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Fp ati neta Miptincsl! Oe ps aorta Fyeetitgen rng whe ee SPE NI Yen i NT eS Pee PE aw Foe reagan pe lets APR ont niet noena sot - y > - ~ + eines waa a at eee SO en ee Sinhtigoegetaniiges hen ck eae . Pe peer . eed Fiera oe x Ses" ating, Fa tc hg hong Sal retag esc annis? eve eT Sion ™ San Vere tLPe, palatine ecati Me ee ae eee ke ene Se 2 mee pope amie es me A big a ei aed st ie - os Ty Mie TTR PA age fx, ot 9 atte mel age aah, Caan 2S ik “a on ae fe ~ " ? sera acteiaed > p AS : a Tek wn A Cone, a, Meera AN, Meee te = ie TR ig ny ae ea eat” BIE Rage atts Oy ae ae a en ee SO nad CONTRIBUTIONS OF THE ROYAL ONTARIO MUSEUM OF ZOOLOGY No. 6: A STUDY OF THE SHARP-TAILED GROUSE, BY L. L. SNYDER. (Reprinted from University of Toronto Studies, Biological Series, No. 40). 1935 NORTHERN SHARP-TAILED GROUSE Pedioecetes phasianellus phastanellus (Linn.) From a painting by T. M. Shortt. University of Toronto Studies Biological Series, Wo. +0 A STUDY OF THE SHARP-TAILED GROUSE By L: L. SNYDER ROYAL ONTARIO MUSEUM OF ZOOLOGY TORONTO THE UNIVERSITY OF TORONTO PRESS 1935 Digitized by the Internet Archive in 2011 with funding from University of Toronto http://www.archive.org/details/studyofsharptail0Osnyd CONTENTS Introduction and acknowledgements. .......... Ue ai 5 PART I THE 1932 EMIGRATION OF Pedtoecetes phasianellus phasianellus AND RELATED TOPICS The races of sharp-tailed grouse and their distribution.... 7 Periodicity in the sharp-tailed grouse................. r( On the seasonal distribution of sharp-tailed grouse in POEM -OSSCEEIA INGE IO TECE ))\iiale wilde ou Gaceave kw 12 SIE ERT ARGUE i ool Qi Nias Gi lk, Sh UNE 13 IN TARE CREEL Ul 6 y's bias) ay dlegien dongs wa. Goede yaks 13 PN MLA le cares ia shall lad eta ai teres ches ee alan tod 15 The period of the flight, its progress, and the TUL NA ATUL UV UNA Cn UNTER EN! big 15 On the fall and winter habits of emigrating sharp- Bate) SOUS pie ia ee NAN as) ar akan Vespy lala MOS ahi 17 BA ES AN eNO ON GM He sua ails ter aaa UCN AS Li Lote MA RECOGNI MEI OK OA RH DENUMSE RPRAS He 18 CCS OE a NRE NR Bee OND SLL TaN 18 Le Tee MME ENTE SEES A an ce RAs ea eee 18 IOS CAMARO A aC Mg RMON cL PR RRA RE 19 BAO ie cia) Soe tale) alle al cu nah eh al anal Aa el 19 PU een IL Sanayi Cait INP Wie, Meh 20 Specimens from the 1932 flight and the distribution of Pre TE NY UPR EG Ne NEON a A A 22 REN sta OM lr uc sey Co dal Cag UE SR TON 22 ESCA OUT ON MOOT i efi iklel an). via aualatanetnaa i tars 23 The status of the sharp-tailed grouse during the year subsequent to the 1932 emigration.................. 24 Reports from south of Canadian National Trans- POM erat aay i te ha ae 24 Reports from north of Canadian National Trans- Peaveemhi a i) ce) MM ie | ar a A RR ee A 26 4 A STUDY OF THE SHARP-TAILED GROUSE PAGE Notes on the mating habits of the northern sharp-tailed OMNI nia Saige a'ol view be 4b cee ea a te eles aa A 26 Factors which worked to diminish the numbers of north- ern sharp-tailed grouse in the east following the emiavation OF 1UB2. Aster eo ewe Gade ere Catala 27 A discussion of the final phases of the emigration....... 29 Possible causes and directional factors of the emigration 31 PART-II A PRELIMINARY TAXONOMIC STUDY OF THE GENUS Pedioecetes The taxonomy and nomenclature of the sharp-tailed F498 AOR LO aN OP ANP Nici sy abs © 36 On the distribution of characters of the sharp-tailed ERE: 4) cca s sh Rik nie eis 8 <5 ie ee a 40 Remarks on the forms of the sharp-tailed grouse......... 48 On Pedtioecetes phasianellus phasianellus (Linnaeus) 43 A description of Pedioecetes phastanellus phasi- UELIOES CU MIIOIIS))2, . . sas. odes ge kee ene 44 RR A aoa eS a eee art eek ee re 44 Fall and winter NDR NN Peat Naner iy 19.6) 45 Females tine e ais) s0.e 6. ae Oe Dea eee 47 Pav nnd SVATEG OE: cos /c a poate oak, Re ea ee 47 Aucbunlte ‘tn summer OS oy ees wea ee 47 PUM A cite Ganraiadt aye et ee ee ae 47 INSERM yen idce Gala ee eye Oe cee en 48 On Pedioecetes phasianellus kennicottu (Suckley).... 48 On Pedtoecetes phasianellus columbianus (Ord)...... 53 On Pedtoecetes phasianellus campestris Ridgway..... 55 On Pedioecetes phasianellus jamest Lincoln.......... 56 On what appears to be an unnamed race of Pedioe- GObES PUUSTOHENUS: oo 6 caudal Be ae ae ee ee 57 Summary. of taxonomic Giscussion... (ss: 5¢< hs ens 59 A theoretical discussion on the origin and dispersal of the races, of Pedtoeccies Diastanellus. co. 4 5 alc ee aa se Be 60 A STUDY OF THE SHARP-TAILED GROUSE L. L. SNYDER INTRODUCTION AND ACKNOWLEDGEMENTS When it became evident in the fall of 1932 that a south- ward emigration of sharp-tailed grouse was taking place in parts of northern Ontario and Quebec, the Royal Ontario Museum of Zoology undertook the gathering of facts relative to the population involved and on the status of sharp-tailed grouse in general. A questionnaire was circulated among correspondents and observers distributed throughout Canada who kindly co-operated by contributing a rather extensive record of observations. The compilation of this record gave a very satisfactory account of the numbers, distribution, habits, and behaviour of these northern grouse during their southward movement in the fall and winter of 1932-3, and a general idea as to the status of the species as a whole in Canada. To allow for the incorporation of some remarks on the aftermath of the emigration of northern sharp-tailed grouse in the east, a report on the subject was delayed and a later questionnaire distributed the following year. This second questionnaire was directed toward the status of the species, more particularly the emigrants, during the spring, summer, and fall of 1933. The circulation of questionnaires constitutes a survey which cannot be accomplished in any other way. The Museum wishes to thank the correspondents who have co- operated, collectively, since the large number of persons con- cerned prevents individual acknowledgements here. As a further aid, the following persons forwarded speci- mens to the Museum: Stuart Criddle, Treesbank, Man.; E. S. Davison, Fort Frances, Ont.; L. S. Dear, Port Arthur, Ont.; R. Edleston, Timmins, Ont.; W. R. Gardner, Cochrane, Ont.; E. R. S. Hall, Kenora, Ont.; L. J. Houston, Nakina, 5 6 A STUDY OF THE SHARP-TAILED GROUSE Ont.; W. Kishbaugh, Iroquois Falls, Ont.; G. M. Parks, North Bay, Ont.; H. H. Parsons, Armstrong, Ont.; E. Robert- son, Toronto; J. A. Simard, Ville Marie, P.Q.; R. A. Smith, Toronto; E. Sullivan, Emo, Ont.; T. A. C. Tyrrell, Kirkland lake, Ont.; F. H. Warner, Fort Frances, Ont.; S. Waller, Gypsumville, Man.; R. V. Whelan, Smoky Falls, Ont. The assistance rendered in this way is gratefully acknowledged. The writer is indebted to a number of institutions and individuals for the use of comparative material which was generously forwarded to the Royal Ontario Museum of Zoology upon request. These are as follows: Carnegie Museum, Pittsburg, through Mr. W. E. C. Todd; Fleming Collection, Toronto, by