Changes in the Fauna of Ontario: A Contribution of the Division of Zoology and Palaeontology 9781487583699

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Changes in the Fauna of Ontario

Edited by

F. A. URQUHART

Published by UNIVERSITY OF TORONTO PRESS for the Division of Zoology and Palaeontology Royal Ontario Museum

R©M IOYAL ONTA.BIO IIUSEUII

Copyright©, Canada, 1957, by University of Toronto Press Printed in Canada London: Oxford University Press

Reprinted in 2018 ISBN 978-1-4875-8234-0 (paper)

FOREWORD IT IS INDEED most difficult to study, with any degree of accuracy, the possible changes which may have taken place over a period of years in any defined geographical area with respect to the distribution of the various species of animals, when no accurate distributional notes have been recorded in print for that period. We rather glibly speak about .a certain species of animal as being "new to the district" or having "arrived within recent times" with little, if any, factual data to substantiate our claims. For our distributional data we are forced only too often to rely on the words of some early voyager or inexperienced naturalist who had recorded scattered notes and observations in his diaries, and, as so often happens, we find that there have been numerous errors with respect to identification. During the past forty-three years our Division of Zoology of the Royal Ontario Museum has been most active in collecting various species of animals throughout the Province of Ontario, with particular emphasis on the local fauna of the Toronto region. During this time a number of outstanding naturalists have not only made careful collections of animals, but have also recorded accurate observations on the species with which they were familiar. Those who have been active in this field of research have been impressed by the changes that apparently have taken place during the past few decades with respect to the distribution of the various species of animals, with particular reference to our indigenous fauna. Having observed such changes, we immediately ask, "What are the possible factors responsible for these changes?" It is, of course, most difficult to analyze and evaluate the effect of factors operating within a geographical area that might account for such changes in the distribution of the fauna without knowing exactly what changes have definitely taken place and to what extent. We can evaluate such changes only in the light of our experience over a period of many years. In order that we do this, it is essential that observations covering short periods of time be regularly recorded. With such accurate data available covering short, consecutive periods, we can then compare these data with changes which have been recorded respecting meteorological or other factors. This publication, we most sincerely hope, will be the first of many similar contributions to the study of the distribution of the fauna of Ontario. P.A. U.

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CONTENTS FOREWORD

iii

Introduction: Factors Influencing the Changing Fauna of Ontario. By F. A. URQUHART

1

Changes in the Insect Fauna of Ontario (with Special Reference to the Orthoptera). By E. M. WALKER

4

Changes in the Amphibian and Reptilian Fauna of Ontario. By E. B. S. LooIER

13

Changes in the Fish Fauna of Ontario. By W. B. ScoTT

19

Changes in the Avifauna of Ontario. By L. L. SNYDER

26

Changes in the Mammalian Fauna of Ontario. By RANDOLPH L. PETERSON

43

Changes in the Climate of Ontario. By M. K. THOMAS

59

iv

INTRODUCTION: FACTORS INFLUENCING THE CHANGING FAUNA OF ONTARIO BY

F. A. URQUHART

THERE WERE, apparently, three centres of ice-sheet formation on the North American continent during the glacial period of the Pleistocene epoch: Cordilleran, Keewatin, and Labrador. Of these, two, namely, the Keewatin and the Labrador, covered the greater part of Canada and the eastern United States, and these two were the most important in affecting the distribution of Ontario's indigenous fauna. The Keewatin ice sheet was centred west of Hudson Bay, in the Keewatin region. It extended as far west as the Rocky Mountains, as far south as latitude 38° N., and east over much of the Hudson Bay area. The Labrador ice sheet was centred near the southeast end of James Bay. It covered most of eastern Canada and a great part of the northeastern United States. As the Keewatin ice sheet withdrew northward, a large glacial lake was formed, named Lake Agassiz, which occupied parts of SaskatchewaR, Manitoba, and western Ontario. Remnants of Lake Agassiz still remain, forming Lake Winnipeg as well as other lakes in Manitoba. Coincident with the withdrawal to the north of the Labrador sheet, Lake Algonquin, a glacial lake that occupied the area which is now covered by Lakes Superior, Michigan, and Huron, was formed. In addition to this large glacial lake, a second and smaller one, termed Lake Iroquois, was also formed which occupied a much smaller area, including the area now occupied by Lake Ontario. As the Keewatin ice sheet withdrew from the Labrador ice sheet, a land space was formed between Lake Agassiz to the northwest and Lake Algonquin to the southeast. A land space also occurred between Lake Algonquin to the west and Lake Iroquois on the east. Animals moving northward into Ontario as a result of the retreat of these large ice sheets, would encounter three large water barriers: Lake Agassiz to the northwest, Lake Algonquin to the north, and Lake Iroquois to the east. Such animals, however, would find easy land access to Ontario by moving northwestward, or north between Lake Agassiz and Lake Algonquin, and thence eastward, arriving thus at the land area to the north of Lake Algonquin. They could also move straight northward entering Ontario between Lake Algonquin and Lake Iroquois. Finally, they could move north or northeastward around the east end of Lake Iroquois and thence westward. As viewed at the present time, there are, therefore, three main routes that could have been taken by southern animals entering into Ontario as they

pushed their distributional limits northward. One of these, from west to east, would be around the north shores of Lake Superior and Lake Huron. A .second route would be from south to north in the vicinity of the St. Clair River, the west end of Lake Erie, and the Niagara Peninsula between Lake Erie and Lake Ontario. A third route would be across the St. Lawrence River in the vicinity of the Thousand Islands. It would appear that the invasion into Ontario of southern animals might follow the routes as outlined. If such is true, we might expect more species to enter by the St. Clair River and the Niagara Peninsula route than by the western route north of the Great Lakes and the eastern route east of Lake Ontario, because of the greater distance involved in the latter, together with physiographic difficulties. We should not expect as much, if any, movement from east to west or north to south, since such would be simply movements within the area on the part of established populations now located in the province. There have been numerous minor and a few major fluctuations in temperature and humidity throughout the province during the past hundreds of years, with minor fluctuations, of some importance, during the past century, as pointed out in the chapter on climate change in Ontario by M. K. Thomas. If temperature changes are taking place, it will be of considerable importance to record accurately any changes that may take place in the fauna of the province as being possibly correlated with such changes in meteorological factors. Such changes will be evident not only through the appearance of new species entering the province from the south, but also through the movement northward of southern species already indigenous to the province. In addition to meteorological changes, there are also physiographic changes going on over the province which bring other factors into play in limiting the distribution of the various species of animals. Trees are being removed, thus converting forested areas into grassland and permitting grassland species to spread over a far greater area. Coincident with the removal of trees and the resultant destruction to the forest undergrowth, there is a greater amount of water run-off, causing rivers and lakes to change their character from clear bodies of water with free gravel beds to muddy waters with silt-covered beds. Such changes may result in the complete disappearance of some species and the invasion into such changed waters by other species. With the construction of artificial waterways, dams, locks, and the like, various aquatic species will be permitted to spread from one body of water to another. During the past many years we have witnessed the disappearance, and in some cases the complete annihilation, of some species of animals. As examples of this, mention could be made of the Passenger Pigeon as a species of bird which has been completely extirpated, and the Wild Turkey, which has been extirpated from the Province of Ontario but is still found in other localities in North America. We have witnessed also the introduction of certain species of animals from other countries, e.g., the Starling and the European Hare. Such changes in our fauna are interesting from the point of view of the change 2

that is taking place, but it has very little, if any, real significance in the study of climatic factors and other factors affecting the distribution of our present indigenous fauna. It may be, however, that with the destruction of forest areas some species of animals in the province which are now of rare occurrence may be extirpated. Such changes can only be followed with accuracy by making accurate observations over a period of years and recording such observations in the literature of the period. As these changes in climate, physiography, and distribution of the various animals take place, it is essential that zoologists and naturalists make accurate observations, identifications, and recordings of pertinent data in order that such information will be available to assist the student of the future to understand the causes of changes in the fauna of any particular area. It is quite likely that such studies will help not only to explain what has happened in the past, but to forecast what may take place in the future.

3

CHANGES IN THE INSECT FAUNA OF ONTARIO (WITH SPECIAL REFERENCE TO THE ORTHOPTERA) BY

E. M. WALKER

THE INEXPERIENCED COLLECTOR of insects is not concerned with the question as to what sort of collecting the distant future may bring. It is not until he has collected some special group intensively over a limited area and has obtained a detailed knowledge of the regional species, that he is in a position to appreciate any future changes in the composition of that group, which may take place from time to time. Having collected and observed insects myself for over sixty years, chiefly in southern Ontario, I have noted many changes in the insect fauna of this region. We shall not be concerned in this paper with those changes which are directly due to man's interference with the original environment nor to the introduction of exotic insects, but to changes in the distribution of native insects resulting in the disappearance of some species from localities where they were formerly established and the immigration of other species into new localities. After a few years of general collecting, which commenced in August, 1890, I became specially interested in the Orthoptera. Most of my collecting was done in the vicinity of DeGrassi Point, Lake Simcoe, and in the course of a few years I had found all the species of this order that were living within ten miles or more of my summer residence. I also became familiar with the Orthoptera of the Toronto district but I shall select DeGrassi Point as the representative locality, since it has suffered little from human interference, whereas Toronto has been ruined as a collecting area. Only the grasshoppers and crickets will be considered since the non-saltatorial species of Orthoptera of our selected region are insignificant from our point of view. Most of the biotypes of grasshoppers and crickets are represented in this area but not quite all. There are no extensive beaches nor sand dunes and no good examples known to me of the open sphagnum bogs that are so common on the Canadian Shield. The beach and dune biotype, however, is well represented in Simcoe County by Wasaga Beach and, in the same general latitude, by the beach at Southampton, Bruce County, where I collected in 1901 with Professor John Macoun and again in 1935 with Dr. John Oughton. In the eighteen-nineties it is probable that the Orthoptera of Lake Simcoe were nearly all species that were resident there when the land was first cleared for settlement. There were only thirty-six species of grasshoppers and crickets native to the vicinity of DeGrassi Point arid of these original species, as we may call them, at least seven have disappeared from DeGrassi 4

