NORTHWESTERN NATURALIST 84:1–19 SPRING 2003

GROUSE OF THE LEWIS AND CLARK EXPEDITION, 1803 TO 1806

FRED CZWICKEL1 Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 CANADA

MICHAEL ASCHROEDER Washington Department of Fish and Wildlife, PO Box 1077, Bridgeport, Washington 98813 USA

ABSTRACT—Members of the Lewis and Clark expedition produced written descriptions of many species of wildlife, including 6 species of grouse. We reviewed the accounts of 83 grouse observations, plus summary descriptions and indirect references to grouse by Meriwether Lewis, William Clark, and 3 of their sergeants. We then assigned them to species based on described characteristics, known distributions, habitat, behavior, and/or other written clues. Fifty-nine (71%) observations were con- sidered relatively unambiguous as to identity because of a bird’s morphology and appearance, a descriptive name, a non-overlapping range, and/or other clues. These included greater prairie-chick- ens (Tympanuchus cupido), sharp-tailed grouse (T. phasianellus), greater sage-grouse (Centrocercus uro- phasianus), blue grouse (Dendragapus obscurus), spruce grouse (Falcipennis canadensis), and ruffed grouse (Bonasa umbellus). Other observations were assigned to ‘possible’ species based on available evidence. Our evaluation of Lewis’s and Clark’s written descriptions differed in some cases from earlier reviews. Most notable differences involved blue, spruce, and ruffed grouse and we offer sug- gestions for changes. We also used journal records to compare pre-settlement (by Euro-Americans) distributions of grouse as indicated by the journal records with those today and to speculate on changes in abundance. Greatest changes have occurred in distributions and abundances of the 3 prairie grouse, species whose habitats have been most impacted by settlement. Other highlights of the expedition include the 1st written descriptions of blue grouse and greater sage-grouse and of undescribed subspecies of sharp-tailed, spruce, and ruffed grouse.

Keywords: Lewis, Clark, tetraonines, grouse, distribution, abundance

In the early 1800s, Meriwether Lewis and with abundant observations of fauna and flora. William Clark led an expedition of exploration The faunal list included 6 species of tetraoni- from the mouth of the Missouri River to its nes: greater prairie-chicken (Tympanuchus cupi- headwaters, across the continental divide, do pinnatus), sharp-tailed grouse (T. phasianellus down the Columbia River to the Pacific Ocean, jamesi and T. p. columbianus), greater sage- and returned along a similar route (Fig. 1). The grouse (Centrocercus urophasianus urophasianus principal stated objective was to seek a feasible and C. u. phaios), blue grouse (Dendragapus ob- water route from the Mississippi River to the scurus pallidus, D. o. fuliginosus, and D. o. sierrae), Pacific. A secondary mandate was to document spruce grouse (Falcipennis canadensis franklinii), natural history and biological resources along and ruffed grouse (Bonasa umbellus umbelloides the route. To fulfill these objectives, Lewis was and B. u. castanea). Greater sage-grouse, blue given several months of schooling in natural grouse, and subspecies T. p. columbianus, F. c. history and other sciences prior to start of the franklinii, and B. u. castanea were new to science. expedition (Moulton 1986; Botkin 1995). We assigned subspecific names on the basis of Collectively, Lewis and Clark and 3 of their current taxonomy. Henceforth, we use ‘prairie- sergeants maintained almost daily records, chicken’and ‘sage-grouse’ to refer to the great- er prairie-chicken and greater sage-grouse, un- less necessary for clarity. 1 Present address: Box 81, Manson’s Landing, British Natural history observations in expedition Columbia V0P 1K0 CANADA. journals have been reviewed by Coues (1893;

1 2 NORTHWESTERN NATURALIST 84(1)

FIGURE 1. Primary outbound (westward) and return (eastward) routes of the Lewis and Clark expedition. Primary routes are illustrated with a solid line, major deviations with dashed lines, and state boundaries with dotted lines. Locations of winter camps and major delays for weather, portages, and canoe building are shown.

Vols 1 to 4), Burroughs (1961), Cutright (1969), knowledge. Other objectives are to compare the Holmgren (1984), and Moulton (1986–1997; probable early distributions and abundances of Vols 2 to 11), and identifications of species of the different species with those today. grouse are not always consistent among au- Except for 1 reference to ‘sharp-tailed grouse’, thors. This is partly caused by the plethora of none of the current common or scientific names common names used by expedition members of NorthAmerican grouse was used in the jour- to refer to individual species, a paucity of basic nals (Table 1) and this sometimes makes it dif- information about them, in some cases inade- ficult to identify species to which journal au- quateorambiguous descriptions, and misin- thors refer. For example, references to sage- terpretations by reviewing authors. One of our grouse included mountain cock, cock of the objectives is to resolve identifications we be- plains, prairie hen, heath hen, prairie fowl, and lieve are questionable in the light of current long-tailed heath cock, among others; different

TABLE1. Common names of grouse used by members of the Lewis and Clark expedition. Current spellings are used in this table.

Current common name Names used by expedition members Greater prairie-chicken Heath hen, prairie fowl, prairie hen, grouse, prairie fowl common to the Illinois Sharp-tailed grouse Heath hen, prairie fowl, prairie hen, pointed-tail prairie fowl, pointed- tail prairie hen, grouse, pheasant, sharp-tailed grouse Greater sage-grouse Heath hen, prairie fowl, prairie hen, mountain cock, cock of the plains, prairie cock, fowl of the plains, long-tailed heath cock, long-tailed grouse, large pheasant Blue grouse Black pheasant, large black pheasant, large black and white pheasant, dark brown pheasant, large dark brown pheasant, large speckled pheasant, large domminecker (or dommanicker) Spruce grouse Small speckled pheasant, small black pheasant, small brown pheasant, small pheasant Ruffed grouse Common pheasant, pheasant of the Atlantic states, pheasant of the United States, reddish brown pheasant, partridge SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 3 names and spelling were often used by a single ferences we had between us after discussions of author. Further, heath hen, prairie hen, and theevidence. prairie fowl all appear at least once to refer to We sometimes used other clues to help assign otherwise clear observations of prairie-chick- identities to birds. Pattern of use of ‘grouse’ and ens, sharp-tailed grouse, and sage-grouse. It is ‘pheasant’, and occasionally other common no wonder later authors have differed in iden- names, revealed some distinct tendencies on tifying some of the birds encountered. the part of Lewis and Clark. Clear references to prairie-chickens and sharp-tailed grouse were METHODS most often identified as grouse, not pheasants (grouse in 31 journal entries, pheasant in 2). All We examined all grouse observations of ex- sage-grouse were identifiable by other clues pedition members as indexed in the recent and and were twice referred to as pheasants, in 1 monumental republication of the Lewis and instance as ‘‘large pheasents’’ (Clark, 18 Octo- Clark journals by Moulton (Vols 2 to 11 and 13). ber 1805). Clear references to ruffed, blue, and This work includes the unedited writings of spruce grouse were referred to as pheasants in Lewis and Clark (Vols 2 to 8) and sergeants 44 entries and grouse in 1. We did not use those John Ordway (Vol 9), Patrick Gass (Vol 10), and clues to identify birds reported only by the ser- Joseph Whitehouse (Vol 11), with many insight- geants because they were less consistent with ful annotations by Moulton. Unless otherwise names. In direct quotes from the journals, we specified, all quotations are from that work. We retain spelling, grammar, punctuation, and also read in their entirety Moulton’s Volumes 4 capitalizations of the authors as reported in and 5, Fort Mandan to the lower Columbia Riv- Moulton. Items or comments in brackets are er,tosearch for references that might not have ours except that those with asterisks are from beenindexed and found none. We believe we Moulton. have examined virtually all, if not all, referenc- Finally, we compared locations of birds we es to grouse in the Moulton edition of the jour- consider identifiable with current distributions nals. and speculate on changes from presettlement We summarized all observations and sum- (by Euro-Americans) distributions and abun- mary descriptions by date, observer, and spe- dances. Locations must be considered general cies(as identified in Moulton). Journals of the because the only information available in most different authors often included references to cases is the approximate vicinity of campsites, the same observation (up to 4 authors for 1 ob- with the party often covering up to 30 km or servation), many of which appear to have been more in a given day. copied from others. We treat multiple referenc- es to the same observation as 1 and in such cas- BACKGROUND es lean most heavily on those of Lewis because of his position as naturalist and because he Aprincipal objective of the expedition was to usually provided more clues to identity than find a water route across the continent and others. There were at least 83 observations of most travel was concentrated along the 2 major grouse in the journals, plus indirect references rivers, outbound or westward up the Missouri and summary descriptions of grouse. River and down the Columbia River and the re- We visited many areas (where possible) at verse on the return journey, although Clark and which expedition members had reported hav- some members of the party returned down the ing encountered grouse to provide us with Yellowstone River to its confluence with the background on local habitats. We each then Missouri. Numbers and kinds of observations compared identities reported by Coues (1893), were undoubtedly related in part to speed of Burroughs (1961), Cutright (1969), Holmgren travel, faster downstream than upstream. Prin- (1984), and Moulton with descriptions in the cipal overland routes involved approaching journals, known distributions, habitats, behav- and crossing the Rocky and Bitterroot moun- iors, morphologies, and plumages of the differ- tains. Most exploration was concentrated in the ent species. We then independently assigned outbound journey, with returning home the current species names to each observation and/ principal objective in the spring and summer of or general description and resolved the few dif- 1806. Seasons in which particular areas were 4 NORTHWESTERN NATURALIST 84(1) traversed are important considerations in eval- flewof[f]* and having no gun with me did not per- uating journal entries. sue it’’ The expedition was dependent upon living 2 September 1806.—[Clark, near confluence of the offthe land, and while most of the party was Jamesand Missouri rivers]: ‘‘I saw 4 prairie fowls Common to the Illinois, those are the highest up engaged in moving canoes and supplies along which have been Seen.’’ the rivers, some members were regularly del- egated as hunters. Lewis, especially, and Clark That Lewis and Clark made no effort to de- spent much time exploring on land while sup- scribe this species in detail is additional evi- plies were being moved by water. They and the dence of their acquaintance with it. hunters likely would have been flanking the rivers ahead of the main party much of the Sharp-tailed Grouse time, not more than 5 to 10 km from them in Three journal entries, especially, are helpful most cases. Except when crossing the moun- in providing identifications of sharp-tailed tains, faunal records are mainly from or near grouse: the river valleys. Other exceptions involve oc- 1 October 1804.—[Clark, in present Dewey Coun- casional short stops to rest, replenish food sup- ty, South Dakota, a few km above the mouth of the plies, or talk with natives; over-winter sojourns Cheyenne River]: ‘‘worthy of remark that the at Camp Dubois, Illinois (1803 to 1804), Fort Grouse or Prarie hen is Booted, the Toes of their Mandan, North Dakota (1804 to 1805), and Fort feet So constructed as to walk on the Snow, and the Clatsop, Oregon (1805 to 1806); about 30 days Tail Short with 2 long Stiff feathers in the middle.’’ while moving supplies around the Great Falls 15 April 1805.—[Lewis, Little Knife River area, of the Missouri River, Montana; 11 days near Mountrail County, North Dakota]: ‘‘I also met with Orofino, Idaho, while resting and building ca- great numbers of Grouse or prairie hens as they are noes to move down the Clearwater and Colum- called by the English traders of the N.W. these bia rivers; and about 6 weeks at, or near, Ka- birds appeared to be mating; the note of the male is kuck, kuck, kuck, coo, coo, coo. the male also miah and Weippe Prairie, Idaho, waiting for dubbs[EC:drums]* [NB:with his wings]* some- snow to melt so they could return east across thing like the pheasant, but by no means as loud.’’ the Bitterroot Mountains. Lewis and Clark and 1March 1806.—[Lewis, at Fort Clatsop; descrip- the hunters likely ranged farther from the river tion of sharp-tailed grouse]: ‘‘The Grouse or Prarie bottoms at these times. hen is peculiarly the inhabitant of the Grait Plains of Columbia they do not differ from those of the CLUES TO SPECIES IDENTITIES upper portion of the Missouri, the tail of which is On 1st encountering a new species, Lewis pointed or the feathers in it’s center much longer and Clark usually provided at least some clue than those on the sides. this Species differs essen- as to what it was and from which they often as- tially in the construction of this part of the plum- signed a descriptive name. While in winter age from those of the Illinois [prairie-chicken] which have their tails composed of fathers of equal camp at Fort Clatsop, they prepared more ex- length. in the winter season this bird is booted tensive descriptions of most species seen ear- eventothe 1st joint of it’s toes . . . Their colour is lier. We provide parts of these descriptions and a mixture of dark brown redish and yellowish selected quotations from the journals here that brownand white confusedly mixed in which the provide clues to identities. If other authors’ in- redish brown prevails most on upper parts of the terpretations differ from ours, we discuss why body wings and tail and the white underneath the we have assigned a particular species to an ob- belley and lower parts of the breast and tail.’’ servation. These quotes provide solid support for the clar- Greater Prairie-chicken ity of their observations of this species and for separating it from its closest relative, the prai- Lewis was no doubt familiar with the prairie- rie-chicken. chicken,asubspecies of the eastern heath hen (T. c . cupido)(Moulton 1986). Two journal en- Greater Sage-grouse tries illustrate this point: At least3journal entries provide clues for 16 November 1803.—[Lewis, near confluence of identifying this species: the Missouri and Mississippi rivers, 1st reference to any grouse]: ‘‘saw a heath hen or grows which 5June 1805.—[Lewis, near mouth of the Marias SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 5