Mr. J. H. Fleming; National Museum of Canada, Ottawa, through Mr. P. A. Taverner; Museum of Natural History, State University of Iowa, Iowa City, through Professor H. R. Dill; United States National Museum, Washington, through Dr. A. Wetmore and Dr. H. Freidmann. Important data on various matters have been contributed to this study. The National Parks of Canada Branch, Ottawa, through Mr. Hoyes Lloyd, submitted its files and clippings relative to the emigration of sharp-tailed grouse and other matters. The Bureau of Biological Survey, Food Habits Research, through Mr. W. L. McAtee, examined the stomach specimens forwarded to it and completed a return of its quantitative and qualitative analyses. For this co- operation grateful acknowledgement is here made. Also the writer wishes to thank his associates for aid in various ways. The Ontario bibliography of birds compiled by Mr. J. L. Baillie has been most useful and convenient; Mr. T. M. Shortt has made the maps and drawings used in the paper. Aid rendered by several persons is acknowledged in the Text. In conclusion the writer wishes to thank Mr. J. H. Fleming, Honorary Curator of Ornithology, and Professor J. R. Dymond, Director of the Royal Ontario Museum of Zoology, for criticisms of the manuscript. PART) IL THE 1932 EMIGRATION OF PEDIOECETES PHASIAN- ELLUS PHASIANELLUS AND RELATED TOPICS THE RACES OF SHARP-TAILED GROUSE AND THEIR DISTRIBUTION For purposes of the following discussion on the emigration of northern sharp- tailed grouse and the status of the species in general, the most recent conception of the grouping of racial variants is here transcribed from the American Ornithol- ogists’ Union Check-list of North American Birds (19381). Genus Pedioecetes Pedioecetes phasianellus phasianellus (Linnaeus). NORTHERN SHARP-TAILED GROUSE Range.—Central Alaska, northern Manitoba, and north- ern Quebec (Ungava) to Lake Superior, and casually to Parry Sound district, Ont., and the Saguenay river, P.Q. Pedioecetes phasianellus columbianus (Ord). COLUMBIAN SHARP-TAILED GROUSE Range.—Interior lowland of British Columbia south to north-eastern California (formerly), Utah, Colorado, and northern New Mexico. Pedioecetes phasianellus campestris Ridgway. PRAIRIE SHARP-TAILED GROUSE Range.—Southern Alberta and southern Manitoba to Wyoming, eastern Colorado, central Nebraska, eastern South Dakota, Minnesota, and western Wisconsin, formerly to Kansas, western Michigan, and north-eastern IIlinois. PERIODICITY IN THE SHARP-TAILED GROUSE It is almost common knowledge that certain animal populations undergo periods of density, alternating with periods of scarcity. Naturalists have taken considerable interest in recent years in this phenomenon and the number of species whose populations have been discovered to be numerically unstable is ever growing. Grouse are among the 7 8 A STUDY OF THE SHARP-TAILED GROUSE birds whose populations fluctuate periodically. Leopold (1931) has recently examined conditions, past and present, relative to the prairie chicken (greater, Tympanuchus cupido americanus) and the sharp-tailed grouse (prairie, Pedioecetes phasianellus campestris) in the north-central United States, and in the summary of his findings he states that ‘‘chickens [meaning both species] like other grouse, are cyclic’. Since we are here particularly concerned with the sharp- tailed grouse, a more specific reference can be cited, namely that of Criddle (1930) which deals with the prairie form of this species. Criddle has shown that this grouse reaches a peak of numbers lasting two to three years, after which the population declines rapidly and does not reach another peak until approximately eight years have elapsed. His records, covering a period of thirty-four years, show rather strikingly that there is not only a periodic numerical change, but that the periods are rhythmic, approximately ten years being involved in the complete cycle. Criddle’s data for P. p. campestris cover a period of thirty-four years, from 1895 to 1929, and show three peak periods, namely 1902-4, 1912-3, and 1923-5. Although corresponding periods of numbers in the population of the northern race, P. p. phasianellus, might be expected, our ornithological literature and the Museum’s questionnaire do not offer sufficient information to demon- strate the point fully. The principal reason for this paucity of data is that the normal home range of typical phasianellus is in the inaccessible and little-known north. The Museum’s request for information on the sharp-tailed grouse in general was given wide distribution mostly in Canada, and it is of interest here to note the status of the species during the period of 1931-3. In British Columbia the population of this grouse (P. p. columbianus) was apparently not at aconspicuous maximum. Previous periods of numbers there were variously estimated as 1911 and 1923. In Alaska (A.O.U. P. p. phastanellus) there was a notable increase in 1931 and an abundance of the birds in 1932. In Alberta (A.0.U. Psp, campestris and P. p. phasianellus) a steady increase was noted since 1926, which was apparently the last minimum period, with great numbers being present in the winter of 1932-3. Saskatchewan (A.O.U. P. p. campesiris EMIGRATION OF THE SHARP-TAILED GROUSE 9 and P. p. phasianellus) reports show a gradual increase from a period of scarcity prior to 1929 to an abundance in 19382. Manitoba (A.O.U. P. p. campestris and P. p. phasianellus) reports show that the last period of scarcity was between 1926 and 1928 with a gradual increase since that time to 1932-3. In western Ontario (A.O.U. P. p. campestris) the reports show that the last period of abundance was between 1918 and 1923 with a period of scarcity following in 1926 and 1927, and further show a steady increase over the subsequent period with the grouse being plentiful in 1932-3. Reports from available sources in northern Ontario (north of Canadian National Transcontinental Railway) were largely confined to a few settlements and areas through which there are travelled routes. These are in agreement that the sharp-tailed grouse (P. p. phasianellus) has been approaching plentiful numbers since 1930, with the greatest numbers in years being present in the fall of 1932. There were occasional reports here and there through the Prairie Provinces and into Ontario, in the territory of A.O.U. campestris, which indicate that the numbers of the prairie sharp-tailed grouse during the period from 1931 to 1933 were exceptional. This suggests the possibility of a major peak, one in excess of Criddle’s (loc. cit.) ten-year peak of numbers,* although Mr. Stuart Criddle, brother of the late author to whom reference has been made, cannot substantiate this belief. Although our available information, both from the literature and from the Museum’s questionnaire, does not satisfactorily cover the question of numbers over a long period of years, there is considerable evidence that the increase in numbers of sharp-tailed grouse in the far north, during the period from 1930 to 1932 in Alaska and northern Ontario, and since 1926 in northern Alberta, coincides in period with the increasing population of sharp-tailed grouse on the prairie. There is ample evidence that the north-eastern population of sharp-tailed grouse does, at times, experience periods of surplus of numbers and although it cannot now be demon- *B. W. ieee in his weekly nature article for the Winnipeg Tribune, states that an old hunter who has kept close track of game conditions in Manitoba informs him that sharp-tailed grouse were more numerous during 1933 than they had been for forty years. 