Point itself and all but one or two of these species have apparently vanished from the entire district. The seven species are as follows: ACRIDIDAE ( Short-horned

Grasshoppers) Stethophyma lineatum (Scudder) Stethophyma gracile (Scudder) Circotettix verruculatus (Kirby) Zubovskya glacialis variegata (Scudder) Melanoplus fasciatus (F. Walker)

TETTIGONIIDAE

Scudderia pistillata (Brunner) Orchelimum vulgare Harris

Stethophyma lineatum is a large handsome grasshopper widely distributed in open sedgy marshes. It was formerly common in a small marsh of that type at the mouth of our nearest stream, about half a mile west of the Point. This marsh has been gradually changing in character, as the bay into which the stream flows is slowly filling in. Alders are invading the marsh and the sedges, grasses, and other emergent plants are giving way to boneset, asters, and other plants indicative of a drier type of marsh. The disappearance of S. lineatum is probably due entirely to ecological changes that are independent of any alteration of climate. Stethophyma gracile is a boreal grasshopper that formerly occupied a narrow irregular zone between the open marsh inhabited by S. lineatum and a tamarack swamp. It was partly shaded by the tamaracks but mainly open and there was a belt of leatherleaf ( Chamaedaphne calyculata) between the open sedgy marsh and the bog vegetation of the tamarack swamp. With the cutting away of most of the tamaracks this small habitat of S. gracile was altered and the species became locally exterminated. It was close to its southern limit of distribution at this station and its disappearance may have been due entirely to local causes although climatic changes, such as higher temperature, are not excluded as a possible factor. The next on our list is Circotettix verruculatus. This is the dark gray grasshopper with yellow hind wings that makes a loud snapping noise as it flies over rocks and open sandy barrens, all across the wooded parts of Canada from Newfoundland to British Columbia. When I first collected insects at DeGrassi Point it was common within a mile of the Point, near the village of Lefroy. Here is an area of some ten to fifteen acres of sandy soil, overlying a bed of gravel, part of the old lake beach. Blackened logs and stumps of white pine showed that this area had been burned over, probably many years ago, following lumbering operations. Small aspen poplar, staghorn sumach, and a few second growth pine, with plenty of bracken and thistles occupied this site when I first saw it, and the loud crackling of Circotettix could be heard on any hot sunny day in late July or August. About half a mile northeast of this area is the railway station of Lefroy and I often saw and heard 5

Circotettix here, as it flew over the cinders of the railway tracks. It also occurred at Bradford, ten miles farther south, where I found it flying over the sawdust of a sawmill. South of Bradford I have recorded it from the vicinity of Aurora, but it was certainly rare or very local there even at that time and was unknown nearer Toronto. It has now completely disappeared from this entire territory, although it persists on the high rocky escarpment at Credit Forks in Peel County unless it has vanished within the last ten or twelve years. It flies over the sand dunes at Wasaga Beach in the extreme north of Simcoe County. Unfortunately we have no record of the time of its disappearance from the vicinity of DeGrassi Point and Lefroy, but it was certainly more than thirty years ago. The gravelly area where Circotettix used to be abundant has been waste land ever since. It has been burned over twice but the sumachs and bracken are still there and it is still largely open. A thin growth of grass (Danthonia spicata), which now covers much of the ground that was once nearly devoid of grass, may have influenced unfavourably the ovipositing habits of Circotettix, but the general disappearance of this insect from an area of some hundreds of square miles must be due. to some other factor. It is still common from Muskoka northward or was so within the last decade. Zubovskya glacialis is another northern insect whose southern limit in Ontario, so far as known, was the wooded swamps in Simcoe and Bruce counties. The genus Zubovskya is a near relative of Melanoplus, the genus to which our commonest grasshoppers belong, but it is completely wingless, the male black above with a creamy yellow stripe along the whole length of its thorax and abdomen, whereas the female, which is much larger, is olive green. I mention this type of coloration and Zubovskya's wingless condition because they are so often associated with the alpine habitat, and the typical subspecies of Z. glacialis comes from the White Mountains of New Hampshire, where it has been found above the tree line, although it also lives at sea level in Maine and the Maritime Provinces. In Ontario it occurs as a slightly different subspecies (Z. g. canadensis Walk.) in Algonquin Park, and here it is not confined to the lowlands. Southward, however, it becomes restricted to the boreal type of swamp, where tamarack, balsam fir, spruce, white cedar, and paper birch grow, particularly where there are raspberry bushes. Here it is regarded as a slightly different subspecies, Z. g. variegata Scudd. Even in my earliest days of collecting this was the rarest and most local of all the Orthoptera at DeGrassi Point, and although it persisted until about ten years ago, I believe it has finally died out. It is quite possible that it may still occur in some of the swamps in Simcoe or York County, but few of these swamps remain that have not been severely cut over. Melanoplus fasciatus, another northern grasshopper, whose habitat is dry, open, sandy woods or blueberry barrens, was formerly found not only at DeGrassi Point but even in High Park, Toronto. It is never a plentiful species but it is everywhere common in the north and was formerly easy to find at DeGrassi Point along the sandy borders of woods, especially pine woods. 6

Its habitat at DeGrassi Point has not been interfered with during the period we are considering, but M. fasciatus has nevertheless disappeared or become very rare. Dr. Urquhart tells me, however, that he has taken it recently at Point Pelee, Ontario, and its disappearance from DeGrassi Point would thus appear to be due to a local cause. The Point is now much more heavily wooded than it was fifty years ago and it may be that the woods are now too shady for M. fasciatus. Its rarer associate, M. islandicus, is still present, but it is undoubtedly more tolerant of shade than is M. fasciatus. Turning now to the Tettigoniidae, the two species listed were formerly abundant at DeGrassi Point but are now extinct or very rare. These are the Broad-winged Bush Katydid (Scudderia pistillata) and the Common Meadow Grasshopper ( Orchelimum vulgare). Both of these insects were at one time common in an area of about ten acres immediately behind a row of cottages on the lake shore facing east. This area was once pasture land with a few clumps of trees, but was allowed to become naturally reforested. Among the first plants to become dominant in this site after it was fenced off were two native grasses, the Indian Grass (Sorghastrum nutans) and the Forked Beard-grass (Andropogon Gerardi), each five or six feet high. Both species of grasshopper throve in this meadow for many years, but in the course of time the meadow became wooded and both meadow grasshopper and bush katydid disappeared. And even though an adjoining piece of pasture had meanwhile been enclosed and is still largely covered with the same two grasses, both of the grasshoppers have gone. Scudderia pistillata still occurs here and there within a few miles of DeGrassi Point but apparently only in open marshy places, as it did in the Toronto district fifty years ago. It is a northern species and, in the more favourable parts of its range, is generally distributed. As the southern limit is approached, however, pistillata becomes restricted to the cooler marshy or swampy areas. Thus the evidence from the Lake Simcoe region suggests that this species is retreating northward. The case of Orchelimum vulgare is different. It is a relatively southern species and the population at DeGrassi Point may have been an isolated one. Like Scudderia pistillata it flourished for years in the tall grass that grew in abundance in what was formerly a pasture but, as this area become afforested it did not spread to the adjoining meadow but died out. It was also common about Toronto and probably still is, but a factor in determining the presence of Orchelimum is the observer's ability to hear the stridulation of the male, a test which I am no longer able to apply satisfactorily. The apparent disappearance of Orchelimum vulgare from the vicinity of DeGrassi Point remains at present a puzzle, the solution of which requires more data. Having noted the partial or complete disappearance of seven species of Orthoptera from the vicinity of DeGrassi Point, Lake Simcoe, during approximately the last fifty years, we may now ask the question as to what other species, if any, have immigrated to this locality. Of the Acrididae the first new arrival was Encoptolophus sordidus 7

(Burm.). My first specimen of this insect was taken in Toronto on the Exhibition Grounds in September, 1892, and I was unable to find another in the Toronto district for several years, although it was plentiful along the Niagara River. In a few years it had become common at Toronto but it was not seen at DeGrassi Point until about 1900. Since then it has been very abundant and is thoroughly established as a resident species. Although belonging to the same subfamily (Oedipodinae) as the vanished Circotettix verruculatus, it cannot be said to have replaced this species since it inhabits dry grassy fields, not the bare rocky or sandy places preferred by Circotettix. Much later, however, in the late thirties, another member of the Oedipodinae appeared at DeGrassi Point, known as Spharagemon bolli Scudder. This species I had known from early collecting days in High Park, Toronto, but it was certainly absent from DeGrassi Point or any of my Lake Simcoe haunts. It was first seen at the Point on a little-used road in a dry, somewhat worn-out pasture, along the fringe of a wood, a very typical habitat for its kind. For several years it was very scarce and its status as an addition to the local fauna remained uncertain. Now, however, it has definitely increased in numbers and, having invaded the area formerly occupied by Circotettix verruculatus, it may be said to have replaced this species in some degree. Like the latter species it is a dark insect with the hind wings yellow and black bordered, and during flight it makes a low crackling sound with its wings. It is, however, much less conspicuous in both flight and stridulation. The distributional ranges of the two species are also very different. In general, C. verruculatus inhabits the northern half of North America, S. bolli the southern half, i.e., the United States from Texas northward, although it has now passed Lake Simcoe and has reached well into the Muskoka and Parry Sound Districts. In fact there were isolated colonies in the Muskoka District as early as 1903. Another species of Spharagemon, S. collare wyomingianum (Thomas), was first found in Ontario in 1901 on the sand dunes of Point Pelee, Essex County, and later at the southern end of Lake Huron. It was certainly not resident near Toronto, nor at DeGrassi Point, but it appeared at both localities during the nineteen-forties. A few years ago Dr. F. A. Urquhart told me it was one of the commonest grasshoppers in the vicinity of his home in West Hill, east of Toronto, and about the same time it had reached DeGrassi Point, where I saw it almost at my cottage door. It eventually spread to nearly all the sterile pastures and open sandy areas between DeGrassi Point and Lefroy, including the old bum where Circotettix was once so common. Like the other two oedipodine invaders wyomingianum comes from the south side of the international boundary, although the typical S. collare has long been known as an indigenous species in the Prairie Provinces. The species of Acrididae that have disappeared from the swamps, unlike those from the barrens and dry fields, have not been replaced by others, so that the total number of species of this family in the region typified by DeGrassi Point is still one less than it was in the nineties. 8

