Williams 596), 1888. Wyoming: Laramie Peak (A. Nelson 1627), 1895. Colorado: Denver (Coulter), 1873; (Wolf 398, 587), 1873 ; Pen Gulch (Vasey) ; Lake Eanch (G. H. French) ; Silver Plume, altitude 3,952 meters (Shear 691, 696; Eydberg 2470), 1895; Georgetown, altitude 2,512 meters (M. E. Jones 500), 1878; Georgetown (Shear 614), 1895; Clear Creek (Parry 365, 368), 1861; Pikes Peak (Canby, T. A. Williams 2170, 2188, 2188£), . 1896; Manitou, El Paso County, altitude 3,040 meters (Clements 42), 1896; Colorado Mine (Eydberg 2380), 1895; Mount Princeton, altitude 3,344 meters
19
(Sheldon 607) ; Beaver Creek, Laramie County, altitude 3,344 meters (Pammel), 1896; Buenavista, Chaffee County, altitude 3,040 to 3,192 meters (Clements 309; Shear 1016), 1896. Alberta: Sheep Mountains (Macoun, 13113, G. S. C.)i Morley (Macoun), 1885. Idaho: (Wheeler Expedition 798, in part), 1871. Utah: (Ward) 1875. Nevada: Humboldt Mountains, altitude 3,040 meters (Watson 1291). Alaska: (McDonald), 1864; along Yukon Eiver (Funston), 1893. North- west Territory: Tagish Lake, latitude 62° (DawBon), 1887; Lewis River (Daw- son), 1887; Fort Pelly Banks (Dawson), 1887; Bennetts Lake (Ogilvie), 1887. British Columbia: Donald (Macoun), 1885; Kicking Horse Lake (Macoun), 1885; Dease Eiver (Dawson), 1887. Washington: Wenatchie region (Tweedy 650), 1883; Mount Stuart, altitude 2,280 to 2,432 meters (Sandberg and Leiberg 825), 1893. Oregon: Baker City (Nevius), 1873. California: Redwoods (Bolander); Mount Dana, altitude 3,748 meters (Bolander 5071).
C. purpurascens has been confused by some authors with the very distinct C. sylva- Hca (Schrad. ) D. C. of Europe, which is a taller, less ca^spitose grass with broad, flat, lax, leaf-blades; longer and looser panicle; and broader, less pointed, thinner, and nearly glabrous empty glumes.
CALAMAGROSTIS PURPURASCENS ARCTICA (Vasey) Kearney, n. comb. C. Arctica Vasey, 111. ST. Am. Grasses 2 : Pt. 2, No. 55 (1892).
Depauperate, perfectly smooth and glabrous up to the inflorescence. Culms ascend- ing, 18 cm. or less in length. Leaf-blades short, the longest 7 cm. long, compara- tively broad. Panicle 2 to 3 cm. long, barely 1 cm. wide. Spikelets about 5 mm. long. Awn short, about 7 mm. long.
Type specimen in the United States National Herbarium, collected on St. Paul Island, Pribilof Islands, by James M. Macoun (38), July 31, 1891.
8. Calamagrostis vaseyiBeal, Grasses N. Am. 2 : 344(1896). C. purpurascens Vasey,
Monog. Grasses IT. S., Contr. TT. S. Nat. Herb. 3 : 83 (1892), not R. Br. Mountains of Washington and Oregon.1
Type specimen collected in the Cascade Mountains, Washington, by G. R. Vasey, in 1889.
Specimens examined. — Washington: Cascade Mountains (G. R. Vasey), 1889; Goat Mountains (O. D. Allen 177), 1896; Skamania County (Suksdorf, 201, 909, 1025), 1886-1890; South Olympia Mountains, altitude 1,368 meters (Henderson 1855), 1890; (Brandegee 177), 1882. Oregon: Siskiyou Mountains (Howell), 1887.
Imtermediate between C. purpurascens and C. howellii, the type specimens with shorter awns and denser panicles being nearer the former. From C. purpurascens it differs in being very strongly caespitose from a hard, knotted rootstock, with less rigid and more slender culms; marcescent basal sheaths very short and less closely enveloping the culm; panicle usually looser, fewer flowered, broader, less colored, with longer, straighter, more slender, and less scabrous branches; thinner and smoother, less strongly keeled empty glumes ; usually much longer and more slender awn attached higher ; and longer callus hairs. From C. howellii it differs in its shorter, flatter, and more rigid leaf-blades; and contracted, usually purplish panicle. The spikelets are much like those of the European C. sylvatica, but in other respects the plant is quite different.
tt Not ewspitose, soft in texture, leaf-blades flat.
9. Calamagrostis tweedyi Scribner; Vasey, Monog. Grasses U. S., Contr. U. S. Nat.
Herb. 3 : 83 (1892). Deyeuxia tweedyi Scribn., Bull. Torr. Club 10 : 64 (1883). Type specimen collected in the Cascade Mountains, Washington, by Frank Tweedy. Specimens examined. — Washington: Cascade Mountains (G. R. Vasey), 1889. A unique species with no near ally.
1 Northward to Alaska according to Beal, but this is probably a mistako.
20
b. Awn shorter than or not much exceeding the empty glumes. * Spikelets strongly compressed, empty glumes sharply keeled. t Plant low, leaf-blades almost filiform — Rocky Mountain species.
10. Calamagrostis montanensis Scribn. ; Vasey, Monog. Grasses U.