10 A STUDY OF THE SHARP-TAILED GROUSE strated that there is a lesser cycle of numbers approximating ten years, asin the prairie region, it is here suggested that there may be a major cycle approximating something more than thirty years. (See below.) This may or may not be a feature in connection with long-period changes of population of prairie sharp-tailed grouse. Substantiation of both these points will, of course, rest with observations made in the future. It seems safe to assume that the pronounced periodic southern occurrences of northern sharp-tailed grouse (1896 and 1932) indicate an abundance of individuals but in examin- ing the records for what might be termed lesser periodic peaks of numbers of this race, the question is complicated. First, reports of occurrences of sharp-tailed grouse south of the Canadian National Transcontinental Railway, in Ontario, a line which may be regarded as approximating the normal southern limits of P. p. phasianellus, may not refer to typical examples of that race. It seems fairly certain that A.O.U. campestris (and possibly intermediate birds, 2.e., campestris X phasianellus) may occur eastward, more particularly during the period just prior to a peak of numbers, into territory around lake Nipigon and Long Lac, Thunder bay district, and even to the vicinity of Franz and Oba of Algoma district.* Second, a confusion is also caused by birds being left in central and northern situations, south of their normal limits, following a major peak emigration of the northern race. This seems certainly to have been the case subsequent to the 1896 emigration. These birds apparently occurred locally in diminishing numbers for several years. Unfortunately, few substantiating specimens have been collected to determine some of these questions. Three specimens in the Royal Ontario Museum of Zoology collection taken at lake Temis- kaming, Ont., in November, 1902, possibly represent remnant birds (a point to be discussed later) from the 1896 flight. According to reports, such birds persisted in that region until 1910. *It is also possible that birds representing A.O.U. campestris may hereafter be found in central Ontario, having come in from the upper peninsula of Michigan where they occur as native birds and where eastward transplants are reported. EMIGRATION OF THE SHARP-TAILED GROUSE 11 In a published note (anonymous, 1867) it is stated that a specimen of ‘‘pin-tailed grouse’’ was shot at Sault Ste. Marie, Ont., and exhibited at a meeting of the Canadian Institute held in Toronto on January 13, 1866. The note further remarks that this grouse was to be found in more northerly parts of the province. Although there can be nothing certain from such a record, it rather strongly suggests that the speci- men referred to represented the northern race. In attempting to ascertain the distribution of the prairie race during the year 1866, it is not altogether certain as to the exact eastern front in this latitude. Coues (1874) writing eight years after the Sault specimen was taken, marks the eastern limit of “‘columbianus’’ (=campestris) as western Iowa and Minnesota, but, probably because of incomplete data, could but loosely define the eastern limits of the species as a whole, thus leaving us to suppose that it probably occurred eastward in the great lake region, but in such a case, birds of the area were considered to be referable to the north- ern race. Cory (1909) states that campestris was resident in northern Wisconsin in 1877. Considerably later, and therefore less important in a con- sideration of the distribution of sharp-tailed grouse in 1866, Mcllwraith (1886) in his first edition of the Bzrds of Ontario, stated that the species has been reported from Sault Ste. Marie and apparently considered the form concerned to be P. p. phastanellus. Macoun (1909), however, interprets Mc- Ilwraith’s statement as birds “being found on the market at Sault Ste. Marie’. Also, more recently, Wood (1905) and Leopold (loc. cit.) record the species from Isle Royale and the upper peninsula of Michigan. Since great changes in the forest of this region had taken place by the time of these later reports, no certainty can be placed on them as indicating an ancient range. Another consideration is the possibility of an artificial introduction. The only note on this subject which has come to the writer’s attention is of too late a date to be of concern here.* ) *Toronto Mail and Empire for October 6, 1894, states that seven pairs from Manitoba were liberated at Galt, Ont., previous to that date and that two nestings had been observed. The introduction failed, however. 12 A STUDY OF THE SHARP-TAILED GROUSE Assuming that the Sault Ste. Marie specimen represented a true Pedioecetes phasianellus phasianellus on the basis that P. p. campestris and intermediates did not occupy the region in 1866, its occurrence there would, therefore, be readily explained by a fall and winter emigration of birds from the typical northern locality of Hudson bay, an event which has twice been experienced since that date. Such a movement must have been a major one to have reached the latitude of Sault Ste. Marie, and according to the meagre reports on the bird it may have occurred in the fall and winter of 1865-6, or if the bird was a remnant, a point to be discussed later, the emigration may have occurred a year or more prior to that time. Thirty-one (or perhaps more) years after this hypothetical emigration, a conspicuous flight occurred. Fleming (1906) has recorded this flight of typical phasianellus which took place in the fall of 1896. Although he states it “‘was not large’’ it was of sufficient proportions to culminate in records of the birds from as far south as Beaumaris, Huntsville, Port Cockburn, and Bracebridge in Muskoka district and Emsdale and elsewhere in Parry Sound district. After another thirty-six years a major movement has again occurred, namely the flight of 1932-3. The writer is con- vinced from the information at hand that these flights mark at least the greater peak periods of numbers of the northern sharp-tailed grouse populations which occupy the Hudson bay region. ON THE SEASONAL DISTRIBUTION OF SHARP-TAILED GROUSE IN NORTH-EASTERN NORTH AMERICA Consequent to the gradual enlargement of our knowledge of the avifauna of eastern Canada, it is apparent that the sharp-tailed