Referring now to immigrants of the family Tettigoniidae, we have already noted that Scudderia pistillata has vanished from DeGrassi Point but is still regional, although restricted to open marshy places. Its former habitat where Indian Grass and Forked Beard-grass grow is now occupied by another immigrant from the south, the common conehead or sword-bearer ( N eoconocephalus ensiger [Harris]). It was already in Toronto when I began to collect in 1890, but it did not reach DeGrassi Point until many years later. It was, however, one of the earliest immigrants there, arriving well before 1930. It is now a common and settled inhabitant of Simcoe County and has penetrated much farther northward into the Muskoka District. We also have a new bush katydid, Scudderia texensis Saussure-Pictet, which I first found in August, 1953, in a grassy field at the western limits of DeGrassi Point. Since then it has greatly increased in numbers and in range. Being a southern species, long known in Canada only from Point Pelee, Lake Erie, it was not expected at DeGrassi Point, although records during the last twenty years have extended its range to include the St. Clair River delta, Prince Edward County, and even Montreal. To summarize the changes in the Orthoptera of DeGrassi Point and its immediate vicinity, as described above, seven species have vanished and five species invaded and become established in this locality. These may be called respectively the emigrants and the immigrants. They are as follows: EMIGRANTS

Stethophyma lineatum Stethophyma gracile Circotettix verruculatus Zubovskya gracialis Melanoplus fasciatus Scudderia pistillata Orchelimum vulgare

IMMIGRANTS

Encoptolophus sordidus Spharagemon bolli Spharagemon c. wyomingianum Neoconocephalus ensiger Scudderia texensis

Five of the seven emigrants are definitely northern species, that is, all but the first and last, whereas all of the immigrants are relatively southern, their distributional ranges being chiefly in the United States. Before leaving the Orthoptera we may recall our remark that another biotype of this Order is found in Simcoe County, namely, the wide type of beach and sand dunes, which in Ontario is characteristic of the Great Lakes. It is best represented in Simcoe County by Wasaga Beach on Georgian Bay but similar beaches at about the same latitude are found at Southampton and Sauble Beach, Bruce County, on Lake Huron. At Wasaga Beach there are two species of Acrididae that are distinct from any known from the shores of Lake Simcoe. These are Melanoplus stonei Rehn and Trimerotropis huroniana Walker. The former species does not concern us here but the latter species does. I first discovered it in 1901 on a collecting trip with Professor John Macoun in southwestern Ontario. 9

We had collected at various stations along Lake Erie and at Point Edward on Lake Huron and everywhere along the beaches I was struck with the abundance of the Sea-beach Grasshopper (Trimerotropis maritima [Harris]), a maritime relict which I first met with on Toronto Island and later found on the sea beach of New Jersey. We next visited Goderich, whose steep cliffs and narrow beach apparently created a barrier against the spread of T. maritima farther north. At our next stop, Southampton, however, there was a fine broad beach, but instead of T. maritima again, I found a new species of Trimerotropis apparently occupying precisely the same biotype on the beach, the zone of beach grass (Ammophila breviligulata Fern.). I recognized it as different from maritima first by its interrupted stridulation. It turned out to be a closer relative of one of the species of the Great Plains than to maritima but its pale, speckled coloration was so similar to that of maritima that it could· be easily mistaken for that species. During later years I found this new species (T. huroniana Walk.) on the beach of the Giant's Tomb Island, Georgian Bay, and still later on Wasaga Beach, also on Georgian Bay. In 1935, thirty-four years after T. huroniana was discovered at Southampton, Dr. John Oughton and I made a similar but more detailed survey of the same· parts of southern Ontario as I had covered in 1901 with Professor Macoun. I was curious to know where the range of T. maritima would end and that of huroniana would begin. So we stopped at many stations all along the coast of Lake Huron. First we discovered that the cliffs of Goderich were no barrier after all to the northward spread of T. maritima, which continued wherever we went, even to Southampton, where it had completely displaced huroniana, the latter species not having been met with anywhere. Unfortunately it was necessary for us to return to Toronto, so it was not determined whether maritima had colonized all the beach areas of the Bruce Peninsula or not. I regard it as highly probable that this will prove to be the case. Here again, at any rate, is another instance of a southern species pushing northward and a more northern species retreating and, in this instance, being apparently ·directly displaced by it. These observations on Orthoptera are all casual ones, since my faunistic studies on this Order were discontinued many years ago in favour of the Odonata. It may therefore· be asked why I did not choose the latter Order instead of the Orthoptera to illustrate the subject of this paper. The reason is that dragonflies are great wanderers and their tremendous powers of flight often carry them great distances. I have known certain species to appear suddenly and iii numbers on a familiar stream whose odonate inhabitants were well known to me. I have known them to mate and oviposit, sometimes persist for a· few years, but finally die out again. The wandering propensities of dragonflies make them an unsuitable group for our present purpose. Yet I am satisfied that the same kind of change is taking place in this Order as in all the others. Naturally I would not venture to discuss this matter in detail with reference 10

to any of the other Orders of insects but, having been a general collector in my youth, I may perhaps be pardoned for mentioning some changes in the distribution of certain well known and easily recognized species of several Orders. Among the butterflies that have disappeared from the vicinity of DeGrassi Point about the tum of the century are Polygonia faunus Edw. and Argynnis atlantis Edw., both relatively northern species. The former was not uncommon along an unused corduroy road through the swamp at DeGrassi Point. As a young collector I valued much more the more southerly P. interrogationis Fab. because it was a rarity at that time and place. Now it is common and faunus has long departed. Argynnis atlantis was abundant in the same swamp and now it too has vanished. Another common butterfly at DeGrassi Point was Limenitis arthemis Dru., 1 sometimes called the White Admiral from the broad white band crossing the black wings. A variety of this butterfly without the white band or with only traces of it was formerly restricted, or almost restricted to the Eastern States, immediately south of the international boundary. But in the early thirties specimens of this dark form began to appear in the vicinity of Toronto and at Credit Forks, and in 1936 at DeGrassi Point. In a few years it was as common as the typical arthemis and has continued to be. Although the dark form is not a distinct species its northward spread appears to be comparable to that of the various species mentioned as having extended their range northward. The most spectacular case of northward range extension among butterflies, however, is that of the Buckeye (Junonia coenia Hbn.). Although reported from southern Ontario as an occasional stray in my early years of collecting, I had never observed it in any part of Canada until about ten years ago, when it began to be seen occasionally in the vicinity of Toronto. During the last four years I have seen it at DeGrassi Point and, in 1953, I noticed that it was already common there. As to other Orders of insects I have no doubt that there are plenty of examples to illustrate both the northward retreat of some species from our selected locality and the immigration of others generally from the south, these movements being, of course, all a part of the same process. One conspicuous species from each of the three largest Orders is all that I shall mention. Of Coleoptera the most conspicuous example that I have noticed is that of Pelidnota punctata L. This beetle was included by Dr. Saunders in his Insects Injurious to Fruits (1889) and was certainly known from the southernmost parts of Ontario in my early years of collecting, but I never saw a specimen until about the beginning of the nineteen-forties, long after I had ceased to look for beetles. First specimens began to appear about Toronto and almost immediately afterwards at DeGrassi Point. Of Hymenoptera the best example that has come to my notice is Sphex ichneumonea L. This is our largest digger-wasp and the orange and steel !Commonly known as Basilarchea arthemis. 11

blue body is so distinctive that it cannot be mistaken for any other species. It is seen in summer about barren gravelly or sandy places, where the female digs her nest and provisions it with grasshoppers. It was not uncommon at High Park, Toronto, in the nineties but it was certainly not present at DeGrassi Point. During the last three or four years, however, I have often seen it along the same road where the newly arrived grasshoppers of the genus Spharagemon were first observed. I attach no significance to this fact except that the dry and now almost unused road makes it a favourable habitat for various insects that prefer gravelly sterile soils. Finally, I would select from the Diptera a large hairy robber-fly, Proctacanthus milberti Macq., a species that is found in the same biotype as Sphex ichneumonea and has had a similar history as far as DeGrassi Point is concerned. I remember it on the sandy areas of High Park many years ago, although it was not very common there. At DeGrassi Point there are similar sandy and gravelly spots, although on a smaller scale, and I have spent many hours there through several decades, but until about fifteen years ago there were no Proctacanthus in this region. In the last few years, however, particularly in the hot, dry summer of 1955, they have been very common indeed. It is obvious that the material presented herewith is offered as evidence of climatic change in southern Ontario, although I am aware that the problem is not as simple as it appears to be. Many insects like plants are found at their southern limit in the coolest situations, namely, higher altitudes or swamps and bogs. In southern Ontario variations in altitude are not of much significance, although they do play a minor role in distribution. More important are the swamps and bogs, and in their undisturbed condition they support many northern plants far south of their main area of distribution. And with these northern plants are associated northern insects. But when the trees are cut away a rank jungle of willows and other shrubs is likely to replace the former growth and the northern insects as well as the northern plants soon disappear. The lowering of the water table, which has proceeded steadily with the clearing of the land, is another factor that affects the flora and fauna of swamps and may thus be instrumental in the destruction of boreal plants and animals. But when we make due allowance for the effects of these factors there still remain the many inhabitants of drier areas, the grasshoppers of semiarid habitats, butterflies and others that are moving northward. They seem to offer evidence of a general rising of temperature, but we shall leave the discussion of this problem to the meteorologists.