River, Montana, 1st contact]: ‘‘I . . . saw . . . a flock ‘‘I also saw two fesants today of a dark brown col- of the mountain cock, or a large species of heath ourmuch larger than the phesant of the U’States hen with a long pointed tail which the Indians in- [ruffed grouse].’’ formeduswerecommon to the Rockey Mountains 1August 1805.—[Lewis, Bull or Tobacco Root ...’’ Mts, Jefferson County, Montana]: ‘‘as I passed 17 October 1805.—[Clark, vicinity of present these mountains I saw a flock of the black or dark Kennewick, Washington]: ‘‘Send out Hunters to brown phesants;...this bird is fully a third larger Shute the Prarie Cock . . . Several of which I have than the common phesant of the Atlantic states killed, they are the Size of a Small turkey, of the [ruffed grouse]. it’s form is much the same ...the pheasant kind, one I killed on the water edge to male has not the tufts of long black feathers on the daymeasured from Beek to the end of the toe 2 feet sides of the neck which are so conspicuous in those 6&¾ inches; from the extremeties of its wings 3 of the Atlantic. their colour is a uniform dark feet 6 inches; the tale feathers is 13 inches long: brown with a small mixture of yellow or yelloish they feed on grasshoppers and the seed of the wild brown specks on some of the feathers particularly plant which is also peculiar to this river and the those of the tail, tho’ the extremeties of these are upper parts of the Missoury somewhat resembling perfectly black for about one inch [birds here are the whins’’ [identified as big sagebrush by Moul- D. o. pallidus which have indistinct, or no, gray tail ton (1988)]. bands, hence the black tips] ...feathersofthetail 2March 1806.—[Lewis, at Fort Clatsop, descrip- are reather longer than that of our phesant or pat- tion of sage-grouse]: ‘‘The Cock of the Plains is tridge as they are Called in the Eastern States; are found in the plains of Columbia and are in Great the same in number or eighteen and all nearly of abundancefrom the entrance of the S.E. fork of the the same length,...[D. o. pallidus usually has 20 Columbia [Snake River] to that of Clark’s River rectrices (70%) but the number varies from 17 to [Deschutes River]. this bird is about 2/3rds the 22 (Zwickel and others 1991)] the flesh of this bird size of a turkey . . . the colour is an uniform mix- is white and agreeably flavored.’’ ture of dark brown reather borde[r]*ing on a dove 3March 1806.—[Lewis, at Fort Clatsop; descrip- colour, redish and yellowish brown with some tion of blue grouse]: ‘‘The large black and white small black specks . . . the tail is composed of 19 pheasant is peculiar to that portion of the Rocky feathers [usually 18 or 20] of which that in the cen- Mountain watered by the Columbia river. at least ter is the longest, and the remaining 9 on each side we did not see them in these mountains until I we deminish by pairs as they receede from the center; reached the waters of that river or since we have ...the tail when foalded comes to a very sharp left those mountains. they are about the size of a point and appears long in proportion to the body well grown hen. the contour of the bird is much ... the motion of the wings is much that of the that of the redish brown pheasant common to our pheasant and Grouse . . . the wings are also pro- country [ruffed grouse]. the tail is proportionably portionably short, reather more so than those of as longand is composed of eighteen feathers of the pheasant or grouse. the habits of this bird are equal length, of an uniform dark brown tiped with much the same as those of the grouse. only that the black.the feathersofthe body are of a dark brown food of this fowl is almost entirely that of the leaf black and white. the black is that which most pre- and buds of the pulpy leafed thorn [likely sage- dominates, and wh[i]*te feathers are irregularly brush, Artemisia sp; Moulton (1988) identifies it as intermixed with those of the black and dark brown Sarcobatus vermiculatus]; nor do I ever recollect see- on every part, but in greater proportion about the ing this bird but in the neighbourhood of that neck breast and belley. this mixture gives it very shrub . . . the gizzard of it is large and much less much the appearance of that kind of dunghill fowl compressed and muscular than in most fowls;... which the hen-wives of our country call domman- theflesh ...is dark, and only tolerable in point of icker.inthe brest of some of these birds the white flavor.—it is invariably found in the plains.’’ predominates most. they are not furnished with tufts of long feathers on the neck as our pheasants Lewis’s description of sage-grouse while at Fort are, but have a space on each side of the neck about Clatsop and measurements by Clark provide a 2 ½ inches long and 1 In. in width on which no number of unambiguous clues. feathers grow, tho’ tis concealed by the feathers which are inserted on the hinder and front part of Blue Grouse the neck . . . they have a narrow stripe of vermil- Four journal entries considered together lion colour above each eye which consists of a clearly identify this species: fleshey substance not protuberant but uneven with anumber of minute rounded dots . . . it feeds on 21 July 1805.—[Lewis, general vicinity of Can- wild fruits, particularly the berry of the sac-a-com- yonFerry Dam, e. of Helena, Montana, 1st contact]: mis, and much also on the seed of the pine and fir.’’ 6 NORTHWESTERN NATURALIST 84(1)