grouse does not regularly inhabit the region which can be roughly defined as from the lake Nipigon region eastward through Ontario into Quebec, and south of the Canadian National Transcontinental Railway. From south of lake Nipigon north-westward, suitable territory is occupied by birds referable to the race campestris within the A.O.U. application of the name. Because of the possibility of inter- EMIGRATION OF THE SHARP-TAILED GROUSE iS mediate birds, 2.e., campestris & phasianellus, occurring in the north-western part of Ontario, a consideration of the normal status of typical phasianellus is confined to the more northern and eastern parts of its range. Summer Distribution The record applying to the earliest time chronologically (prior to 1768, Preble, 1902) is that of Forster (Sclater, 1882), which states that this grouse breeds in the Severn river region and at Fort Albany. Richardson’s record (1831) referring to a later period and attributing the observation to a Mr. Hutchins who spent a considerable time at Fort Severn, and in the Severn river region generally, might be doubted on the grounds that the description of the eggs (“‘white with coloured spots’) appears to indicate confusion of species, but the general statement of breeding is substantiated by Forster’s earlier record. Fleming (loc. cit.) has recorded a set of eggs taken on May 20, 1889, at Fort George (Quebec) by A. P. Lowe, and Macoun (Joc. cit.) remarks that young birds ‘‘able to fly’’ were seen between the mouth of the Hannah river and East Point on James bay on July 8, 1904, by Wm. Spreadborough. The information gathered by the Museum’s questionnaire indicates that the status of the species as a breeding bird varies from time to time. The occupancy of nesting territory in the upper (more southern) portions of the Hudson bay and James bay watersheds, is probably somewhat irregular, but with a noticeable increase of frequency during the year or two prior to a peak of numbers and for at least one year subsequent to an emigration. This seems certainly to have been the case prior to, and following, the emigration of 1932-3. In other words, the population of northern sharp-tailed grouse somewhat expands its nesting territory southward when numerical pressure is present. Winter Distribution Many authors (such as Roberts, 1932) remark on the seasonal movements of the prairie sharp-tailed grouse (P. p. campestris) and for the most part these movements are regarded as habitat changes. Flocks leave the nesting 14 A STUDY OF THE SHARP-TAILED GROUSE territory for areas more completely and extensively covered with trees and shrubs. This is apparently an alimental necessity or convenience, since food can be obtained more abundantly and readily above the ground than on the ground after snow falls. There is, apparently, no obvious north- south movement which can be interpreted as a marked climatic change of residence for this form of the sharp-tailed grouse. In the case of the eastern population of the northern sharp-tailed grouse (P. p. phasianellus), there is a similar seasonal movement, and Preble (1908) states that the sharp- tailed grouse of the north-west “‘to some extent is migratory” and observers more recently concur with this. Probably because of food advantages, this movement has a southward direction in the fall. Fleming (loc. cit.) states that this bird occurs regularly zm October at lake Abitibi. The Museum’s work on lake Abitibi (Snyder, 1928b) did not discover the species there in summer, and residents informed the writer that these grouse occur in the fall and winter, not in summer. At lake Nipigon a similar condition prevails, since the species was not found during the Museum’s two summers of survey work in the region (Snyder, 1928a). The species is, however, to be found there fairly regularly in winter according to residents, and a specimen record was secured. The form could not be determined, however, because of the incomplete nature of the specimen. Typical phasizanellus probably appears there occasionally in winter. Thisis definitely known to be the case as a major period of numbers is approached. In 1931 birds which probably represented the northern form appeared ‘‘plentifully’’ at Hudson and in November of the following year, at the time of the marked emigration, a typical specimen of the northern race was taken at Armstrong. They were also present in the Kapuskasing and Cochrane areas in the fall of 1931. In general, available information indicates that the Canadian National Transcontinental Rail- way forms a convenient line to indicate the approximate southern front of the northern sharp-tailed grouse in winter when the population of these birds is not at a minimum, and at a period of maximum numbers they can be expected well south of this line. tn EMIGRATION OF THE SHARP-TAILED GROUSE 15 THE 1932 EMIGRATION The Period of the Flight, Its Progress, and the Numbers Involved As has been discussed, sharp-tailed grouse, over most of its range in Canada, had become progressively more numerous during the two or three years prior to the fall of 1932. A peak of numbers was apparently approached or reached in that year, and as a culmination of these extraordinary numbers there was an exodus of birds from the Hudson bay and James bay watersheds of Ontario and Quebec. ‘The place of origin of the birds in flight is determined not only on the direction as observed at Moosonee but by a comparison of specimens secured, as will be shown later. The movement was largely in a southward direction and involved the eastern population of the A.O.U. race Pedioecetes phastanellus phasianellus (see map I). It is of interest to mention at the outset that a similar movement (massed emigration) has not been reported in the case of northern sharp-tailed grouse inhabiting the northern parts of the Prairie Provinces, Mackenzie and Alaska regions, either in 1932 or any other year. In October, 1932, the grouse appeared in unusual numbers at Moosonee, at the south end of James bay. Mr. George S. Cotter, writing from that situation, stated that ‘‘flocks, both large and small, appeared to be coming from the east [Ungava] and north [Patricia] and working down south and west’’. Another observation from Moosonee, made near the end of October by Mr. R. L. Lamb, concerns these grouse as they ‘passed over Moosonee in vast flocks reminding him of the stories he had heard of the passenger pigeon in years gone by’. He further states that this migration continued approximately at the same rate for about three weeks and was still going on, on November 26, although on a very much diminished scale. Mr. R. V. Whelan of Smoky Falls, Ont., approximately one hundred miles south-west of Moosonee, states that the flight was first noted there on October 17. ‘“‘Up to forty were counted in a flock, and flocks were passing over several sections’ of his territory. Mr. R. R. Cockburn, referring to the district from Cochrane west to Nakina (Thunder bay district) states that ‘‘from October 12th to November 20th [these birds were there] in thousands’”’. Mr. W. R. Gardner, 16 A STUDY OF THE SHARP-TAILED GROUSE who refers to the whole district north of Cochrane to Moos- onee, states that ‘‘during the past fall [1932] it was nothing to see flocks of from 20 to 100 all over the district and at Mile 126 north, where they were not hunted and food plentiful, they were in, I believe, thousands’. Mr. Walter Kishbaugh, writing of the lake Abitibi region, states that during the ‘‘fall of 1932, large flocks migrated into this general district and into Quebec bordering lake Abitibi’. Mr. C. S. Jessup, writing of the territory south of lake Abitibi, states: ‘‘These birds started to come in last fall [1932] during the partridge season, early October, and were present in hundreds, at least until severe weather set in, in December.”’ The flight continued south of the Canadian National Transcontinental Railway, beyond what might be considered the normal wintering territory, and in reports from Kirkland lake, Englehart, the Elk lake region, New Liskeard, and Haileybury in northern Timiskaming district, Ont., and in the same general region in Quebec (see Dery, 19383), large numbers were reported over a widening territory. The time of arrival in this section of Ontario and Quebec was variously stated as from mid-October to the end of the month. It is obvious that the flight during this period was quite sudden, continuous, and with decided direction, between two and three hundred miles being encroached upon, from Moosonee, during approximately the first two or three weeks. The majority of the reports of first appearances south of lake Timiskaming are for late October and November. The flight, although uninterrupted, was apparently losing impetus, and statements, such as that of Mr. W. B. Greenwood, that the birds ‘‘worked south steadily in large numbers’’ during November,.picture the flight toward its limits in late fall and early winter. The sharp-tailed grouse reached the region of North Bay in December. Record dates, from central and southern Algoma, and Sudbury districts of Ontario, although approximately on the same latitude as those from Timis- kaming, were for slightly later periods, namely from late October and November. The flight reached its western limits in this latitude on the shores of lake Superior, from sections somewhat north of Sault Ste. Marie to Rossport. Records from northern and eastern Thunder bay district, EMIGRATION OF THE SHARP-TAILED GROUSE 17 from Rossport, Long Lac, Armstrong (specimen), eic., were probably of birds which came more or less directly from the Hudson bay watershed and not from the concentrated flight which poured south from the end of James bay. A specimen taken on the Albany river north of Nakina on October 22 is particularly suggestive on this point. It would appear that the fan-like shape of the area covered by the flight as indicated by the accompanying map was approximately correct in outline; that the shape was not entirely produced by the broadness of the southern territory from which reports could be obtained and the narrowness of the line of points from which observations were made in the north. By late November the flight was practically spent, but further extralimital records were reported, probably through the casual but more or less southward wandering of flocks throughout the winter. The extreme southern record which is definite is that of a specimen taken in the third week of March near Bracebridge, Muskoka district, in Ontario, although there is a further report by Mr. G. B. Greene of ‘birds in appreciable numbers’’ a few miles south of this at Gravenhurst. In Quebec the flight reached the southern parts of Pontiac and Papineau counties according to Mr. James C. McCuaig. It might be pointed out that the above is largely con- cerned with the time and numbers of the first appearances of these birds in various sections. The actual numbers in the more southern or outlying areas varied somewhat and nearly all were, of course, fewer than those noted in the north. It should also be repeated that the flight was continuous in the extreme north for at least three weeks, and that great numbers of birds remained along the route here and there as the flight progressed, and were present along the entire line of flight until well into the winter. Additional statements of numbers of birds observed follow in the section of this paper which deals with the habits of the northern sharp-tailed grouse. On the Falland Winter Habits of Emigrating Shar p-tatled Grouse Flocking It is apparent from the above remarks that the northern sharp-tailed grouse appeared in flocks during their emigration 18 A STUDY OF THE SHARP-TAILED GROUSE and that the flocks were somewhat larger toward the source of the flight. The counts and estimates of the number of individuals composing these flocks varied from four to ‘‘more than one hundred’’, with fifteen to twenty-five appearing to be an average, or a flock-size most commonly observed. At times several flocks of these birds congregated in very local areas to feed. Perching The reports invariably indicate that this grouse prefers open situations, a point to be discussed later. Trees, how- ever, serve as perches, both normally and after alarm. Mr. A. H. Burk writes that ‘‘over fifty pin-tails were counted in one dead pine tree’’ on January 11, in Beaumont township, Sudbury district. Many observers point out that they use the extreme tops of evergreens as perches and are frequently seen on post-like rampikes. When alarmed, the birds fly to the woods, or at least to trees, usually settling within one or two hundred yards of where they were put up. Several reports state specifically that the birds fly to the tops of “the tallest trees’’ near at hand. It would appear quite obvious that the northern sharp-tailed grouse selects open or lookout perches rather than cover. Flight Although little could be conveyed as to the nature of this grouse’s flight, it is apparent from the reports that the observers considered it swift and direct. There was no inclination to fly a dodging course through cover but rather to fly over wooded territory encountered in flight. Habitat The answers to the Museum’s questions relative to the habitat of the northern sharp-tailed grouse give a rather complete list of open situations to be found in the north. No observer suggests that the forest proper was the usual habitat. The following is a compilation of situations where the birds were reported to be found: sparse tree growth and barren sand plains where trees are scattered; ‘‘cranberry EMIGRATION OF THE SHARP-TAILED GROUSE 19 marshes’ and muskegs; recently burned-over areas and older brulés; exposed rock outcroppings; clearings and burns with low second growth; borders and surfaces of small frozen lakes and ponds; logging camps; roads; grain fields; farm-yards; railroad right-of-ways; inhabited clearings and settlements; urban gardens, streets, and even pavements. These are sufficient to indicate that the northern form of the sharp- tailed grouse frequents open situations not unlike the prairie representatives of the species. Roosting In answer to a specific question concerning the habit of burrowing into snow for roosting, there is general agreement that the birdsdoso. This, too, is like the sharp-tailed grouse of the prairie. | Fearlessness In regard to the exhibition of fearlessness in the northern sharp-tailed grouse, a few quotations bearing on this point are as follows: Mr. G. B. Greene writes: “In Timmins, on the afternoon of Tuesday, November 29th, a friend of mine living in the heart of the town counted 42 [northern sharp-tailed grouse] feeding in the yard with his chickens.’ Mr. Frank A. Carter, in comparing the behaviour of the sharp-tailed grouse with other native species, remarks that they are ‘‘very much tamer, coming close to buildings and in cases feeding with chickens’. Mr. W. G. Armstrong states that “they alighted on the houses and in the streets of Elk Lake; on houses and in the streets and on the railway platform of Swastika; and around the mines at Kirkland’’. Only one observer records observations more extreme than the above. He states that these birds could be “killed with a stick’’, but gives no further particulars on the matter. To assume that these emigrating birds were in a state of bewilderment seems unwarranted. It is not clear to the writer whether these descriptions of fearless habits can be interpreted as abnormal or not, since so little is known of their normal behaviour. A comparison with the winter behaviour of prairie sharp-tailed grouse suggests a similarity and therefore it would appear that the 20 A STUDY OF THE SHARP-TAILED GROUSE fearlessness of the northern grouse can be interpreted as not abnormal. Food Thirty-three crops and stomachs of birds secured in the flesh were preserved at the Royal Ontario Museum of Zoology. These specimens, all of which contained food, have been examined, largely by the Food Habits Research Branch of the Bureau of Biological Survey, Washington, but some material was analysed by Mr. C. H. D. Clarke, Department of Biology, University of Toronto. Dery (loc. cit.) has reported on eighteen well-filled stomachs from birds taken during the 1932-3 emigration and this information also has been used. In the report made by Dery, catkins and buds of Ostrya virginiana are recorded as comprising a high percentage of the items taken by the sharp-tailed grouse. Since this tree, namely ironwood or hop hornbeam, does not grow within the region concerned, it is obvious that the materials found were parts of paper birch, Betula papyrifera, which many authors agree resembles the ironwood very closely. The error is a natural one, especially since the food studies could not be checked by work in the field. Incidentally, the correction is entirely consistent with the results of many other stomach examinations of birds originating in the same general region. The birds collected were in good condition, and, except for one listed by Dery, and not used here in estimating percen- tages, crops and gizzards were well filled. The results of the food examinations for all specimens have been compiled in the table given below and the percen- tages worked out for a convenient summary.* ‘The first column of figures indicates the bulk quantity of the various items of food while the second column suggests the general acceptance or availability of the various items. *Since the above was written and the table compiled, twenty-five additional examination reports have been received from Mr. Dery. A comparison of these data with the table shows that the inclusion of these reports would not greatly alter the percentages of the variousitems. A revision of the table has, therefore, been considered unwarranted in the circumstances. EMIGRATION OF THE SHARP-TAILED GROUSE 21 Foop oF 51 NORTHERN SHARP-TAILED GROUSE TAKEN FROM OCTOBER, 1932, TO Marcu, 1933 Per cent. of Bulk percent.| birds which Food items found in 51 stomachs of all food | fed on each consumed item Catkins, buds, and twigs of paper birch, Betula alba and Betula sp. 46.9 88 Fruit, seeds, and twigs of mountain ash, Pyrus amert- cana 17.9 51 Stems, buds, and fruit of blueberries, Vaccintum sp., and small cranberry, Vaccinium oxycoccus 7.6 37 Buds and twigs of willow, Salix sp. 3.9 10 Buds and twigs of aspen poplar, Populus tremuloides 3.2 26 Fruit and seeds of wild rose, Rosa sp. 3.0 43 Catkins, buds, and twigs of beaked hazel, Corylus rostrata Pan | 24 Catkins and leaves of speckled alder, Alnus incana and Alnus sp. 2.5 30 Seeds of sedge, Carex flexuosa, C. oligosperma, and C. tenella 2.4 16 Buds, seeds, and twigs of black cherry, Prunus sero- tina, and pin cherry, Prunus pennsylvanica 2.3 20 Buds, leaves, and twigs of Labrador tea, Ledum groenlandicum 1.8 8 Seeds of juniper, Juniperus horizontalis and J. communis 1.6 20 Seeds of bunch berry, Cornus canadensis Bin 36 22 A STUDY OF THE SHARP-TAILED GROUSE Foop or 51 NORTHERN SHARP-TAILED GROUSE TAKEN FROM OCTOBER, 1932, TO Marcu, 1983—Continued Per cent. of Bulk percent.| birds which Food items found in 51 stomachs of all food fed on each consumed item Buds and twigs of tamarack, Larix laricina; leaves of vetch, Vicia sp.; leaves of shield fern, Aspidium intermedium; seeds of Solomon’s seal, Polygonatum biflorum; seeds of bramble, Rubus sp.; seeds of mayflower, Maianthemum canadense; seeds of arrow-wood, Vibernum opulus; seeds of dogwood, Cornus paniculata; black bindweed, Polygonum convolvulus; leaves of white clover, Trifolium re- pens; flower-heads of wild lettuce, Lactuca sp.; seeds of green brier, Smilax sp. 3.1 40 Seeds of bristly sarsparilla, Aralia hispida; fruit of hawthorn, Crataegus sp.; seeds of crab grass, Digit- aria sanguinalis; seeds of galingale, Cyperus fili- culmis; leaves of saxifrage, Saxzfraga sp.; leaves of ground ivy, Glacoma hederacea; seeds of sumac, Rhus sp.; beetles, Buprestidae, Curculionidae, Carabidae, Cantharts (larva), Lathrobiwm. Trace Gravel a 25 Feathers a less than 2 SPECIMENS FROM THE 1932 FLIGHT AND THE DISTRIBUTION OF RECORDS Specimens So far as it has been ascertained, sixty-nine specimens from this flight have been preserved in scientific collections. The writer has examined twenty-four of these and it is noteworthy that this series is remarkably uniform. The characters of the race are constantly displayed and the range of individual variation is not at all wide. The present occasion is appa- rently the first afforded to examine a satisfactorily extensive series of specimens which had their origin in the Hudson and James bay region. EMIGRATION OF THE SHARP-TAILED GROUSE 23 An attempt was made to ascertain the proportions of the sexes as determined from dissected specimens. The sex of sixty-five specimens has been determined, forty-eight of which were females and seventeen males. No reason can be suggested why the sex ratio of the specimens collected may not represent the sex ratio of the bird in nature, at least during the 1932 emigration. Distribution of Records The accompanying map shows in some detail the distribu- tion of the records of occurrence of the northern sharp-tailed grouse in Ontario and Quebec during its recent emigration. Reports which aroused an element of doubt as to their reference to the form concerned have not been included. Actual specimens substantiate a large number of the place records indicated. ’ i} U ' Re { { | if Map J.