12

CHANGES IN THE AMPHIBIAN AND REPTILIAN FAUNA OF ONTARIO BY

E. B. S. LOGIER

IN PRESENTING the case for the amphibians and reptiles, special difficulties are encountered for two reasons: There are no figures for numbers seen, caught, or killed, and the identification of forms reported may be open to question. The term "bullfrog" for example, in popular accounts, does not necessarily mean Rana catesbeiana, and even if the Latin name happens to be used, we still cannot always be sure that it was correctly applied. The same is true of such terms as, toad, tree frog, lizard ( often applied to salamanders), grass snake, rattlesnake, puff-adder (a name sometimes applied to the hognosed snake, but also extended occasionally to any snake that hisses when frightened), snapping turtle, mud turtle, etc. Apparent extensions of ranges, or the filling in of gaps within ranges, in recent years, are probably due to an increasing interest in natural history, the presence of more observers, and extended field-work. For example, Bleakney (1954, pp. 165-171) published upon range extensions in Canada of six species of salamanders and eight species of frogs, and these extensions vary from 50 to 600 miles beyond the previously known range limits. Certainly, nobody would contend that these slow-moving, earthbound creatures have actually extended their ranges by such distances in recent years. This is plainly a case of more knowledge of conditions that have existed for a long time. In the case of the wood turtle (Clemmys insculpta) Ure (1858, p. 58) reported it from Toronto, York County, Ontario. Provancher (1874, p. 296) reported it from St. Gertrude, Nicolet County, Quebec. Williamson (1907, p. 138) reported it from Searchmont, Algoma District, Ontario. A New Brunswick specimen without definite locality came into our possession in 1925. In the years from 1933 to 1955 the former gaps have been filling in and the eastward range has been extended. There are records from four Nova Scotia counties, Cumberland, Halifax, Hants, and Kings (Bleakney, 1952, p. 127); from three more Quebec counties, Jacques Cartier, Joliette, and Sherbrooke; and from four additional Ontario counties, Carleton, Halton, Huron, and Renfrew; besides further specimens from counties previously cited. Again we must concede that this animal had a wide and more or less continuous distribution throughout the vast area covered by these records, but had been overlooked. Several species reported by Ure in 1858 from the Toronto region have not, to my knowledge, been found there since; a number of them certainly did not occur there, so it is difficult to assess on his authority the probability of those that might have occurred. Among the latter may be mentioned: the soft-shelled turtle (Trionyx ferox [spinifera]); the spotted turtle (Emys 13

[= Clemmys] guttata) (we assume that a single specimen caught in a stream in Lawrence Park within the city in 1933 was a liberated or escaped pet); the map turtle (Emys [= Graptemys] geographica); the musk turtle (Sternotherus odoratus) ; and a black snake reported as Coluber constrictor but possibly referring to Elaphe obsoleta. From the Natural History of the Toronto Region published by the Canadian Institute in 1913 these species were omitted. A glance at the present range maps will show that the musk, map, and soft-shelled turtles miss Toronto by a distance of about 40 miles at their nearest approach westward at Hamilton, and the blue-tailed skink (Eumeces f asciatus) by about an equal distance southward at St. Catharines ( or by about 70 miles around the lake shore). They all occur again east of Toronto, the nearest points being: the musk turtle in Peterborough County, about 70 or more miles away; the map turtle at Rice Lake in Northumberland County, about 60 miles away; the soft-shelled turtle not nearer than Ottawa, more than 200 miles away; and the skink in Peterborough County, about 70 or more miles away. The spotted turtle does not, so far as we know, occur east of Toronto in Canada (although reported by Provancher, 1874, p. 295, from Nicolet and Quebec counties in Quebec). The "black snake" cited as Coluber constrictor is represented in Ontario by the race flaviventris, which is not black, but bluish or greenish, and is known only from Essex and Huron counties. The pilot blacksnake (Elaphe obsoleta) on the other hand, occurs along Lake Erie and extends into Lincoln County, and occurs again in Frontenac and Leeds counties. It was reported from a locality probably between Rice Lake and Port Hope by Fothergill in his notes of 1816 to 1837 (Black, 1934, p. 161). The ranges of the musk turtle and skink appear to be continuous north of Toronto; those of the map turtle and pilot blacksnake discontinuous. We have then an area of about 40 X 100 miles unpopulated by any single one of these species, though near the center of their ranges in Canada. Until 1939 Blanding's turtle (Emys blandingi) showed a similar pattern, apparently missing Toronto by about 100 miles in any direction. Since then it has been found at Ashbridge's Bay, Armour Heights, Mount Dennis, the Humber marsh, and Grenadier Pond in Toronto; also at Highland Creek east of Toronto and Mimico Creek west of Toronto. At the first, second and fourth Toronto localities named above, two specimens were found in each case. Has it been introduced in recent years? In the writings of early travellers in Canada there are many references to rattlesnakes, but the identity of the species is not always clear and one is left to infer it from other details in the narratives. The two species found in eastern Canada, the massasauga (Sistrurus catenatus) and the timber rattlesnake ( Crotalus horridus) are restricted to southern Ontario. From the records and legends available, it appears that these snakes were more numerous and widely distributed in former years than they are today. The earliest identifiable reference of which we have knowledge was to 14

the timber rattlesnake, and appeared in the journal of Rene de Brehart Galinee for September, 1669. Galinee was a member of M. de La Salle's party and described the latter's encounter with three large rattlesnakes on a rocky eminence near a small lake, now called Lake Medad, just inside Halton County and not far from Waterdown in Wentworth County. The description given of the timber rattlesnake is unmistakeable. The facts were obtained from the translation by B. E. Charlton (1884,. pp. 81-82). Robert C. Gourlay ( 1822, pp. 185-186) recognized both the massasauga and the timber rattlesnake, and wrote, "There are two species of rattlesnakes, vulgarly distinguished by the names yellow, or large, and black, or small, rattlesnake. The former is from four to five feet in length, and the middle of the body seven or eight inches in circumference from whence it tapers toward the head and tail. The neck is small, head flattened, . . ." According to Gourlay, the small rattlesnake was the commoner. He cites no localities where specimens were seen or collected. In the Royal Ontario Museum collection there is a timber rattlesnake labelled as from Fitzwilliam Island, August, 1886, but with no other data. The species is still found sparingly in the Niagara Gorge in the vicinity of the Glen, so that it seems likely from the scanty evidence extant, that its range in Ontario was once coextensive with the limestone outcropping northward to Manitoulin District. If we assume, as I think we may, that a specimen killed at Point Pelee in September, 1918 was not indigenous, all Canadian records of this species known to us to be supported by specimens, since the Fitzwilliam Island record of 1886, were from the vicinity of the Niagara Gorge. The massasauga is still found in southern Ontario from Welland County and Hamilton westward along Lake Erie, northward to the tip of Bruce Peninsula, and around the coastal region of Georgian Bay to Manitoulin District. Eastward it reaches the Muskoka Lakes. Judging by reports of early writers it was more abundant and its range more extensive formerly than is now the case. Mrs. John Graves Simcoe, in her diary for the years 1792 to 1797, tells of rattlesnakes being numerous during the cutting of the forest for the opening of Dundas Street. For September 24, 1793 she had an entry, "I hear that they kill rattlesnakes every day, yet not a man has been bitten although they have been among them for six weeks." For December, 1795 she had an entry, "Mr. Jones, the Surveyor, says seven hundred rattlesnakes were killed near Burlington Bay this summer. They live in caves, and in very dry weather go down to the lake to drink; they are sluggish, and as they move in numbers at a time, probably would be easier destroyed than many other reptiles. The man is quite recovered who was bitten by one last August." The above diary notes possibly refer to both species, but would seem to relate mostly to the massasauga, if we may judge by the infrequency of bites, and the apparent lack of fatal accidents with men working where the snakes were numerous. 15

In Charles Fothergill's notes on the natural history of eastern Canada, 1816 to 1837, p. 273 (Black, loc. cit.), we find the following remarks about "the rattlesnake," species not named, "It is common in the Midland District and has been frequently seen and killed in the Home District--east and west of the District of Newcastle--yet in the latter district, notwithstanding there is every variety of ground, I cannot hear of a single instance where it has been killed." According to Smith's Canadian Gazetteer, 1846, the Midland District consisted of the counties of Frontenac, Lennox and Addington; the Home District of York County; and the Newcastle District of Northumberland and Durham counties. We believe these references (if they are really to rattlesnakes) are to the massasauga rather than to the timber rattlesnake, and a glance at the maps of their present ranges in eastern North America supports this view. It is apparent that the rattlesnake referred to occupied an extensive territory north of Lake Ontario through York County and eastward to Frontenac County, being absent apparently from the counties of Northumberland and Durham. Fothergill's statement lends weight to reports of its former occurrence in Prince Edward County, which, if valid, indicate that it was abundant there about 150 years ago, and was occasionally reported as killed there until 1929, when two were said to have been killed at Demorestville. What are we to make of all this? In the case of those species which leave a "vacant lot" so widely surrounding Toronto, the mere fact that they are so distributed today tells us nothing about faunal changes. Today, the hog-nosed snake (Heterodon platyrhinos) shows a similar pattern, but it did occur at Toronto, at least until 1907, when the last specimen (of which we have knowledge) was collected at High Park. This snake is especially poorly adapted to survive in settled areas for two reasons: Its habit of bluffing and threatening when confronted by man, coupled with its slow gait when seeking cover, make it a ready target for sudden death; while its restricted feeding habits would ensure its disappearance from areas rendered unsuitable for the habitation of toads or frogs. These factors would readily account for its disappearance from Toronto, but would hardly explain its apparent absence from a forty-mile radius around Toronto: we suspect that it is present in that area but that those seen or killed have not been collected or reported. For the other species, the musk turtle, spotted turtle, map turtle, and soft-shelled turtle, and the blue-tailed skink:, we know of no convincing evidence that they ever occurred in the area, while neither climate, geology, nor degree of settlement would seem to account for their absence, and thus we seem to come to a dead end. It is to be expected that rattlesnakes would be singled out for especial destruction in settled areas, and this fact, without doubt, had much to do with reducing the Ontario populations of these snakes, but does not seem to be the only factor. Why, for instance, has the massasauga disappeared within a century from such a large area of its former range when much of that area is still wilder and less settled, and certainly not more settled, than much of 16