Confusion between this species and spruce cies, most likely D. o. sierrae, based on the lo- grouse is sometimes caused by an authors’ use cation. Also, cervical apteria of sierrae males of black and white pheasant and speckled are yellow, those of interior males red; sierrae pheasant interchangeably for the 2 species, but usually have 18 rectrices (Zwickel and others the confusion is eliminated when the names are 1991), and their tail feathers are tipped with preceded by ‘large’ or ‘small’ because these light grey (bluish white of Lewis). Character- birds are grossly different in size. Coues (1893) istics noted here add support to the argument and Burroughs (1961) identified this as the that the ‘large black and white pheasant’ de- spruce grouse and Holmgren (1984) as the scribed at Fort Clatsop is the blue grouse. Dark ruffed grouse, but we disagree with both iden- brown pheasants are likely females in most cas- tifications. Lewis described this bird as large, es, perhaps large juvenile, or yearling, males in size of a well-grown hen—not a spruce grouse other cases. on this basis. The only clue that might indicate spruce grouse is the vermillion eye stripe for Spruce Grouse that of blue grouse is usually yellow, except in Two entries provide good clues for identify- males in peak excitement during spring dis- ing male and female spruce grouse: play,when it may be bright red. Since this entry 3March 1806.—Lewis, at Fort Clatsop; descrip- was written in March 1806, this could easily be tion of the male]: ‘‘The small speckled pheasant a lapse in memory on Lewis’s part or a confu- found in the same country with that above discri- sion between blue and spruce grouse, which he bed [large black and white pheasant or blue considered very similar except in point of size. grouse], differs from it only in point of size and Nordo18tail feathers fit spruce grouse, which somewhat in colour. it is scarcely half the size of almost always have 16 (Short 1967, Boag and the other; ascociates in much larger flocks and is Schroeder 1992). The bare space on the side of very gentle. the black is more predominant and the the neck (lateral cervical apteria), conspicuous dark bron feathersless frequent in this than the in male blue grouse but not in spruce or ruffed larger species. the mixture of white is also more grouse, is distinctive. Size of the bird, number general on every part of this bird. it is considerably smaller than our pheasant [ruffed grouse] and the of tail feathers, and conspicuous cervical apter- body reather more round. in other particulars they ia far outweigh the ‘vermillion eye stripe’ as differ not at all from the large black and white clues to identity. With reference to Holmgren’s pheasant. this by way of distinction I have called identification, Lewis noted that they do not the speckled pheasant. the flesh of both these spe- have the ‘‘tufts of long feathers on the neck as ciesofparty coloured phesants is of a dark colour our pheasants’’ [ruffed grouse]. This compari- and with the means we had of cooking them not son to ruffed grouse as being different rules out very well flavored.’’ that species. The bird described is clearly the Coues (1893) identified the small speckled blue grouse, not spruce or ruffed grouse. pheasant as a female or juvenile spruce grouse. 16 April 1806.—[Lewis, Rockfort Camp, The Lewis’s description of it, however, is clearly Dalles, Oregon]: ‘‘Joseph Feilds brought me a black that of an adult male, not a female or juvenile. pheasant which he had killed; this I found on ex- The small size, darker coloration [than blue amination to be the large black or dark brown grouse], and vermillion eye stripe (see descrip- pheasant I had met with on the upper part of the tion of female, below) identify this species and Missouri. it is as large as a well grown fowl the iris its sex. of the eye is of a dark yellowish brown, the puple black,the legs are booted to the toes, the tail is 3March 1806.—[Lewis, at Fort Clatsop; descrip- composed of 18 black feathers tiped with bluish tion of female spruce grouse]: The small brown white, of which the two in the center are reather pheasant is an inhabitant of the same country and shorter than the others which are all of the same is of the size and shape of the specled pheasant length. over the eye there is a stripe of a ¼ of an which it also resembles in it’s economy and habits. inch in width uncovered with feathers of a fine or- the stripe above the eye in this species is scarcely range yellow. the wide spaces void of feathers on perceptable, and is when closely examined of a yel- the side of the neck are also of the same colour.’’ lowororrange colour instead of the vermillion of the outhers. its colour is an uniform mixture of This entry, made on their way home, is clearly dark and yellowish brown with a slight mixture of an adult male blue grouse of a coastal subspe- brownish white on the breast belley and the feath- SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 7

ers underneath the tail . . . this is also booted to the Lewis may have been referring to coverage, toes. the flesh of this is preferable to either of the which may extend to or near the toes even others and that of the breast is as white as the though the point of insertion is higher. Other pheasant of the Atlantic coast.—the redish brown characteristics noted are clearly references to pheasant [ruffed grouse] has been previously dis- ruffed grouse. Lewis provided only a brief de- cribed—’’. scription of ruffed grouse, indicating his famil- Spruce, blue, and ruffed grouse caused much iarity with it. confusion among those attempting to identify forest tetraonines, and it is apparent from Lew- SUGGESTIONS FOR CHANGES OF is’s description of the small brown pheasant SPECIES IDENTITIES that he believed it was a different species than Here,weconsider other journal entries and the small speckled pheasant, as noted by Bur- relevant annotations from Moulton for which roughs (1961).The small brown pheasant was we believe assigned species are in error or identified as the ruffed grouse by Coues (1893) questionable. Comments on locations and hab- and Burroughs and as the blue grouse by itats are ours. Holmgren (1984), all of which we consider in 15 December 1803.—[Clark]: ‘‘hunters [k]*illed error. This is the female spruce grouse on the Some grouse’’. Camp Dubois (Wood River winter basis of habitat, eye stripe, size, shape, econo- camp), Illinois. my [presumably food habits], and habits. The reference to reddish brown pheasant clearly re- Moulton (1986) suggested these birds may have ferstothe ruffed grouse and eliminates the been ruffed grouse. We think it likely they were ‘‘yellowish brown’’ bird as a ruffed grouse, as prairie-chickens because of the usual use of pointed out by Moulton (1990). As well, authors ‘grouse’ for clear references to prairie-chickens of the journals sometimes noted its food habits: and ‘pheasant’ for clear references to ruffed budsand needles of conifers (presumably from grouse. As well, this was near the southern end crop contents), which separates it from all oth- of the range of ruffed grouse in this region er species but blue grouse, the 2nd largest spe- (Rusch and others 2000), where densities were cies encountered—see Lewis’s comment about likely low. foods of the small brown pheasant, 16 June 25 July 1804.—[Clark]: ‘‘Several Grous Seen in 1806 (below). the Prarie’’. Whitefish Camp, Iowa, about 17 km above the mouth of the Platte River [Clark made 2 Ruffed Grouse nearly identical entries on this day, presumably re- There are more references to ruffed grouse in ferring to the same observations]. the journals than observations because Lewis Moulton (1986) suggests these were probably and Clark often compared birds they observed ruffed grouse. Because Clark identified them as or killed to ‘pheasants of the Atlantic or United grouse, rather than pheasants, and in this States’, illustrating their acquaintance with the quote, refers to them as on the prairie, we con- species. For example: sider it most likely he was referring to prairie- 5February 1806.—[Lewis, at Fort Clatsop]: chickens. ‘‘Filds brought with him a phesant which differed 17 September 1804.—on this date, Lewis remarks little from those common to the Atlantic states; it’s abouta‘‘perfectly white’’ turkey, ‘‘booted as low brownisreather brighter and more of a reddish as the toes’’, and ‘‘of the Black Hills’’ [based on in- tint. it has eighteen feathers in the tale of about six formation received from a ‘‘french lad’’ who win- inches in length. this bird is also booted as low as tered with the Cheyenne Indians]. the toes. the two tufts of long black feathers on 1 October 1804.—a French trader, Jon Vallie,was each side of the neck most conspicuous in the male contacted in the vicinity of the Cheyenne River. He of those of the Atlantic states is also observable in had wintered some 300 leagues up this river, ‘‘un- every particular with this.’’ der the Black Mountains’’, where ‘‘white booted One characteristic noted here, ‘‘booted as low turkeys’’ were living (presumably the birds noted on 17 Sep). as the toes’’, does not exactly fit ruffed grouse for their metatarsi are not normally completely Coues (1893) referred to this bird as a prairie- feathered. This feathering usually refers to at- cock and suggested it was the sage-grouse. We tachment of the feathers on the metatarsi and find no reference to prairie cock in Moulton’s 8 NORTHWESTERN NATURALIST 84(1)