—The 1932 flight of Pedioecetes p. phasianellus based on observations of occurrence and on collected specimens. 24 A STUDY OF THE SHARP-TAILED GROUSE THE STATUS OF THE SHARP-TAILED (GROUSE DURING THE YEAR SUBSEQUENT TO THE 1932 EMIGRATION The Museum's first questionnaire, sent out during the winter of 1932-3, requested information on the sharp-tailed grouse in general and on the emigration of northern sharp- tailed grouse in particular. During the following winter, 1933-4, another questionnaire was distributed, which at- tempted to ascertain a possible diminution of sharp-tailed in general, but was more particularly directed toward the status of the northern sharp-tailed grouse during the year subsequent to its emigration in 1932-3 in the area covered by its flight. It may be here mentioned that in general the numbers of A.O.U. P.p. campestris following the winter of 1932-3 remained about the same, 7.e., the prairie sharp-tailed grouse was still plentiful. Dr. H. F. Lewis writes ‘that ‘‘when travelling from Winnipeg to Kenora by train on March 3, 1934, I counted 47 individuals of this species from the train window’. The population of the eastern limits of this form in western Thunder bay and the Rainy river district appa- rently showed some local decrease by the fall of 1933. Several observers estimate the numbers as being only ten per cent. of those found the previous year. In other sections of the area and into Manitoba and westward, however, the popula- tion was not notably changed, the grouse remaining plentiful. Criddle (loc. cit.) has shown that peaks of numbers of this form last from two to three years.* Reports from South of Canadian National Transcontinental Railway Several of the first reports on the flight of P. p. phasianellus in the east record the disappearance of these birds during midwinter or when the snow and cold weather setin. Others reported flocks remaining through the winter. The reports as a whole demonstrate that these birds disappeared suddenly in several cases, and gradually in others, largely during the *Reports from Rainy River District, Ont., and the Winnipeg area of Man- itoba in February, 1935, indicate that the sharp-tailed grouse population was greatly reduced. EMIGRATION OF THE SHARP-TAILED GROUSE 25 winter of 1932-3, but that locally they persisted longer. Here and there throughout the area south of the National Transcontinental Railway in Ontario and Quebec, in territory considered by the writer as more or less extralimital to the race, birds were observed until spring, after which they were not observed. Only three observers south of this line in Ontario and Quebec report, without details, that birds appa- rently migrating northward were seen in the spring of 1933. Dr. H. F. Lewis reports that during an extensive trip in Cochrane district to Moosonee and the Abitibi region in March, 1934, he was particularly on the lookout for sharp- tailed grouse and saw but ‘‘three individuals in one group in tamarack trees near the Canadian National Railways track just east of Norembega, Ontario’. A total of seven observations of young birds or nests with eggs was reported from south of the normal breeding area in the summer of 1933, while six other observers reported summer occurrences of adults. These observations are from points well distributed through the districts of Timiskaming (Ontario and Quebec), Nipissing, Sudbury, Algoma, and the southern part of Cochrane (Ontario). There may be error in any one of these reports but in the aggregate there is unquestionable evidence that the species occurred and nested in the summer of 1933 in territory occupied by the emigrants. Some of the reported summer occurrences were of stray individuals, while others were of flocks which remained together throughout this season. This may have a relation to the sex ratio of the emigrants as previously noted, which will be mentioned again in a following section of this paper. Little can be recorded on the nesting of the northern sharp- tailed grouse in this extralimital territory. Some miscel- laneous details may, however, be included here. In con- nection with the number of eggs or young, one observer mentions a nest with nine eggs (of a “greenish tint’’) while another reports eight downy young with a female. Concern- ing nesting sites, one report states that the species nested in an ‘‘area burned over in 1932”’. 26 A STUDY OF THE SHARP-TAILED GROUSE Reports from North of Canadian National Transcontinental Railway A number of observers in this area were inclined to believe that there was a northward movement of sharp-tailed grouse in the early spring of 1933. Several observers record nestings and summer (1933) occurrences in numbers which might be expected except when this grouse is at an exceedingly low status of numbers. They nested numerously about Coch- rane in that year but disappeared in August. No notable influx or obvious concerted movement southward or in any other direction was noticed in the fall of 1933 in this region although the species was again noted sparingly from early November onward. NOTES ON THE MATING HABITS OF THE NORTHERN SHARP-TAILED GROUSE Although the mating habits or the ‘“‘dance’’ of male prairie sharp-tailed grouse have been fully and repeatedly described, it is of interest to note that the habit is also practised by these northern birds. Mr. R. V. Whelan of Smoky Falls, Ont., observed this performance on May 14, 1933, and writes: As I prepared to leave my house at 7.00 a.m., I heard a peculiar swishing sound close to my greenhouse, which is about 75 feet from my house. Looking in that direction, I saw that it was caused by the tail of a sharp-tailed grouse in a mating dance. With the tail held perpendicular to the body and fanned out, it was vibrated rapidly from left to right; with wings out- stretched and head close to the ground so that the erect yellow combs were plainly discernible, neck swollen so that it showed up plainly purple, he rapidly circled the female at high speed with short mincing steps, moving as though he were floating. The female apparently paid no attention to him, but went on feeding. He paid no attention to my approach and I watched him with glasses for some time. Finally the approach of another observer put the female to flight and the male followed. I watched him with the glasses during the flight and observed that the yellow comb remained visible. EMIGRATION OF THE SHARP-TAILED GROUSE OF FACTORS WHICH WORKED TO DIMINISH THE NUMBERS