that in which it still survives? It is now mostly restricted, so far as we know, to a belt of not much more than twenty miles wide around lakes Erie and Huron and Georgian Bay, touching Lake Ontario at Hamilton. The farthest inland report of the current century of which we have knowledge is that recorded by Dr. W. S. Fox (1951, p. 39) at Mount Forest, Wellington County, 42 miles inland from Lake Huron. The next farthest in were at Gravenhurst, Muskoka District, and Newbury, Middlesex County, (the latter two were established by specimens collected). Contrast this with its former apparent range east to Frontenac County and probably including Prince Edward County. Could any of Fothergill's reports of "rattlesnakes" be based upon misidentifications of other snakes? The timber rattlesnake appears to have been even less successful in holding its own. The fact that it is a larger and less timid snake would not be to its advantage in this case. However, parts of its former range ( apart from the Niagara Gorge) appear to be still sufficiently rugged and unsettled to afford sanctuary for at least a skeleton population to maintain itself for a while longer; nevertheless, it has apparently failed. The populations in question are in the northern extremities of the ranges of their species and would thus be the first to be affected by a climatic change toward colder, but there has been no such change, the trend being rather toward warmer, and well within the tolerance of these animals, which range far to the south. While human intervention has without doubt been a significant factor locally in some regions, and is ever becoming more so, it does not appear to be the answer for what has apparently happened over most of the former ranges of these snakes, nor does it appear to explain the "vacant lot" around Toronto for four species of turtles and the skink. In view of the paucity of information, it was with tongue in cheek that I complied with the request to prepare this paper. Yet since it does bring together and put on record the few facts available from the past, along with a statement of the situation that exists at the present time, and such comparison as it is possible to make, it may serve a useful purpose in later years. If somebody competent to identify the animals ( or infer probable identity from facts that might have bearing on it) had prepared a paper of this nature 50 or 60 years ago, I would be very glad to see it today, and some of our questions might be answered. The answers, indeed, may exist in scattered notebooks and old letters that have never come to the attention of zoologists, or the light of publication. LITERATURE CITED

R. D. 1934. Charles Fothergill's notes on the natural history of eastern Canada, 1816-1837. Trans. R. Canad. Inst., vol. 20, pt. 1, pp. 141-168. BLEAKNEY, S. 1952. The amphibians and reptiles of Nova Scotia. Canad. Fld.-Nat., vol. 66,no. 5,pp. 125-129.

BLACK,

17

1954. Range extensions of amphibians in eastern Canada. Canad. Fld.-Nat., vol. 68, no. 4, pp. 165-171. CHARLTON' B. E. 1884. The discovery of Burlington Bay. J.Proc. Hamilton Assoc., 18821884, vol. 1, pp. 41-57. Fox, W.S. 1951. The Little Swamp Rattlesnake in inland Ontario. Canad. Fld.-Nat., vol. 65, no. 1, p. 39. GOURLAY, R. C. 1822. Statistical account of Upper Canada. Simpkins and Marshall, London, vol. 1, pp. i-xxiv, 1-625. PROVANCHER, L. 1874. Faune Canadien, les reptiles. Nat. Canad., vol. 6, no. 10, pp. 289-298. URE, G. P. 1858. Handbook of Toronto. Lovell and Gibson, Toronto, pp. i-viii, 9-272. WILLIAMSON, E. B. 1907. A collecting trip to north of Sault Ste. Marie, Ontario. Ohio Nat., vol. 7, pp. 129-148.

18

CHANGES IN THE FISH FAUNA OF ONTARIO BY

W.

B. SCOTT

IN 1944, Rad.forth published an account of the distribution of Ontario fishes in a paper entitled "Some considerations on the distribution of fishes in Ontario." However, many fish distributional changes have occurred since the publication of Radforth's paper, and therefore the primary purpose of the present paper is to record these changes. BRIEF GEOLOGIC HISTORY

The indigenous fishes occurring in Ontario at the present time are generally considered to have moved into the area from at least two regions, perhaps three, in North America. These three regions, Atlantic coastal plain, Mississippi Valley, and the Alaska-Yukon area, were relatively unaffected by glaciation. There is considerable doubt concerning the contribution made by the Alaska-Yukon refugium to the fish fauna of Ontario (Walters 1955). Following the last glacial retreat, the fishes which found refuge in these unglaciated regions reinvaded the ice-free land by means of the available drainage systems (Rad.forth, 1944). After the retreat of the last ice-sheet an arm of the Atlantic Ocean extended up the Ottawa Valley and into the basin of Lake Ontario. These marine waters eventually withdrew. Evidence of their presence is available in the form of marine fish species which were left isolated in some of our inland waters. (See Rad.forth, 1944, for a more complete account of the geologic history of Ontario's drainage systems.) EXTENSION OF RANGE DUE TO REMOVAL OF NATURAL BARRIERS

Fishes which moved into Lake Ontario from the Atlantic coastal area were prevented from further westward movement by the Niagara escarpment, notably Niagara Falls. Such species as the sea lamprey (Petromywn marinus), the Atlantic salmon (Salmo salar), the alewife (Pomolobus pseudoharengus) 1 and the threespine stickleback ( Gasterosteus aculeatus) were apparently indigenous to Lake Ontario or at least have occurred there for many decades. The completion of the Welland Canal in 1824, linking Lake Ontario with Lake Erie, opened the door to the upper Great Lakes, thus allowing fishes to move northwestward into waters new to them. However, the movement of fishes into Lake Erie appears to have been very slow indeed. 1The origin of the alewife is open to some doubt. It has been abundant in Lake Ontario since at least 1890 and was known to occur in that lake many years prior to that date.

19

SEA LAMPREY

Consider, for example, the sea lamprey. The first specimen from Lake Erie was recorded in 1921 (Dymond 1922). The subsequent spread of the lamprey through the upper Great Lakes, including Lake Superior, is a phenomenon of re.cent occurrence. The path of destruction left by the sea lamprey has been amply recorded in the literature (Applegate, 1950; Trautman, 1949). Considering that the Welland Canal was in existence for almost 100 years before evidence of its use by the lamprey to gain access to the upper lakes was obtained, the movement through the canal and establishment in the upper lakes was surprisingly slow. ALEWIFE

The westward movement of the alewife, which has been abundant in Lake Ontario since at least 1890, appears to have been as slow as that of the sea lamprey. Dymond (1937) recorded the first known specimens from Lake Erie in 1931, while MacKay (1934) noted that the first specimens were taken in Lake Huron in 1933. The alewife is currently abundant in Lake Erie and rapidly becoming more numerous in Lake Huron. In 1955 it was reported to be the most abundant species in the experimental fishery operated in South Bay, Manitoulin Island. A better appreciation of the rate of increase of the alewife in Lake Huron may be gained from the following production figures for the experimental fishery operated at South Bay by the Ontario Department of Lands and Forests: 19 511 specimen ( 4 oz.) 19525 specimens 1953-53 specimens (6 pds.) 1954- 2,316 pounds 1955-16,302 pounds It would appear to be only a matter of time until the alewife becomes established in Lake Superior. It has been suggested by Van Oosten (1935), Radforth (1944), and others, that the alewife, sea lamprey and the American eel (Anguilla rostrata) gained access to Georgian Bay and Lake Huron via the Trent Valley Canal system. This means of access necessitates passage through Lake Simcoe, from which lake none of these species has ever been reported. There is no direct evidence to support the theory that these fishes have used this waterway to gain access to the upper Great Lakes, and in view of the recently constructed power dams and other obstacles on the northern portion of the system, it seems unlikely that it will be used as a means of access in the future. (Morone americana) A species new to the Ontario fauna has appeared in Lake Ontario in the past few years, having gained access via New York State. This is the white

WHITE PERCH

20

perch which now occurs along the north shore of Lake Ontario westward at least to the mouth of Highland Creek, York County, 10-12 miles east of Toronto ( Scott and Christie, ms.). This species may have gained access to Lake Ontario via the New York State Bridge Canal (Greeley, pers. comm., and Dence, 1952). Specimens in the collections of the Royal Ontario Museum indicate a range from Lennox and Addington County westward to at least York County. The first Ontario record is a specimen taken in 1952 off Prince Edward County. The white perch appears to be established and to be increasing in numbers along the north shore of Lake Ontario. This species was recorded from the Pennsylvania waters of Lake Erie by Larsen (1954) but only three specimens were taken and no further reports of its occurrence in Lake Erie waters have appeared. EXTENSION OF ORIGINAL RANGE OF NATIVE FISHES BY PLANTING AMERICAN SMELT ( Osmerus

mordax)

This fish is indigenous to several lakes in the Ottawa Valley ( Radforth, 1944; Dymond, 1944) , but its occurrence in all of the Great Lakes is a result of planting. The subsequent invasion of the Great Lakes following the Michigan plantings in the early 1900's has been recorded by Dymond (1944) and Van Oosten (1940). Although the species had become common in lakes Michigan and Huron in the early 1930's, they were only occasionally caught off Lake Erie's north shore even as late as 1946. Since that time, however, there has been a steady increase in the smelt population of Lake Erie. The magnitude of the increase of this species can best be appreciated by the catches of the Ontario commercial fishery in Lake Erie which landed over one million pounds in 1953, one and a quarter million pounds in 1954, and over two million pounds in 1955. ATLANTIC SALMON

(Salmo salar)

Although the Atlantic Salmon no longer occurs in Lake Ontario, it was an indigenous species until its final extinction about 1890-1900 (Huntsman, 1944). Experimental attempts to re-establish the species in Lake Ontario have been unsuccessful (Mccrimmon, 1950). The species still exists in Ontario as a result of plantings in Trout Lake, Nipissing District, where the population appears to be maintaining itself by natural reproduction. Attempts to establish the species in many other Ontario lakes have been unsuccessful. BROOK TROUT

(Salvelinus fontinalis)

Because this fish is a desirable game species, it now occurs, due to plantings, in many Ontario waters. For example, its original distribution did not include the southern portion of western Ontario, but it occurs in that region now as a result of planted stock. 21