(1987) journal entries for 17 September or 1 Oc- birds and was talking about others in his party— tober, for Lewis or Clark. Holmgren (1984) he wasnot with them this day. thought the turkey was likely the sage-grouse Coues (1893) identified the pheasants in Clark’s [but noted that it could be the wild turkey (Me- passage as ruffed grouse. Entries by Clark and leagris gallapovo)and, except for size, resembles Gass clearly refer to the same birds and that by white-tailed ptarmigan (Lagopus leucurus)]. The Gass suggests they were likely sharptails, the description as a ‘white turkey’ is puzzling, but only prairie grouse in the area other than sage- with language differences, turkey could be grouse, 1 of which was separately, and clearly, used as a general descriptor like ‘pheasant’, identified by Clark. ‘grouse’, and ‘fowl’, in which case size of the bird may be irrelevant. Identity of this bird is 3 September 1805.—[Whitehouse]: ‘‘then passed unclear but because of its all white color we down a Steep hill in to the head of a cove and branch where we camped after a disagreeable days doubt it is the sage-grouse. The location too is march of only 11 miles with much fatigue and hun- unclear; the ‘Black Mountains’ may or may not ger as nothing has been killed this day only 2 or 3 be the Black Hills of Wyoming and South Da- fessents, and have no meat of any kind.’’ Party kota as that was a general descriptor for coni- heading north in Gibbonsville, Idaho-Lost Trail fer-covered mountains by early travelers in the Pass area. According to Coues (1893) they camped area, including Lewis and Clark. As well, Vallie about10kmbelow Lost Trail Pass, but Moulton described the mountains as very high, with notes that there is much confusion as to exactly parts retaining snow all summer, and these where they camped; habitat likely patchy shrub- were not likely the Black Hills. The next nearest steppe at lower elevations, merging into montane mountains that might hold snow in summer are forest and open ridges at higher elevations. the Laramie Mountains in southeast Wyoming These birds are not identified in Moulton, but or the Bighorn Mountains in northcentral Wy- Coues (1893) indicated they were ruffed oming, neither of which has been reported to grouse. Judging from the relatively high eleva- have white-tailed ptarmigan. The nearest tion and plant communities passed through, it known ptarmigan are in the Snowy Mountains seems more likely they were blue and/or in southeast Wyoming (C Braun, Grouse, Inc., spruce grouse. Tucson, AZ, pers comm). We consider this bird 6 September 1805.—[Clark]: ‘‘rained this evening unidentifiable. nothing to eate but berries, our flour out, and but 23 August 1805.—[Gass]: ‘‘They killed three little Corn, the hunters killed 2 pheasents only’’. prairie hens, or pheasants’’. Gass was with Clark’s Party appears to have started from near Warm reconnaisance party, headwaters of the North Fork Springs, Montana, where they apparently spent of the Salmon River in Idaho, perhaps near the the nights of the 4th and 5th. Judging from dis- mouthofDump or Moose Creek; habitat was likely tances in Coues (1893), they probably camped shrub-steppe at lower elevations, merging into somewhere near Medicine Tree along the Bitter- montane forest with increasing elevation. root River on the 6th, about 17 km north of Darby, Montana; party was now moving north along the Moulton (1988) suggests these may have been river bottom, which becomes perhaps 1 to 5 km sage-grouse. We think it more likely they were wide as they proceed. Lower S-facing slopes are sharp-tailed grouse for the party was virtually generally in grassland merging upward into open always clearly identifying sage-grouse as large ponderosa pine then Douglas-fir, with more Doug- and different from other species [see 24 August las-fir on N-facing slopes. (Clark), below]. Coues (1893) identified these birds as ruffed 24 August 1805.—[Gass]: ‘‘We caught some small grouse, but they could be blue grouse, or a fish to day, and our hunters killed 5 prairie fowls. mixed bag, especially if any of the hunters were These were all we had to subsist on.’’ ranging away from the valley bottom. Another 24 August 1805.— [Clark]: ‘‘the party had killed possibility is sharp-tailed grouse as the party Several phesents and Caught a fiew Small fish on wasnow entering potential habitat of this spe- which they had Subsisted in my absence. also a cies, but this is less likely since Clark identified heath hen, near the Size of a Small turkey.’’ Gass, them as pheasants, not grouse. with other members of Clark’s party, was moving up the Salmon River this day; habitat likely as on 13 September 1805.—[Clark]: ‘‘. . . I shot 4 Pheas- 23 August. Clark was not likely to have seen these ants of the Common Kind except the taile was SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 9

black.’’ Party started this day near Lolo Hot Pheasants, a large black species, with some white Springs, headed west up the Lolo Trail, crossed feathers irregularly scattered on the brest neck and the Montana-Idaho state line, and camped at Pack- belley a smaller kind of a dark uniform colour with er Meadownear Lolo Pass. Areas traversed were a red stripe above the eye, and a brown and yellow mostly forested, with ponderosa pine and Doug- species that a good deel resembles the phesant las-fir at lower elevations, spruce-fir and lodgepole common to the Atlantic States.’’ [Written on this pine at upper elevations; some open glades and date but appears to be merely descriptive and not meadows occur along creeks. related to birds seen or taken this day]. See Ord- way, this date, above for location and habitat. Moulton (1988) identified these birds as spruce grouse. The black tail rules out ruffed grouse Cutright (1969) identified these birds as blue and suggests spruce or blue grouse. Descrip- grouse, spruce grouse, and ruffed grouse, and tion of the birds as ‘‘of the common kind’’ Moulton (1988) apparently accepted all 3. We (brown or reddish brown) tends to rule out the agree with the first 2, but not the 3rd. Judging black (spruce) or blue-gray (blue) males of both from the area they had recently traversed, high species. We believe they would have been fe- along the Lolo Trail, and color of the 3rd bird, male spruce or blue grouse, or a mixed bag. we believe it refers to female spruce grouse, not Habitats traversed could be appropriate for ei- ruffed grouse—female spruce grouse are more ther. yellowish brown (see Lewis’s description of the small brown pheasant, above) than the more 20 September 1805.—[Ordway]: ‘‘came a Short distance and found a line which Capt Clark had reddish ruffed grouse. Also, Lewis and Clark left with the meat of a horse which they found in appear to have considered male and female the woods and killed for our use as they had killed spruce grouse as separate species. But, this one nothing but 1 or 2 phasants after they left us.’’ might be considered confusing because Lewis [Whitehouse made 2 similar entries this day and compared it closely to eastern ruffed grouse, in 1 reported, ‘‘three prarie hens or Phesants’’]. which superficially resemble female spruce Ordway and Whitehouse appear to have been with grouse. Lewis and the main party—they passed Hearty Meal Camp on this day and stayed at Full Stomach 16 June 1806.—[Lewis]: ‘‘I killed a small brown Camp—between Dollar and Sixbits creeks, Idaho pheasant today, it feeds on the tender leaves and County, Idaho. Specific habitat not known but both budsofthe fir and pitch pine.’’ [Clark had a similar parties were well into the mountains and traveling entry referring to this bird]. Party heading east, at relatively high elevations, moving west toward camped this night at Horsesteak Meadow on Hun- Weippe Prairie. gery Creek, Idaho County, Idaho, below Windy Saddle; specific habitat not known, but well into Clark, with 6 others, had moved ahead of the the mountains when written. main party on 18 September. Ordway and Whi- tehouse were with Lewis and talking about Moulton (1993) identifies this bird as perhaps a birds shot by Clark’s party. The entire party ruffed grouse. There are 2 clues here: reference wasnot united again until Lewis arrived at to a ‘small’ brown pheasant and to its feeding Weippe Prairie. These pheasants would have on leaves [needles] and buds of fir and pine. Be- been taken between 18 and 20 September and ing small and feeding on conifers, a common maybe2mentioned by Clark on 19 September. food of spruce grouse but not ruffed grouse, Considering the elevations traversed, they were suggests the former. Being brown, this was a fe- almost certainly blue and/or spruce grouse. male. Holmgren (1984) indicates ruffed grouse were 26 June 1806.—[Lewis]: ‘‘on our way up this taken on the 20th, presumably these birds, and mountain about the border of the snowey region identifies 1 name used by the party for this spe- we killed 2 of the small black pheasant and a fe- ciesas‘large black and white pheasant’, which male of the large dommanicker or speckled pheas- clearly is not the ruffed grouse (see Lewis’s de- ant, the former have 16 fathers in their tail and the scription written at Fort Clatsop, above). We latter 20 while the common pheasant have only 18. don’t understand Whitehouse’s ‘‘prarie hens’’ the indians informed us that neither of these spe- because Clark and party were nowhere near ceis drumed; they appear to be very silent birds for the ranges of any of the prairie grouse. Ineverheared either of them make a noise in any situation.’’ [Clark’s entry on this date is almost 20 September 1805.—[Lewis]: ‘‘Three species of word for word, a copy of that of Lewis]. Party 10 NORTHWESTERN NATURALIST 84(1)

heading east on the Lolo Trail and camped at 2100 m to about 1400 m and camped near Warm Greensward Camp, northeast of Hungery Creek, Springs Creek, a few km northwest of Sula, Mon- Idaho; specfic habitat not known, but well into the tana; likely passed through spruce-fir, Douglas-fir, mountains at upper elevations. andponderosa pine forest in their descent, with open ponderosa pine-grassland in the Warm Moulton (1993) called the 1st birds blue grouse Springs area. and the latter a spruce grouse, perhaps because Lewis and Clark at times interchanged the Killing 12 grouse in 1 thicket suggests blue names, ‘black pheasant’ and ‘speckled pheas- and/or spruce grouse, perhaps in a flock or ant’, for the 2 species, or because of Coues’ flocks, or perhaps a mixed bag, likely not (1893, p 870–872) misidentifications in his gen- ruffed grouse because of the relatively high el- eral discussion of species. Coues (p 1054) iden- evations most of the day. tified the small black pheasants correctly, as 7 September 1805.—[Ordway]: ‘‘. . . hunters killed spruce grouse, but the large dommanicker or 1goose 1 crain Several pheasants and a hawk.’’ speckled pheasant incorrectly, as ‘‘the same Partyheading north along the Bitterroot River, species’’. Burroughs (1961) also identified the camped near Corvallis, Montana; generally, the large dommanicker or speckled pheasant as valley bottom is now grassland or cultivated, spruce grouse, perhaps following Coues’ lead. merging upward into ponderosa pine-grass on Two clear clues to identity are ‘large’ and both sides of the river, then denser forest (likely ‘small’ and numbers of tail feathers—spruce Douglas-fir). grouse normally have 16, and blue grouse in These pheasants could be blue or ruffed grouse, this region have 20. The small black pheasants oramixed bag, especially if the hunters were are clearly male spruce grouse and the large ranging away from the valley bottom. The par- dommanicker or speckled pheasant is a female ty was now in potential sharptail habitat, but blue grouse. This is a case where Lewis this species is less likely to be part of the bag switched names that he earlier used for species since they were identified as pheasants, not he considered nearly identical except in size grouse. and numbers of tail feathers.