OF NORTHERN SHARP-TAILED GROUSE IN THE EAST FOLLOWING THE EMIGRATION OF 1932 That the great numbers of sharp-tailed grouse which poured southward from James bay over central and east central Ontario and western Quebec during the fall of 19382 and the following winter, disappeared, for the most part, is obvious. There was no northward movement which could be interpreted as a general retreat to customary nesting territory. A few instances of nesting probably took place but more commonly, according to reports, remnant flocks and individuals wandered about during the breeding season. It should be remarked that by the advent of the nesting season the numbers had notably decreased. A careful review of all the reported destructive factors does not uncover a major one which would seem to have caused the disappearance of these birds. Perhaps the great- est number killed by a single known cause were those shot forfood. There was some confusion as to whether these birds were protected under existing game laws but the question did not amount to a general restraint, and there is no doubt that large numbers were secured during the height of their southward movement. As one observer states, ‘‘they were killed very freely in portions of northern Ontario’. It must be remembered that economic conditions during the period constituted an abnormal reason for heavy pressure on any game supply, and locally the appearance of sharp-tailed grouse would probably have been felt as an extremely for- tunate and opportune increase of food. Reports from Quebec and Ontario during the fall of 1932 explain that settlers were “stocking their larders’’, the birds being salted for winter use. In many places the birds were shot during the ‘‘partridge’’ season as game and although probably for the most part bag limits were recognized, there are reports that ‘‘one would have no trouble in bagging 100 [sharp-tailed grouse] in a single day’’. Although the bagging of large numbers of these grouse is generally admitted by observers throughout the north, some of them are positive that this was a minor contributing factor to their disappearance. A few considered 28 A STUDY OF THE SHARP-TAILED GROUSE that humans were largely responsible for their disappearance and the writer is inclined to believe that this factor was of notable importance. The second Museum questionnaire indicated that shooting of these birds was continued in the fall of 1933, in areas where remnant flocks persisted. The few other known destructive factors can fairly be classed as miscellaneous. In no section of the territory covered by the flight were predators obviously more numerous. A few observers remark on a possible slight increase locally in the case of some predaceous forms, but far more observers state that such an increase was nil or that these forms were decidedly scarce. There are no actual observations of north- ern sharp-tailed grouse being caught and devoured by natural enemies. However, it seems reasonable to suppose that a few of the birds from this flight served as food for the various Carnivorous mammals and raptorial birds. Mr. R. V. Whelan mentioned specifically that the northern sharp-tailed grouse was prone to fly into transmission wires and buildings about the town of Smoky Falls, Ont. Reports from other settlements, however, do not suggest that this cause of death was widespread. This observation would seem to be due to a prevalence of hazards around the power plants at Smoky Falls rather than an indication of the so-called ‘‘madness”’ in the flight of this grouse which has been used to explain similar casualties which occur, not infrequently, in the case of other species of grouse. Other similar accidental causes of death probably accounted for the disappearance of a very few sharp-tailed grouse during their southward flight. Adverse weather conditions were frequently suggested by observers in the north as contributing to the disappearance of these grouse. This factor, of course, was usually associated with food supply. There is, however, no unity notable in the reports on weather conditions. Such communications as ‘“‘the lack of snow, together with rain and sleet followed by extreme cold weather had destroyed them in large numbers”’ were received. This statement, forwarded by Mr. C. B. Greene, says further: ‘“‘At least 50 dead [northern sharp- tailed] grouse [were seen] on the privately owned railway right-of-way north of Iroquois Falls, Ontario. There was very little snow on the ground; the temperature that day was EMIGRATION OF THE SHARP-TAILED GROUSE 29 between 30 and 40 degrees below zero, with a heavy wind, which condition followed a bad sleet storm which seemed to be more or less localized between Smooth Rock Falls, Ontario, and La Sarre, P.Q., on the National Transcontinental Rail- way.’’ Further correspondence on this observation was received from Mr. C. B. Davis, who states that the dead grouse ‘‘were found in early December before the heavy snows came. During this period we had exceptionally cold weather and high winds. Locally, it was assumed that the birds were unable to get protection, and froze to death.’’ The habit of the northern sharp-tailed grouse of spending the night beneath the snow, a practice prohibited by the absence of deep snow, lends strength to the plausibility of this explanation. In only two other localities, namely Moose river basin and the vicinity of North Bay, were specimens reported found dead. No cause was apparent in these cases, and it involved but one bird in one case and a very few birds in the other. Although opportunity was not offered for anything like a complete study of the health of emigrating sharp-tailed grouse, eleven post mortem examinations were made by Mr. C. H. D. Clarke of the Department of Biology, University of Toronto. All the birds examined had been shot as speci- mens. No individuals found dead were available for study. For the most part the findings were nil. The absence of parasitic worms in these northern birds was remarkable and there was no excessive case of parasitism. The few Mallo- phaga found on specimens of P. p. phasianellus have been determined as Lipeurus protervus and Goniodes mammulatus by Dr. H. S. Peters of the United States Bureau of Entom- ology. Both species were found on individual birds on occasion. It is of interest to note that these parasites were found on both phasianellus and A.O.U. campestris. A DISCUSSION OF THE FINAL PHASES OF THE EMIGRATION There is no evidence which suggests that the numbers of these emigrating birds were decimated by one or more conspicuous and potent agencies. Shooting for food and sport, however, very probably diminished their numbers very appreciably locally. Other factors, namely, adverse weather conditions and food shortage, natural enemies, accidents, 30 A STUDY OF THE SHARP-TAILED GROUSE parasites, and disease probably took a steady toll, but none was a conspicuous agent of destruction. (ey alm ROLLEI Sm hots ed BP PAOD Bie es