OTHER GAME SPECIES Many other game species such as smallmouth bass (Micropterus dolomieui) and yellow walleye or yellow pickerel (Stizostedion vitreum) now occur, as a result of plantings, in Ontario waters to which they are not native. Such plantings are usually attended by misgivings and should not be done without careful consideration. GIZZARD SHAD (Dorosoma cepedianum) Although it is not definitely known whether the gizzard shad is native to Lake Erie or was introduced, it is now abundant there and has been for many years. Available evidence suggests that it was introduced into Lake Erie waters. It has subsequently spread to Lake Huron and Georgian Bay. As yet there are no records of its occurrence in Lake Superior. NORTHWARD DISPERSAL OF SOUTHERN FISHES

BY

NATURAL MEANS

The western lake chubsucker (Erimyzon sucetta kennerleyi) is native to the Ohio and Michigan drainage system of Lake Erie. In 1949 this species was found to occur in ponds on Point Pelee and subsequently in ponds on Long Point, Lake Erie, and in Lake St. Clair (Scott, 1952). This was apparently the first record of this species in Canadian waters. By 1955 the species had definitely increased and 27 adult specimens were captured in Rondeau Bay, Lake Erie, in hoop nets operated from the 13th to 26th of April under the direction of Mr. R. G. Ferguson, Ontario Department of Lands and Forests. This is the only record in recent years, to my knowledge, of a southern species of fish becoming established in Ontario waters by entirely natural means. 2 This relatively southern species apparently finds conditions suitable for existence in Canadian waters of Lake Erie. EXOTIC INTRODUCTIONS PACIFIC SALMON (genus Oncorhynchus) The Chinook Salmon ( Oncorhynchus tshawytscha) and the Coho Salmon (0. kisutch) were introduced into Lake Ontario (Dymond, Hart and Pritchard, 1929) and Lake Erie, respectively, but a permanent population was not established in either case. In the winter of 1955 approximately one million eggs of the Chum Salmon ( 0. keta) were introduced into the waters of the Winisk and Attawapiskat rivers of the Hudson Bay and James Bay drainage systems. In 1956 more than one half million eggs and fry of the pink salmon ( 0. gorbuscha) were intro2As this publication was going to press another species, new to Ontario's fauna, was recorded. On April 30th, 1957, an 18-pound 2-ounce specimen of Megastomatobus cyprinella, the bigmouth buffalo, ( Catostomidae), was caught by commercial fishermen in Long Point Bay, Lake Erie, and turned over to the Royal Ontario Museum. 22

duced into waters flowing into Hudson Bay. The results of these plantings are not yet known. RAINBOW TROUT (Salmo gairdnerii) The introduction into Ontario waters of this Pacific coast species has been covered by Radforth (1944) and others. The major lakes in which it now occurs and appears well established have been recorded by Scott ( 1954). BROWN TROUT (Salmo trutta) The brown trout is currently established in many lakes and rivers of Ontario (Scott, 1954). For further details of its introduction from European waters see Radforth (1944) and Dymond (1955). FRENCH ALPINE CHAR (Salvelinus alpinus) This species has been introduced into some Ontario waters by the Ontario Department of Lands and Forests. The plantings commenced in 1954. HYBRID TROUT (Wendigo Trout or Splake) Salvelinus fontinali.1 X S. namaycush The re-development of this hybrid trout by fish culturists has resulted in plantings in numerous Ontario waters by the Ontario Department of Lands and Forests ( Scott, 19 5 6) . Plantings commenced in 19 5 3. There is every reason to believe that at least some of the introductions will be successful. CARP ( Cyprinus carpio) Introduced into Ontario waters about 1880 from Asia, via Europe, the Carp is now established in all of the Great Lakes and many inland waters (Scott, 1954). Although a successful introduction, it is regarded as an obnoxious species, and it will undoubtedly spread into many waters in Ontario where it does not yet occur. For further details of introduction see Radforth (1944). GOLDFISH ( Carassius auratus) Although not a planned introduction, the goldfish occurs in many waters in southern Ontario, principally in the Lake Erie District, as a result of release or escapement of domestic stock. EXTENSIONS OF RANGE DUE TO INCREASED KNOWLEDGE Considerable biological activity by biologists of the Ontario Department of Lands and Forests and other agencies has, in recent years, extended the known northward ( and southward) distribution of many fish species. These range extensions appear to be due solely to increased knowledge rather than to movement by the species. For example, the research activities concerning sea lamprey control currently being carried on in the Lake Superior drainage 23

have uncovered many species not previously known to be there. The northern brook lamprey (lchthyomyzon fossor), previously known to occur northward in Ontario only to Grey County, was found in the Chippewa River flowing into Lake Superior in 1954 and 1955. The American brook lamprey (Entosphenus lamottenii) was found (1953) to be the common lamprey in streams flowing into Lake Superior from the north shore, although previously known to occur in Ontario only in drainage of lakes Ontario, Erie, and St. Clair. Prosopium coulteri, the pigmy whitefish, formerly known only from western North America, was found to be fairly common in the deeper waters of Lake Superior (Eschmeyer and Bailey, 1954). This species was caught in a bottom trawl, a type of gear not previously used extensively on the Great Lakes. The first specimen of the goldeye (Amphiodon alosoides) in the St. Lawrence drainage system was reported from Lake Temiskaming, which drains into the Ottawa River. Specimens from the commercial catch were sent in to the Royal Ontario Museum in 1954 by Ontario Department of Lands and Fqrests biologist, Mr. R. Whitfield. The commercially important goldeye was previously known to occur only in the Arctic, Hudson Bay, and James Bay drainages of North America. LITERATURE CITED

v. C. 1950. Natural history of the sea lamprey (Petromywn marinus) in Michigan. U. S. Dep. Int., Fish Wild!. Serv., Spec. Sci. Rpt. no. 55, pp. 1-237. DENCE, W.A. 1952. Establishment of the white perch, Morone americana, in central New York. Copeia, 1952, no. 3, pp. 200-201. DYMOND, J. R. 1922. A provisional list of the fishes of Lake Erie. Univ. Toronto Stud., biol., no. 20, Pub. Ont. Fish. Res. Lab. no. 4, pp. 57-73. 1937. New records of Ontario fishes. Copeia, 1937, no. 1, p. 59. 1944. Spread of the smelt (Osmerus mordax) in the Canadian waters of the Great Lakes. Canad. Fld. Nat., vol. 58, no. 1, pp. 12-14. 1955. The introduction of foreign fishes in Canada. Proc. int. Assoc. Theoret. Applied Limnol., vol. 12, pp. 543-553. DYMOND, J. R., J. L. HART, AND A. L. PRITCHARD 1929. The fishes of the Canadian waters of Lake Ontario. Univ. Toronto Stud. biol. no. 33, Pub. Ont. Fish. Res. Lab., no. 37, 35 pp. EsCHMEYER, PAUL H., AND REEVE M. BAILEY · 1954. The pigmy whitefish, Poragonus coulteri, in Lake Superior. Trans. Amer. Fish. Soc., vol. 84, pp. 161-199. GREELEY, J. R. 1955. Personal communication.

APPLEGATE,

HUNTSMAN, A.G.

1944. Why did Ontario salmon disappear. Trans. R. Soc. Can. Ser. 5, vol. 38, pp. 83-102.

24

A. 1954. First record of the white perch (Marone americana) in Lake Erie. Copeia, 1954, no. 3, p. 154. MACKAY, H. H. 1934. Record of the alewife from Lake Huron. Copeia, 1934, no. 2, p. 97.

LARSEN,

MCCRIMMON'

H. R.

1950. The reintroduction of Atlantic Salmon into tributary streams of Lake Ontario. Trans. Amer. Fish. Soc., vol. 78, pp. 128-132.

RADFORTH, ISOBEL

1944. Some considerations on the distribution of fishes in Ontario. Contr. R. Ont. Mus. Zool., no. 25, pp. 1-116. SCOTT,W.B.

1952. Records of the western lake ch.ubsucker, Erimyzon sucetta kennerleyi, from Ontario, Canada. Copeia, 1952, no. 3, p. 203. 1954. The freshwater fishes of eastern Canada. Univ. Toronto Press, pp. 1-128. 1956. Wendigo-the hybrid trout. R. Ont. Mus., Div. Zool. Palaeont., pp. 1-7. SCOTT, W. B., AND J. CHRISTIE 1957. The white perch (Marone americana) in Ontario. (MS).

TRAUTMAN, M. B.

1949. The invasion, present status, and life history of the sea lamprey in the waters of the Great Lakes, especially the Ohio waters of Lake Erie. Contr. Franz Theodore Stone Lab., mimeo., 7 pp. VANOOSTEN, J. 1935. First record of the alewife (Pomolobus pseudoharengus) for the state of Michigan. Copeia 4, 1935, p. 194. 1940. The smelt (Osmerus mordax) (Mitchill). U. S. Bur. Fish., mimeo., pp. 1-13. WALTERS, V.

1955. Fishes of western arctic America and eastern arctic Siberia. Bull. Amer. Mus. Nat. Hist., vol. 106, art. 5, pp. 259-368.