5August 1806.—[Lewis]: ‘‘we also saw on our DISTRIBUTION AND ABUNDANCE way ...few ducks or prarie hens.’’ Lewis’s party Approximate locations of different tetraoni- wasmoving east along the Missouri River and nes that we believe can be clearly, or very likely, camped near the mouth of Prairie Elk Creek, about 6.5 km west of Wolf Point, Montana; prairie habi- identified are depicted in Fig. 2. These provide tat, mostly now cultivated, with extensive stands an opportunity to compare distributions and of cottonwoods along the river. numbers prior to the arrival of Euro-American settlers with those of today. Moulton (1993) suggests the prairie hens may have been prairie-chickens. The party was well Greater Prairie-chicken outofthe range of this species at this location and deep in sharp-tailed grouse range [see 2 Distribution.—Prairie-chickens were encoun- September 1806 (Clark), above—they passed tered on the lower Missouri from its confluence the mouth of the James River that morning, with the Mississippi River to the vicinity of its heading downstream]. Surely, the birds re- confluence with the James River (Fig. 2A, all ferred to were sharptails, not prairie-chickens. within the range of T. c. pinnatus). The north- ernmost limits are made clearly by Clark in 2 GROUSE NOT IDENTIFIED BY MOULTON OR passages: 2 September 1806 (see above), and in OTHER AUTHORS a‘Postexpeditionary Miscellany’, written some timeafter his exploration of the Yellowstone 4 September 1805.—[Whitehouse]: ‘‘we [de]*scend River: ‘‘The Prarie Fowl common to the Illinois ed the mountain down a rough rockey way and are found as high up as the River Jacque [James along through a large thicket of bolsom fer timber in which we killed a dozen fessents then descend- River] above which the Sharpe tailed Grows ed down in to a large valley on a branch and halted commence 950 Ms.’’ These put their northern to dine’’. Party heading north and went over Lost limit in this area at or near the eastern bound- Trail Pass this day, entering the Bitterroot River ary of Nebraska with South Dakota, far short of watershed—traversed elevations between about the historic range that extended into southern SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 11

FIGURE 2. Distribution and dates of 59 clear, or very likely, identifications of greater prairie-chickens and sharp-tailed grouse (map A), greater sage-grouse and blue grouse (map B), spruce grouse and ruffed grouse (map C) recorded by the Lewis and Clark expedition, and of 24 observations in which identities were not clear and may have involved 1 or more species (map D). Dates are cited as day-month-year. 12 NORTHWESTERN NATURALIST 84(1)