25

CHANGES IN THE AVIFAUNA OF ONTARIO BY

L. L. SNYDER

CHANGES, as here used, refers particularly to alterations in range patterns of Ontario birds. Some consideration of numerical change is naturally involved, but quantitative data are lacking for early, and even our own, periods. With the passage of time, and through agencies of the environment, including man, there have been extirpations of indigenous forms, introductions of exotics, contractions and expansions of range patterns within, or into the province from without. The historic records of cases in no instance exceed 146 years, and the data are admittedly meagre. EXTIRPATIONS

The history of the now extinct Passenger Pigeon (Ectopistes migratorius) in Ontario, has been fully documented by Mitchell (1935). A brief digest of this major and permanent faunal disturbance is as follows: The species once occurred in prodigious numbers in southern Ontario in summer, and scattered beyond. Reports of decline in numbers commenced toward the middle of the 19th century (Bmmycastle, 1841), and extinction came by the opening of the 20th century. The last specimen record, for Sherkston near Niagara, has been fixed as mid-September, 1891. The last unquestioned sight record concerned two birds observed on May 18, 1902, at Penetanguishene. The Turkey (Meleagris gallopavo) once occurred plentifully in the hardwood forests of southern Ontario, south from and including Lambton, Middlesex, Oxford, Brant, and Wentworth counties and with a northward extension to Simcoe County (Devitt, 1943) and east to Durham County (Orr, 1909). Comment on reduction in numbers is found in the literature during the first quarter of the 19th century, and though final extirpation cannot be dated, it came near the close of that century (Clarke, 1948). Reintroductions within the former range are too recent to indicate success or failure. RANGE SHRINKAGES AND FLUCTUATIONS The range boundaries of many Ontario birds have been greatly modified in detail of pattern during the period of human settlement. A once continuous front is now interrupted or presents a moth-eaten pattern. The range of the Ruffed Grouse (Bonasa umbellus, involving B. u. monticola and to a lesser extent B. u. togata) has been so affected. It is reasonable to suppose 26

that such is true of scores of species of forest birds. To single out another probable case, we can mention the American Redstart (Setophaga ruticilla) the nominate race of which probably once occurred throughout our southern forest. Now its summer occurrence in southern counties is notably sporadic. Other species have undergone a more marked change in range by the total disappearance of representatives over extensive peripheral areas. A case in point is the Spruce Grouse ( Canachites canadensis, with particular reference to the race C. c. canace). Morden and Saunders (1882) report a specimen collected near Chatham by E.W. Sandys in the 1870's. Such outlying occurrences, of a non-migratory, non-emigratory species, would appear to be strictly relic. The species was extant in Wellington County in 1905 (Klugh, 1905), and a specimen exists (R.O.M.) which was taken in Caledon Township, Peel County, about 1908. The species has been extirpated from these stations, and the southern range limits have retreated to a line extending from Manitoulin Island to northern Frontenac County. Thus a shrinkage of range, approximating 200 miles, during the past 70 years or so is apparent. An example of range change which represents a fluctuation during historic times is presented by the history of the Bob-white ( Colinus virginianus, nominate race). Oarke (1954) points out that both its geographic range and its numbers were at a maximum between 1840 and 1850. With change from primeval to pioneering conditions, the species expanded its range northward from an indefinite centre in the Lake Erie region to a front extending from Bruce County through southwestern Muskoka District, thence southeastward to Peterborough, and eastward to Kingston. After 1885 the northern front had retreated beyond northern Perth and southern Ontario counties and by early in the present century it had shrunk to approximately its present-day range, probably little beyond its range of the early 19th century. Currently the species occurs from southern Huron County through Middlesex, Oxford, Brant, and Lincoln counties, with the most favourable status existing to the southwest of this line. A range shift of a somewhat different nature is indicated by the history of the Prairie Chicken (Tympanuchus cupido, race T. c. pinnatus). The species was present on the "plains of Essex" County during the period between 1824 and 1836, according to an entry in the Charles Fothergill manuscript (R.O.M., Oendenan volume). It also occurred in Kent County, persisting until 1897 (Saunders, 1910). Long after its extirpation in extreme southern Ontario an effective re-invasion took place in the northern part of southern Ontario, approximately 250 miles north of the early historic range salient. Reappearance in the province first occurred on St. Joseph's Island in the North Channel (Baillie, 1947) about 1925. Further spread to Manitoulin Island took place in 1938 or 1939 where it remains to date, but where hybridization with the Sharp-tailed Grouse (Pedioecetes phasianellus; race uncertain) may present a challenge to its continued existence there. The range of the Sandhill Crane ( Grus canadensis) originally composed of two salients in Ontario, south and north, has been altered in one section 27

and apparently not in the other. The species (represented by the race G. c. tabida) occurred regularly in the marshes of Lake St. Clair as early as 1862 (Mcilwraith, 1886), and there were scattered records of occurrence elsewhere in the southern counties (Walkinshaw, 1949). By about 1888 this inhabitant of open wilderness situations was extirpated in this portion of its range ( Saunders, 18 88). However, it has been present ( race uncertain) in far northern Ontario from the middle of the 19th century (Murray, 1859) to date ( Baillie, 19 51) . There certainly is no evidence or rational basis for supposing that the southern population of this species has withdrawn northward. The case is clearly that of regional extirpation of a species. Probably its numbers are fewer in the north than during primeval times. INTRODUCED EXOTICS

The four cases of exotic species of birds which have become successfully established in Ontario, either from direct introductions or consequent to spread from importations elsewhere in the New World, all constitute important biological changes in our avifauna. The European Partridge (Perdix perdix, presumably the nominate race) was first introduced into southern Ontario shortly before 1907 (Snyder, 1951), and many subsequent introductions have been made in southern, central, and western parts of the province. Currently the species has a scattered population across southern Ontario from Essex to Carleton counties, with the greatest success being evident in eastern areas. It has also persisted in the central east-the regions of North Bay, Timmins, New Liskeard, and Cochrane-and in the west in the vicinity of Fort William and Port Arthur (R.O.M. MSS). The Common Pheasant (Phasianus colchicus) was first introduced in southern Ontario at some time shortly prior to 1892. Liberations were subsequently made on a large scale ( mixed racial stock) and in various parts of the province. It is now apparent that the best pheasant range is in the counties of Essex, Lambton, and Kent, and areas adjacent to the Niagara River and the west shore of Lake Ontario, with optimum conditions on Pelee Island. However, the species has fared fairly well across all counties bordering Lake Erie and to a limited extent inland and along the north shore of Lake Ontario. Marginal limits extend northward to southwestern Huron, northern Middlesex, Oxford, Waterloo, Peel, York, southern Ontario, and Durham counties, with Prince Edward and Carleton counties somewhat isolated as moderately adequate range ( Clarke and Braffette, 1947). The Starling (Sturnus vulgaris, nominate race) spread into Ontario from introduction made in the New World at New York City in 1890. It was first observed in Ontario at Niagara Falls in the autumn of 1914 (Sheppard, et al., 1936). Within 15 years it had become one of the most conspicuously plentiful birds in southern Ontario. Currently it occupies the whole of settled Ontario and occurs sparingly in summer north to the most remote outposts, which probably have to be re-occupied annually. The establishment of the Starling 28

in Ontario is a major biological change affecting the economy of many indigenous forms of wildlife and man. The House Sparrow (Passer domesticus, nominate race) spread into Ontario from liberations in the New World at Brooklyn, New York, in 1851 and 1853 (Barrows, 1889) and subsequent introductions elsewhere in the northeastern United States. Spread into the province took place during the period between 1870 and 1875. Though the species does not now occur in numbers equal to those of the horse and buggy days, it is established throughout southern Ontario and occurs sparingly in the most remote settlements of the north, probably through repeated re-invasion. RANGE EXPANSIONS

In addition to the avifaunal changes grouped in the foregoing paragraphs, a number of range pattern expansions have taken place which involve many non-game, and therefore less familiar, species. These modifications can be meaningfully grouped according to the direction of spread. On this basis it is pertinent and significant to remark that there is no clear evidence that any species of bird is progressively extending its breeding range southward within or into the province. Although the range of the Red-throated Loon ( Gavia stellata) in graphic pattern may appear to indicate a southward extension, the proposition is here rejected. The range of this species in the New World is essentially Arctic and sub-Arctic, reaching northern British Columbia on the Pacific and Newfoundland on the Atlantic. A breeding station on an island five miles east of Thunder Cape on Lake Superior (Baillie and Harrington, 1936-37 and Dear, 1940) is far south of its continuous range boundary in the continental interior. There has been no expansion from this station. This station is more logically regarded as residual-the southern front of range having long since withdrawn to the north-rather than an initial outpost indicating a southward trend. Two other cases of re-dispersal may appear to be instances reflecting southward range extension in the province, namely, the Pileated Woodpecker (Dryocopus pileatus, race D. p. abieticola) and the Raven (Corvus corax, race C. c. principalis). But both of these represent cases of re-occupation of former range following local extirpation. Both persisted to the north and south of the areas now being re-occupied. Instances of eastern species expanding their ranges westward in the province seem limited to two species. Though in initial stages, both apparently show a trend. The Black Duck (Anas rubripes), a species characteristic of the east, has been recorded by Roberts (1932) with some quantitative evidence as having become established as a breeding bird in Minnesota since 1910, and is expanding its range there. A similar trend is indicated for western Ontario and Manitoba (Snyder, 1953), change having taken place during the past three or four decades. 29

The Black-backed Gull (Larus marinus) has long been known to frequent Lake Ontario (Mcllwraith, 1886) in the non-breeding season. In 1927 summer occurrences were noted (Snyder, 1941) on eastern Lake Ontario. With an increase of non-breeding season occurrences west to lakes Erie and Huron there was a breeding record for northern Lake Huron in 1953 (Baillie, 1954). Evidence supporting the occurrence of eastward extensions of range of western species is considerably more plentiful and definite. In addition to certain well-known and probably ancient range salients ( Schmidt, 1938) extending from the plains into extreme southern Ontario, illustrated by the Prairie Chicken and Sandhill Crane mentioned earlier in this paper, there are others of recent origin. Holboell's Grebe (Colymbus grisegena, race C. g. holbollii) is essentially of western distribution in North America (Snyder, 1954), its eastern breeding limits in the south in 1931 being in Minnesota, according to the 4th edition of the Check-list of North American Birds. In 1938 the species was found established as a breeding bird in Wisconsin (Jones, 1938). Since 1931 it has been observed in southern Ontario in summer, and in 1942 a small colony was found nesting at Burlington at the west end of Lake Ontario (Speirs, et al., 1944). This breeding station has persisted. In western sections of northern Ontario the species probably has been long established as a breeding bird. However, field work in the east had not disclosed its presence there in summer until 1955 when a colony was found established and nesting near Cochrane (Baillie, 1955c). These records constitute evidence of a recent eastward expansion of range. The Ring-necked Duck (Aythya collaris) is another species which has expanded its range eastward during the 20th century. Actually its breeding range was poorly understood prior to 1910, except that it was largely in the western Canadian provinces. It was known to breed commonly in Minnesota during the last quarter of the 19th century, but subsequently it almost disappeared there (Roberts, 1932). It may have been a breeding bird of extreme western Ontario at that time, but the first breeding evidence was not established until July, 1919, at Lac Seul (Bent, 1923). The evidence indicates that it has expanded eastward since that time at a phenomenal rate, spreading over the province in a pattern that coincides closely with the Precambrian Shield, and has reached the Atlantic coast region (Mendall, 1938). The Horned Lark (Eremophila alpestris, with particular reference to the prairie race E. a. praticola) apparently moved eastward nearly a century ago. Forbush (1927) has produced evidence that the prairie population of this species reached New England about 1860. However, the 1886 edition of the Check-list of North American Birds ascribes the race to the upper Mississippi Valley with a salient eastward, south of the Great Lakes, for an indefinite distance. It may have occupied the shores, or the sparsely wooded parklands, of extreme southern Ontario in pre-settlement days. Mcllwraith (1886) gives its advent in Ontario as about 1868. There appears to be a sound basis for 30