Saskatchewan and southern Alberta (Aldrich 3rd possible area of occupation may have be- and Duvall 1955; Schroeder and Robb 1993). gun near Kamiah, Idaho, and extended down- Abundance.—Eight journal entries provide stream to the vicinity of The Dalles, Oregon (all clues to abundance. The first 2 were made prior within the range of T. p. columbianus)—they def- to setting up the 1st winter camp, Camp Du- initely occurred between the mouth of the bois, Illinois. On 16 November 1803, Lewis saw Snake River and the Dalles. This bird was thus ‘‘a heath hen or grows’’ and in the 2nd, 22 No- reported in 3 non-contiguous areas extending vember 1803, ‘‘some heth hens or grows’’, 1 of from South Dakota to the lower Columbia Riv- which was killed. Four entries were made by er, with major breaks occurring in the head- Clark while at Camp Dubois: 15 December watersofthe Missouri River and in the Bitter- 1803, ‘‘hunters [k]*illed some grouse’’; 5 Janu- root Mountains. Current range maps (Connelly ary 1804, ‘‘Two men whomeIsenttohunt and others 1998; Schroeder and others 2000a) grouse returned . . .’’; 9 January 1804, ‘‘I. . . indicate they are extirpated along the expedi- Killed Prary fowl . . .’’; and 20 January 1804, tion’s route from the Bitterroot Valley and ‘‘. . . many Grouse Caught to Day . . .’’. Three Salmon River westward. other entries were made as the party moved up Abundance.—Several of the 25 journal entries the Missouri: 25 July 1804 [Clark], ‘‘Several that clearly, or very likely, refer to sharp-tailed Grous Seen in the Prarie’’ and ‘‘Several Grous grouse provide evidence of abundance. First Seen to day.’’ [these may refer to the same ob- mention of this bird was on 12 September 1804, servations]; and 4 August 1804 [Lewis] ‘‘The in the vicinity of the Bijou Hills, South Dakota, prarie hen or grouse, was seen in the praries be- when Clark ‘‘Saw . . . a number of grouse’’. On tween the [mouth of the] Missouri and the river 21 September, near Medicine Creek, South Da- platte’’. These records are too scanty on which kota, he reported, ‘‘Grouse . . . is Common in to speculate much about abundance, but do not those Plains’’. On 6 and 7 October, in an area suggest large numbers. The only indication of now inundated by Oahe Reservoir, Walworth large numbers is in the entry of 20 January, a County, South Dakota, he and Lewis each not- time when birds might be concentrated in mid- ed, ‘‘great numbers’’ of sharptails. winter flocks. While in winter quarters at Fort Mandan, In summary, the prairie-chicken was likely Clark led a hunting expedition in early Febru- common, if not abundant, along the lower Mis- ary. On the 13th, on returning to Fort Mandan, souri River, to as far north as the Nebraska- he reported, ‘‘Saw great numbers of Grouse South Dakota border, a region from which they feeding on the young willows . . .’’ Other en- are now largely extirpated. They expanded into tries from North Dakota include: ‘‘I . . . saw . . . Northand South Dakota, where they are now flocksofGrouse, . . .’’ [Clark, 12 April 1805, sympatric with sharp-tailed grouse, following mouthofthe Little Missouri River area], and, settlement by Euro-Americans. Largest current ‘‘I also met with great numbers of Grouse or densities along the Missouri (small in relation prarie hens . . .’’ [Lewis, 15 April, Little Knife to historic standards) are between its conflu- River area]. ence with the James River and Pierre, South Da- On 30 April, in the vicinity of Brockton, Mon- kota (Fredrickson and others 1999). tana, Clark ‘‘. . . saw Great numbers of Ante- lopes, also Scattering Buffalow,...&Grows’’. Sharp-tailed Grouse On 21 May, in Phillips County, at a site now in- Distribution.—Sharp-tailed grouse were en- undated by Fort Peck Reservoir, Lewis ob- countered in grassland and shrub-steppe hab- served, ‘‘the growse or praire hen are now less itats from southern South Dakota to the west- abundant on the river . . . perhaps they betake ern part of present Phillips County, Montana themselves to the open plains . . . at this sea- (Fig. 2A, all within the range of T. p. jamesi). In son’’. Next day he added, ‘‘. . . passed the en- contrast to the current range (Connelly and trance of grows Creek [probably present Beau- others 1998), none were reported in the Mis- champ Creek according to Moulton (1987] . . . souri watershed above present Fort Peck Lake. this creek we named from seeing a number of They were next noted in the upper Salmon Riv- the pointed tail praire hen . . . these are the first er area, in the vicinity of North Fork, Idaho, and we have seen in such numbers for some days.’’ in the Bitterroot Valley of western Montana. A Summary remarks of Lewis and Clark for May SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 13 include, ‘‘25th . . . the grouse disappear.’’—in present city limits of Missoula. There is no in- the general vicinity of Two-calf Creek, Fergus dication of large numbers in this valley. County, Montana. Sharptails were not again re- After crossing the Bitterroot Mountains on corded until arrival in the upper Salmon River the westward journey, sharptails were 1st re- area, Idaho, west of the continental divide. ported near the confluence of the Snake and However, on the return journey, on 5 August Columbia rivers. On 17 October 1805, Clark 1806, in the vicinity of Prairie Elk Creek, Mon- ‘‘shot . . . Several Grouse . . .’’ On the 26th, he tana, Lewis wrote, ‘‘we . . . saw . . . immence recorded, ‘‘our hunters ...Killed five Deer . . . herds of buffaloe . . . but few prarie hens.’’ and a grouse.’’ (camped near The Dalles, Records for this 1st leg of the journey indi- Oregon), and on the 27th, ‘‘. . . hunters . . . cate sharptails were abundant along the Mis- brought in . . . one grouse . . .’’ (near The souri in North and South Dakota, perhaps less Dalles). This puts sharptails in this region into so in far eastern Montana, and perhaps non-ex- a small area, mouth of the Snake River to The istent or in low density in the Missouri water- Dalles. However, on the return journey, 23 May shed above the upper end of present Fort Peck 1806, at Camp Chopunnish, near Kamiah, Ida- Lake. When crossing the central plains, where ho,‘‘. . . all returned from hunting without hav- large mammals were abundant, prairie-chick- eing killed any thing except a fiew heath hens ensand sharptails were likely taken only inci- &black Pheasants . . .’’ [Clark; Lewis’s entry for dentally, perhaps to add variety to the diet. this day involves the same birds and he men- They may have even been passed by in an effort tions only ‘‘a few pheasants of the dark brown to conserve ammunition and numbers reported kind’’ (blue grouse)]. Moulton (1991) suggests may not reflect their abundance. Clark’s heath hens may have been blue grouse and this cannot be ruled out. By this time, how- After crossing Lemhi Pass into Idaho, Clark ever, we think Clark would have clearly known and 13 others moved down the Salmon River to the difference between blue and sharp-tailed evaluate its possible use as a route to the Pacific. grouse and it is possible sharptails were taken. On 23 August 1805 ‘‘They killed three prairie This would extend their range westward from hens, . . .’’ [Gass]; presumably in the vicinity of heretoatleast the Dalles. Moose or Dump creeks. Next day, ‘‘. . . our Journal entries thus indicate sharptails were hunters killed 5 prairie fowls’’ [Gass], up- common to abundant along the Missouri River stream from the day before. And on the 31st, in from near the mouth of the James River to west- the Tower Creek area of the Lemhi Valley, ‘‘. . . ernNorth Dakota and far eastern Montana. our hunters killed one Deer . . . & Prarie fowl’’ These areas are still occupied (Connelly and [Clark]. others 1998), likely at much lower densities. The entire party was now again united and They were not recorded in the upper Missouri moved up the North Fork of the Salmon River, beyond Two Calf Creek, although Connelly over Lost Trail Pass on 4 September, and en- and others (1998) show them as now occupying tered the Bitterroot Valley, back into Montana. most of eastern Montana. They were likely Moving north down the valley, they were soon common to abundant in the lower Snake River in potential sharptail habitat. On 8 September, and Columbia River areas for while at Fort Clark ‘‘killed a prairie fowl’’—camped near Clatsop, Lewis wrote, ‘‘they associate in large Stevensville. Next day, Clark ‘‘killed 4 deer... flocksinautumn & winter and are frequently &3prarie fowls . . .’’—they camped at Travel- found in flocks of from five to six even in sum- er’s Rest near the mouth of Lolo Creek, where mer ...’’. This passage indicates sharptails they stayed until the 11th. They ‘‘killed 4 deer were more abundant in thelower Columbia ...&3Grouse’’ on the 10th [Clark], presum- than indicated by daily entries, at least as far ably sharptails. These were the only entries for west as The Dalles. They appear to have been this species along the Bitterroot on the out- more sparse in mountainous areas, such as the bound journey, but on the return trip, on 3 July upper Salmon and Bitterroot valleys. 1806, ‘‘we killed a prarie hen with the short and pointed tail she had a number of young which Greater Sage-grouse could just fly’’ [Lewis], between Traveler’s Rest Distribution.—Lewis and Clark were made and the mouth of Rattlesnake Creek, in the aware of sage-grouse by Indians at Fort Man- 14 NORTHWESTERN NATURALIST 84(1) dan, who indicated they would be found in the of the confluence of the Snake and Columbia Rocky Mountains (based on location, C. u. uro- rivers. Here, on 17 October, Clark sent out phasianus), far west of their present range ‘‘Hunters to Shute the Prarie Cock . . . Several (Schroeder and others 1999). The party was un- of which I have killed, they are the Size of a doubtedly watching for them. Swainson and Small turkey . . .’’ On the same day, Gass re- Richardson (1831: p 359), some 25 years later, ported, ‘‘In the plains are . . . a number of reported, ‘‘They do not exist on the banks of the fowls, between the size of a pheasant and tur- river Missouri; nor have they been seen in any key, called heath hens or grous. We killed a placeeastofthe Rocky Mountains.’’, consistent great many of these fowls . . .’’ The next day, with reports by the Mandans. First encounter ‘‘Several Heath hens or large Pheasents lit near with this species was on 5 June 1805, in the low- us & the men killed Six of them’’ [Clark], and er Marias River area, Choteau County, Mon- Whitehouse commented, ‘‘Several Men of our tana (Fig. 2B). If this represented reality, his- party went out in the plains, & killed a number toric records from as far east as North and mor of the Priari or heath hens, which were South Dakota may represent post-expedition very large’’. These were the last reports of sage- expansion. At the very least, if this bird occu- grouse on the outbound journey. Camp that pied the Missouri watershed in far eastern night was near Wallulla, Washington. Montana, they likely were low in numbers. On the return journey, the party was travel- Sage-grouse were next reported in the Horse ing overland, well south of the Snake River on Prairie area near Grant, Montana, just east of 1May 1806, in the vicinity of Waitsburg, Wash- Lemhi Pass, and were seen again in the head- ington. ‘‘we Saw a Great number of Curloos . . . waters of the Salmon River near North Fork, prarie cocks . . . common to the praries’’ Idaho, west of the continental divide. They are [Clark]. still present in these areas. None were then re- Considering the remarks of Clark, Gass, and ported until the party’s arrival in the Lower Whitehouse on 17 and 18 October, sage-grouse Snake River and Columbia River areas (based must have beenatleast common, likely abun- on location, C. u. phaios), wheretheyare now dant, in the lower Snake and Columbia River extirpated. areas. In Lewis’s description of this bird at Fort Abundance.—After Lewis’s 1st encounter Clatsop, he noted, ‘‘The Cock of the Plains’’ with sage-grouse, none was noted in the jour- wasfound in ‘‘Great abundance’’ [from the nals until the party’s arrival at Horse Prairie, al- mouthofthe Snake River to that of the Deschu- most 2 months later—numbers were likely low tes River]. Clark’s 1 May 1806 entry (above) put in the intervening area. sage-grouse about 40 km southeast of the At Horse Prairie, on 12 August, Lewis ‘‘. . . mouthofthe Snake River. saw several of the heath cock with a long point- In summary, sage-grouse were 1st found ed tail . . . but could not kill one of them . . .’’ when nearing the Rocky Mountains, as pre- On 20 August, Clark ‘‘killed a cock of the plains dicted by Indians at Fort Mandan, and were 1st or mountain cock . . . with a long and pointed reported in abundance intheextreme head- tail . . .’’ near Baker, Idaho [their 1st sage- watersofthe Missouri (Horse Prairie). Their grouse in the hand]. And on the 24th, his party presence was established in the Lemhi Valley, killed a sage-grouse in the vicinity of Moose or but with little evidence of numbers. Next en- Dump Creeks in the upper Salmon River area. counters were in the lower Snake and Columbia Lewis, with the main party, left Horse Prairie River areas where they were apparently abun- on 25 Aug, heading for the Lemhi Valley. He dant, a region from which they are now extir- ‘‘saw . . . many of the cock of the plains.’’ Much pated. Historic records in much of eastern of the day would have been spent traversing Montana and into North and South Dakota may Horse Prairie, indicating sage-grouse were at reflect post-expedition expansions or low den- least common, perhaps abundant, in this area, sities. where densities are now apparently low. The 2 killed in the upper Salmon River area by Blue Grouse Clark’s party establish their presence there, but Distribution.—The party’s 1st clear contact are insufficient to speculate on abundance. with blue grouse (21 July 1805, based on loca- Next contact with this bird was in the vicinity tion, D. o. pallidus)was east of the continental SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 15 divide in western Montana (Fig. 2B), about 250 of the Salmon River, Idaho: ‘‘nothing killed this km west of their easternmost range in Montana day ...onlyafiew fessants.’’ [Ordway, 2 Sep- today. This is not surprising since the party tember 1805] and ‘‘nothing has been killed this wasmoving along the river and prime breed- dayonly 2 or 3 fessents . . .’’ [Whitehouse, 3 ing areas for this species were likely not the riv- Sep]. Based on locations and habitat these were er bottoms; they may have been missed east of most likely blue grouse but possibly ruffed that point. Also, many blue grouse may have grouse, or mixed bags. moved upward toward winter range by the On 4 September, the expedition left the Salm- time the expedition was passing through these on River watershed, passed over Lost Trail areas. Clear, or likely, reports of blue grouse Pass, and dropped back into Montana. White- continued in mountainous areas well into coast house noted a kill of a dozen ‘‘fessents’’, most forest in the lower Columbia River valley (5 No- likely blue and/or spruce grouse based on lo- vember 1805; based on location, D. o. fuligino- cation, habitat, and number of birds killed. sus). This range is not greatly different from They camped that night a few km northwest of that of today. Sula. As the party moved down the Bitterroot Abundance.—Numbers of blue, spruce, and Valley and out of likely spruce grouse habitat, ruffed grouse are difficult to assess because pheasants were killed on the 6th [‘‘2 only’’— they are locally sympatric in many areas, were Clark] and 7th [‘‘several’’—Ordway]—most often referred to only as pheasants, and were likely blue and/or ruffed grouse. They arrived likely sometimes taken as mixed bags. These at Traveler’s Rest near the east end of the Lolo species can often be separated, however, at Trail on the 9th and headed west over the Trail leastinLewis and Clark entries, because the on the 11th. authors usually referred to ruffed grouse as the On 12 September, a ‘‘pheasant’’ was killed common pheasant (or pheasant of the Atlantic [Ordway] somewhere east of Lolo Hot Springs or United States) and compared the other spe- that may have been a blue or ruffed grouse, but ciestothis bird, often described spruce grouse likely not a spruce grouse because of the rela- as small and black or brown, and blue grouse tively low elevation and habitat. On the 13th, as large and black or dark brown. The main Clark shot 4 ‘‘pheasents’’ that we identified problem comes from the simple use of pheasant above as either blue or spruce grouse. The next asadescriptor, most prevalent in writing of the day ‘‘2 or 3 pheasants’’ were killed [Gass] in the sergeants, but even these references can often vicinity of the Powell Ranger Station (well into be tentatively assigned to 1 or 2 species from Idaho). These may have been any of the forest location, habitat, or elevation. grouse, or a mixed bag—least likely is spruce The 1st clear reference to blue grouse (2 grouse because of the relatively low elevation. birds), and indicating an unfamiliar species, Clark and 6 men moved ahead of the main wasinthe general vicinity of Canyon Ferry party to hunt on 18 September as provisions Dam, Montana. Lewis apparently saw blue were very low, and arrived at Weippe Prairie on grouse the next day, 22 July 1805, in the Win- the 20th. On the 19th, Whitehouse recorded, ston area, referred to only as ‘‘a few phes- ‘‘One of our Men here killed a Pheasant, .. ,’’— ants’’—most likely blue grouse based on loca- most likely blue or spruce grouse based on el- tion and habitat. In the vicinity of present La- evation and likely habitat. Speaking of Clark’s hood, Montana, on 1 August, he ‘‘saw a flock of party, Whitehouse wrote, on the 20th, ‘‘they the black or dark brown phesants; the young had killed nothing after they left us only three phesant is almost grown we killed one of prarie hens or Phesants.’’—likely juvenile or fe- them.’’ These are the only clear, or likely, jour- male blue grouse for it would be surprising to nal entries concerning this species in the Mis- see the diminutive spruce grouse called prairie souri River watershed and suggest it was not hens; also they were still too far into the moun- common along this part of their route. This may tains for sharptails. On the 21st, 3 pheasants reflect the party’s not getting into higher ele- were killed by the main party [Ordway]. These vations where this bird may have been more were likely blue grouse or perhaps spruce abundant. grouse, but were least likely ruffed grouse; it The 1st likely records of blue grouse west of could have been a mixed bag. They camped the continental divide were in the headwaters that night at Pheasant Camp, near the bound- 16 NORTHWESTERN NATURALIST 84(1) ary between Idaho and Clearwater Counties, few ‘‘pheasants of the dark brown kind’’ were Idaho, 1 day shy of Weippe Prairie. broughtin[Lewis]. Arriving at Weippe Prairie, all members of The party left Camp Chopunnish and moved the party were in poorcondition, having faced to Weippe Prairie on 10 June where they re- possible starvation in passing over the Lolo mained until the 24th, waiting for snow to re- Trail. The few grouse killed, plus the odd deer, cedeathigher elevations. On the 11th, ‘‘Several 1 horse, and a colt, were meagre fare for a party pheasants’’ were killed [Ordway; likely blue of 33. Clearly, grouse were not abundant, al- and/or ruffed grouse; less likely, but possibly though some blue and spruce grouse may have sharp-tailed or spruce grouse]. On the 26th, moved into winter habitats and been relatively along the Lolo Trail, a female blue grouse was invulnerable to hunting. The party then killed [Lewis], the last report of this species. dropped into the Clearwater River bottom near Nowhere did the records indicate large num- Orofino, Idaho, to rest and build canoes and bers. All references that might have included were there until 6 October. A pheasant was re- more than 1 species included blue grouse as 1 ported killed on 25 September [Gass] and an- of the possibilities (Fig. 2D), indicative of the other on the 30th [Ordway], either of which wide range of habitats occupied by this bird. most likely would have been blue or ruffed grouse. These were the last forest grouse re- Spruce Grouse ported until arriving in the lower Columbia Distribution.—Unambiguous reports of River area [but the party was now moving spruce grouse were recorded only on the Lolo mainly by water]. On 5 November, and ap- Trail (Fig. 2C), between the vicinity of Lolo proaching the vicinity of Rainier, Oregon, Pass, Idaho, and just east of Weippe Prairie. Clark ‘‘. . . killed a grouse which was verry fat, However, they may have been included in the and larger than Common.’’ [only 2 species in- bag of 12 taken in the Lost Trail Pass area, like- habit this area, blue grouse (largest) and ruffed ly on the Montana side of the pass. Overall, grouse (Clark’s ‘Common’)]. No blue grouse spruce grouse were encountered where one were reported during the winter stay at Fort might find them today, with no obvious change Clatsop, perhaps because of their relative in- in range. All sightings were within the range of vulnerability to being seen in winter. F. c. franklinii. The next record of this species was of a male Abundance.—Six numerical reports of spruce taken 16 April 1806, near The Dalles. All re- grouse were recorded in the journals—only 1 maining references to possible blue grouse on the westward journey—although there were were made while returning to Camp Chopun- others of unidentified pheasants which likely nish. On 7 May, ‘‘a pheasant’’ [Ordway]; on the included this species. Lewis observed, on 18 8th, ‘‘Several . . . pheasants’’ [Ordway]; and on September 1805, ‘‘there is nothing upon earth the 9th, ‘‘a few pheasants’’ [Lewis] were killed. exeptourselves a few small pheasants, small During this time the party was in the general grey Squirrels,...’’ [they started that morning vicinity of the Clearwater River between Ahs- near Indian Grave Peak in Idaho County, Idaho, anke and Kamiah, Idaho. Most of these birds and camped at Dry Camp, near Bald Mt]. There were likely blue or ruffed grouse, based on hab- almost certainly were more encounters for itat and elevations. Lewis (and Clark) clearly described them while The party was at Camp Chopunnish 14 May at Fort Clatsop on 3 March 1806 (see Clues to to 10 June 1806. On the 14th, Lewis wrote, ‘‘La- Species Identities). buish . . . brought with him several large dark On the return journey, 16 June 1806, Lewis brown phesants . . . Shannon also returned killed a ‘‘small brown pheasant’’. This was east with a few pheasants . . .’’ [Clark’s entry for of Weippe Prairie during an early attempt to this day indicates Shannon’s birds were blue cross the Bitterroots. On 22 June, the hunters grouse], and on the 15th, Gass wrote, ‘‘hunters ‘‘killed nothing but one small pheasant.’’ came in and had killed nothing but some grous [Gass] somewhere east of Weippe Prairie, ...’’ [most likely blue grouse for we have seen where they had camped the night of the 21st. no use of this word to identify ruffed grouse by On the 26th, and moving east on the Lolo Trail, any of the authors]. On the 18th, the hunters Lewis observed, ‘‘on our way up this mountain killed ‘‘some grous’’ [Gass] and on 23 May, a about the border of the snowey region we killed SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 17

2ofthe small black pheasant...Twodayslat- ruffed grouse represents only 1 bird. On 5 Feb- er, ‘‘we killed a small black pheasant; this bird ruary 1806, at Fort Clatsop, Lewis reported, is generally found in the snowey region of the ‘‘Filds brought with him a phesant which dif- mountains and feeds on the leaves of the pine fered little from those common to the Atlantic and fir’’ [Lewis]. They were now on the east states . . .’’ and in the summary remarks of side of the Bitterroots and camped at 13 Mi Lewis and Clark for March 1806 we find, ‘‘7th Camp, Idaho County, Idaho, a few kilometers abird of a scarlet colour as large as a common from Powell Ranger Station. pheasant with a long tail has returned, one of The last clear report of spruce grouse was on them was seen today near the fort by Capt. 29 June: ‘‘I killed a small black pheasant near Clark’s black man . . .’’ On the return journey, the quamash grounds this evening which is the Clark ‘‘killed . . . a Common pheasant’’ on 2 1st I have seen below the snowy region . . .’’ July 1806, near Traveler’s Rest in the Bitterroot [Lewis]. They camped at Lolo Hot Springs, Valley. Some of the unidentified pheasants tak- which is outofwhatwewould consider normal en at lower elevations and along the Lolo Trail spruce grouse habitat and fits with Lewis’s in Montana and Idaho were likely this species. comment about finding the bird at such a low Nevertheless, ruffed grouse couldn’t have been elevation. very abundant along their route. This seems of Most spruce grouse were taken or seen at rel- interest because the expedition traveled in river atively high elevations, as indicated by Lewis’s valleys much of the time, and areas with ripar- comment above. Most reports of this species in- ian vegetation are normally prime habitat for volved killing only 1 or 2 individuals. Taken to- this species. gether, references to spruce grouse indicate In summary, none of the forest grouse was they were not often encountered and usually in noted as abundant and the largest number of lownumbers. However, although the authors clear, or likely, identifications was of blue did not note them as in flocks in their daily en- grouse, with spruce grouse next, then ruffed tries, Lewis, at Fort Clatsop, comparing them to grouse. Many references to pheasants in forest blue grouse, wrote, ‘‘it . . . ascociates in much grouse habitats may have included more than 1 larger flocks and is very gentle.’’ They clearly species. The largest numerical reference was of encountered more birds than documented in 12 ‘pheasants’ killed in the Lost Trail Pass area, the daily reports. with ‘several’ reported on a few occasions. Most references indicated 1, 2, 3, or ‘a few’ Ruffed Grouse killed or seen in a single day. In view of the Distribution.—Ruffed grouse were seldom shortage of food on the journeys through the clearly mentioned in the journals except when Bitterroot Mountains, we think the party comparing other species to those in the eastern would have taken any birds seen. Apparent low states. Evidence of only 3 clear contacts with numbers may reflect, in part, the generally dis- ruffed grouse were found (Fig. 2C), 2 of which persed nature of these species. Unless the au- were at Fort Clatsop. These 3 records span the thors failed to mention many birds taken on area from the mouth of the Columbia River to this leg of the journey, we suspect a similar the Bitterroot Valley. None was reported on the sized party on the same route today would westward journey or east of the continental di- likely see, and be able to kill, as many birds as vide, even though their historic range includes taken by the expedition. Numbers of forest upper portions of the Missouri and Yellow- grouse may not have been vastly different in stone rivers (Rusch and others 2000). If encoun- that area than today. tered, we would expect Lewis and Clark to comment on this species because of their fa- DISCUSSION miliarity with it. Whether this lack of obser- The Lewis and Clark expedition was the 2nd vations represents a historic change in distri- to make an overland journey across continental bution or low densities is unclear. Identifica- NorthAmerica, preceded only by that of Al- tions of ruffed grouse, though few, generally exander Mackenzie in 1792 and 1793. In con- span the known range west of the continental trast to Mackenzie, however, 1 mandate of the divide, as understood today. Lewis and Clark expedition was to document Abundance.—Each clear identification of faunal and floral resources. The almost daily 18 NORTHWESTERN NATURALIST 84(1) journals of the party have provided an oppor- conventional scientific format. Many behavior- tunity to view some of the fauna and flora of the al and other natural history observations were Missouri and Columbia River drainages prior also insightful. In our view, Lewis and Clark to movement of Euro-Americans into these ar- made few significant errors in the materials we eas. examined—perhaps the greatest was in iden- Anumber of authors have examined journals tifying the female spruce grouse as a separate of the expedition for their natural history con- species from the male. They clearly recognized tents and have not always agreed on identities the differences between species and described of the different grouse. Some problems have them well considering the times and their back- been caused by the journal authors’ use of dif- grounds. ferentand sometimes ambiguous common names, inadequate descriptions, and their con- ACKNOWLEDGMENTS tacts with as yet undescribed species. Among We acknowledge the assistance of library staff at 121 identifications of grouse in the journals by the University of British Columbia, Vancouver, sev- other authors and examined by us, 117 were for eral of whom pointed us in the right directions. K observations and descriptions we consider rel- Frieze, Executive Director of the North Dakota Lewis atively unambiguous. Among these, we were in and Clark Bicentennial Foundation in Washburn, agreement with 94 (80%). North Dakota, was helpful in interpreting some of Clearly, the journal authors did not always the Lewis and Clark maps. Staff at the Fort Clatsop record encounters with the different grouse, National Memorial near Astoria, Oregon, provided access to materials in their collections, and volunteer nor did they always provide information on librarian J Warner went out of her way to assist us in abundance. As well, the number of clearly iden- finding pertinent references. DF Nell of the Lewis tifiable contacts for some species was relatively and Clark Trail Heritage Foundation, Bozeman, small. Although this causes potential problems Montana, provided some literature. RM Zwickel as- in evaluating presettlement distributions and sisted with our examination of some of the habitats numbers, we believe 2 conclusions are war- along the Lewis and Clark Trail. CE Braun, PW El- ranted. First, the 3 prairie grouse all appear to liott, J Kristensen, LA Robb, and GF Searing offered have had significant changes in distribution helpful comments on earlier drafts of the manu- from the time of the expedition. In some cases script. GE Moulton provided the edition of the jour- these have been expansions while in others nals from which we primarily worked. they have been extirpations. Along with chang- es in distribution, numbers have declined in LITERATURE CITED most cases; all 3 prairie grouse are in trouble in ALDRICH JW,DUVALL AJ. 1955. Distribution of Amer- many areas. These are the species whose hab- ican gallinaceous game birds. Circular 34, Fish itats have been most impacted by agriculture and Wildlife Service, USDI. Washington DC: U. S. and other human land uses. Second, judging Government Printing Office. 23 p. from the journals, we don’t see similar impacts BOAG,DA,SCHROEDER, MA. 1992. Spruce grouse (Dendragapus canadensis). In: Poole A, Stettenheim on forest grouse, likely reflecting that the Bit- P, Gill F, editors. The birds of North America No. terroot Mountain habitats the expedition tra- 2. Philadelphia PA: The Academy of Natural Sci- versed are still relatively intact. Development ences; Washington DC: The American Ornithol- and increased use of resources in montane ar- ogists’ Union. 27 p. eas could change this situation. BOTKIN,DB. 1995. Our natural history, the lessons of In closing, we believe Lewis and Clark pro- Lewis and Clark. New York, NY: Berkeley Pub- vided excellent 1st descriptions of blue and lishing Group. 300 p. sage-grouse and of some subspecies of other BURROUGHS RD. 1961. The natural history of the western grouse, for example, Franklin’s spruce Lewis and Clark expedition. East Lansing, MI: grouse. Although Swainson and Richardson Michigan State University Press. 340 p. CONNELLY JW, GRATSON MW,REESE KP. 1998. Sharp- (1831: p xiii) thought the descriptions ‘‘too tailed grouse (Tympanuchus phasianellus). In: Poole vague for scientific purposes’’, Coues (1893) A, Gill F, editors. The birds of North America No. and we disagree. With careful reading, we be- 354. Philadelphia, PA: The Birds of North Amer- lieve all species described in some detail and ica, Inc. 20 p. many of the daily observations of grouse can be COUES E, editor. 1893. The history of the Lewis and clearly identified even though not presented in Clark expedition. Vols 1 to 3. Dover reprint of the SPRING 2003 ZWICKEL AND SCHROEDER:GROUSE OF LEWIS AND CLARK 19

1893 Francis P. Harper edition. New York, NY: and Clark expedition. Vol 11. Lincoln, NE: Uni- Dover Publications, Inc. 1364 p. versity of Nebraska Press. 459 p. CUTRIGHT PR. 1969. Lewis and Clark: pioneering nat- MOULTON GE, editor. 2001. The journals of the Lewis uralists. Urbana, IL: University of Illinois Press. and Clark expedition: comprehensive index. Vol 506 p. 13. Lincoln, NE: University of Nebraska Press. FREDRICKSON FC, CROUCH B, HEISMYER GL. 1999. 174 p. Status and management of the greater prairie- RUSCH DH, DESTEFANO S, REYNOLDS MC, LAUTEN L. chicken in South Dakota. In: Svedarsky WD, Hier 2000. Ruffed grouse (Bonasa umbellus). In: Poole RH, Silvy NJ, editors. The greater prairie-chick- A, Gill F, editors. The birds of North America No. en—a national look. Miscellaneous publication 515. Philadelphia, PA: The Birds of North Amer- ica, Inc. 28 p. 99–1999, Minnesota Agricultural Experiment Sta- SCHROEDER MA, HAYS DW, MURPHY MA, PIERCE DJ. tion. St Paul, MN: University of Minnesota. p 75– 2000a. Changes in the distribution and abun- 80. danceofColumbian sharp-tailed grouse in Wash- HOLMGREN VC. 1984. A glossary of bird names cited ington. Northwestern Naturalist 81:95–103. by Lewis and Clark. We Proceeded On 10:28–34. SCHROEDER MA, HAYS DW, LIVINGSTON MF, STREAM MOULTON GE, editor. 1986. The journals of the Lewis LE, JACOBSEN, JE, PIERCE DJ. 2000b. Changes in and Clark expedition. Vol 2. Lincoln, NE: Univer- the distribution and abundance of sage-grouse in sity of Nebraska Press. 612 p. Washington. Northwestern Naturalist 81:104– MOULTON GE, editor. 1987a. The journals of the Lew- 112. is and Clark expedition. Vol 3. Lincoln, NE: Uni- SCHROEDER MA, ROBB LA. 1993. Greater prairie- versity of Nebraska Press. 544 p. chicken (Tympanuchus cupido). In: Poole A, Stet- MOULTON GE, editor. 1987b. The journals of the Lew- tenheim P, Gill F, editors. The birds of North is and Clark expedition. Vol 4. Lincoln, NE: Uni- America No. 36. Philadelphia PA: The Academy versity of Nebraska Press. 464 p. of Natural Sciences; Washington DC: The Amer- MOULTON GE, editor. 1988. The journals of the Lewis ican Ornithologists’ Union. 24 p. and Clark expedition. Vol 5. Lincoln, NE: Univer- SCHROEDER MA, YOUNG JR, BRAUN CE. 1999. Sage- sity of Nebraska Press. 415 p. grouse (Centrocercus urophasianus). In: Poole A MOULTON GE, editor. 1990. The journals of the Lewis and Gill F, editors. The birds of North America and Clark expedition. Vol 6. Lincoln, NE: Univer- No. 425. Philadelphia, PA: The Birds of North sity of Nebraska Press. 531 p. America, Inc. 28 p. SHORT LL JR. 1967. A review of the genera of grouse MOULTON GE, editor. 1991. The journals of the Lewis (Aves, Tetraoninae). American Museum Novita- and Clark expedition. Vol 7. Lincoln, NE: Univer- tes 1189. 39 p. sity of Nebraska Press. 383 p. SWAINSON W, R ICHARDSON J. 1831. Fauna boreali- MOULTON GE, editor. 1993. The journals of the Lewis Americana; or the zoology of the northern parts and Clark expedition. Vol 8. Lincoln, NE: Univer- of British America. Part 2, The birds. London, UK: sity of Nebraska Press. 456 p. John Murray. 523 p. MOULTON GE, editor. 1995. The journals of the Lewis ZWICKEL FC, DEGNER MA, MCKINNON DT, BOAG and Clark expedition. Vol 9. Lincoln, NE: Univer- DA. 1991. Sexual and subspecific variation in sity of Nebraska Press. 419 p. numbers of rectrices of blue grouse. Canadian MOULTON GE, editor. 1996. The journals of the Lewis Journal of Zoology 69:134–140. and Clark expedition. Vol 10. Lincoln, NE: Uni- versity of Nebraska Press. 300 p. Submitted 30 March 2002, accepted 15 November MOULTON GE, editor. 1997. The journals of the Lewis 2002. Corresponding Editor: SR Johnson.