the assumption that it has spread over southern Ontario since that time. Currently its range limits are bounded by a line extending from the Sault Ste. Marie region, Manitoulin Island, Sudbury, Lake Nipissing to Petawawa. Though range extension in Ontario may appear to be northward, from the continental viewpoint it has spread eastward, northward spread being a secondary effect from fanning out. An eastward extension of range of the Western Meadowlark (Sturnella neglecta, nominate race) was first noticed at Port Arthur in 1917. Since then, with progressive frequency, scattered occurrences have been documented east to Moosonee in. the north (R.O.M., specimen) and to Carleton County in the south (Baillie, 1955b). The species is now established as a regular nesting bird west of Lake Superior and in the Sault Ste. Marie region where it entered about 1921 or 1922 (Snyder, et al., 1942). It has been known to nest once to the south, near Hamilton (Baillie, 1955b). Though covering a more recent period, · a similar history pertains to Brewer's Blackbird (Euphagus cyanocephalus). It was first ascertained that a colony was established in the Port Arthur region in 1945 (Allin and Dear, 1947). Numbers and colonies have increased there in the past ten years (Baillie, 1955c). The species has continued its eastward trend south of the upper Great Lakes (White, 1935) and has entered and become established in the Sault Ste. Marie region of Ontario (Baillie, 1954). One of the most remarkable changes concerning eastward penetration of a western species concerns the Evening Grosbeak (Hesperiphona vespertina, nominate race) . Its invasion and spread has been traced by Baillie ( 1940). The species has increased and developed a range pattern across middle Ontario and beyond to the New England States and the Maritime Provinces. Though a winter visitant to the east previous to the present century, its summer range was understood to centre in the west. Summer occurrences east of Manitoba were unknown until 1899, none as far east as the west end of Lake Superior until 1910, and none east of Lake Superior until 1915. Currently the range pattern is a fairly narrow belt corresponding rather closely to the edge where wilderness is broken by pioneering conditions. A spread and widening of this belt, particularly in the clay belt of eastern central Ontario, is now apparent and follows the environmental conditions mentioned. The Clay-colored Sparrow ( Spizella pallida) is another western bird undergoing eastward range expansion. Scattered occurrences in summer east to Fort Albany in the north (R.O.M., specimen) and Grenville County in the south (Terrill, 1952) have taken place since 1924 (Dale, 1927). It is now established as a breeding bird west of Lake Superior (Snyder, 1938 and 1953) and in the Sault Ste. Marie region (Snyder, et al., 1942), and among the numerous summer occurrences for southern Ontario there are now several nesting records (Baillie, 1955a). Occurrence of other western bird species have been noticed in the east, but establishment there is still in question. The Western Kingbird (Tyrannus verticalis) is an example. In its case there is one actual nesting record, for 31

Kent County in 1943, but the trend toward establishment in the east appears to have waned. Also the Double-crested Cormorant (Phalacrocorax auritus, nominate race) has expanded its population across the province from west to east but this is· a filling-in process since the species occupies the east, from the Gulf of St. Lawrence to New England. By far the most numerous examples of range extensions, within and into the province, are northward in trend. First, the Egret (New World race Casmerodius albus egretta) included in the provincial list by Mcilwraith ( 1886) on the basis of a single record, is now established as a breeding species on East Sister Island in Lake Erie. This represents a spread (Baillie, 1953) from West Sister Island (Ohio) where a colony became established in 1946 (Campbell, 1947). Although the present range expansion is linked with population recuperation of a once-threatened species, it is of interest to note that its former breeding range did not extend north to Lake Erie. The Turkey Vulture ( Cathartes aura, apparently both of its racesC. c. septentrionalis in the east and C. a. teter in the west) has extended its range northward in Ontario during the 20th century. Mcilwraith (1886) reports it as frequently seen in Essex County seventy years ago. The first nesting was in 1901 at Dashwood, Huron County, and by 1911 it was established as a breeding species in Middlesex County. Also in 1911 it was found near Kenora (Snyder, 1953). By 1934 the species was found breeding north to Grey County (Baillie and Harrington, 1936-37). Currently the species can be expected west of Lake Superior north to a line running from Sydney Lake southeastward to Wabigoon and eastward to southwestern Thunder Bay District. In the east, range limits extend from Middle Bruce Peninsula to Bracebridge, thence through Peterborough to middle Frontenac County. The population is nowhere dense, and suitable conditions for nesting are wanting in many districts within these limits, but it is obvious that the Turkey Vulture has become established at least 200 miles beyond its range of fifty years ago. The Mourning Dove ( Zenaidura macroura, with particular reference to z. m. carolinensis) has a similar history of range expansion northward. It occurred sparingly in the region of early settlement more than a century ago, but a precise northern limit at any particular early period cannot be drawn. A survey of the literature, and the life-time impressions of observers, leave no doubt of the general increase in the population in the south, and it is apparent that its range frontier has also advanced northward. The region west of Lake Superior must now be considered marginal for the species and there seems little doubt that this is an extension since the country there was opened up for agriculture approximately 65 years ago (Snyder, 1938). In the east the northern limits of its breeding range twenty years ago were considered to be north to Simcoe and Carleton counties (Baillie and Harrington, 1936-37). Southern Algoma District and Renfrew County represent the current limits though there is an increasing number of occurrence records beyond. An estimate of the northward spread during the present century, from the line 32

of sparse breeding status in 1900 to the line of sparse breeding status today, would indicate approximately a 100-mile advance in 50 years. The Black-billed Cuckoo (Coccyzus erythropthalmus) is another species which has obviously advanced its range northward, but precise .limits for early periods cannot be drawn. It was present in areas of early settlement along the north shore of Lake Ontario more than 100 years ago according to the Charles Fothergill Manuscript (R.O.M., Clendenan volume), but its population was sparse. By 1888 it was known to occur in Muskoka District (Baillie and Harrington, 1936-37). At the present time it can be expected in southern Kenora District north to Malachi and Sioux Lookout (Snyder, 19 5 3) and in the east it has been known to nest north to Matheson with several occurrences north of this area, even to Moosonee. It is estimated that northward range extension approximates 200 miles, attained in the present century. The Tufted Titmouse (Parus bicolor) was not observed in the province until May, 1914 (Saunders, 1914) when it appeared at Point Pelee. Since then it has spread slowly and become established sparsely as a nesting resident (Baillie, 1955b) with occurrence records as far north as southern Georgian Bay and east to Toronto. If we consider only breeding evidence as proof of establishment, a spread of 140 miles in 40 years is indicated. Moseley ( 1947) states that the species has spread northward in New York and Michigan, and notes that a general expansion is evident in Ohio. Bewick's Wren (Thyromanes bewickii, race T. b. altus) was first found in the province in 1898 when a specimen was collected in southwestern Middlesex County in December, 1898 (Saunders, 1906). In succeeding years, at first with intervals of several years, but with increasing frequency, it has been reported, for various seasons in southern counties, but more significantly north to York and Grey counties in summer. In 1950 it was found nesting at Point Pelee (Baillie, 1950a). Though not widely established, it ranks as a new element in our avifauna which has moved in from the south. Moseley (194 7) states that the northward extension of range of this wren, from the states south of Ohio, started in the 1870's. The distance from southern Ohio to Point Pelee is somewhat over 200 miles. The Carolina Wren (Thyrothorus ludovicianus;with respect to the nominate race) has a similar history. It first occurred, in Lambton County, in February, 1891 (Anon., 1891). Evidence of its having established itself in our region was first indicated by the taking of fledglings at Point Pelee in 1905 (Klugh, 1906). By 1916 the species was established north to Lambton and Elgin counties. Penetration as a nesting bird has extended to Toronto (Lindsay, 1931 ) , and there are occurrence records north to Woodville ( Baillie and Harrington, 1936-37) and Ottawa (Bourguignon, 1945). Moseley (1947) states that the Carolina Wren began its range expansion northward into Ohio in the 1880's. On this basis the species has advanced northward at least 350 miles in 70 years. The Mockingbird (Mimus polyglottos, concerning the nominate race) was 33

first discovered at Chatham, Ontario, in 1860 (Morden and Saunders, 1882). After a lapse of several years, the first breeding status was established, at Amherstburg (Barrows, 1912). Spread has continued, with periodic interruptions, and establishment as a nesting species has reached as far north as Manitoulin Island (Baillie, 1955c). There are sporadic occurrences north to Fort William and Moosonee (Virtue, 1929). Moseley (1947) refers to the northward spread of the Mockingbird in Pennsylvania and New England, and since he records the species as present in Ohio for many years before the present century, we have but to deal with the Ontario range to determine its spread, which is approximately 240 miles northward, largely during the 20th century. Another case, which though not pronounced indicates a trend toward northward expansion of range, is the Prothonotary Warbler (Protonotaria citrea) . This species was first recorded as a spring wanderer for Hamilton in 1888 (Mcilwraith, 1888). It was not until 1929 that it was found established as a breeding species, at Rondeau Park ( Saunders, 1935) . This remained the only breeding station for eleven years, but now it is established in Essex, Kent, Elgin, Norfolk, and Wentworth counties (Baillie, 1954). The most remote occurrences are set currently at the base of the Bruce Peninsula and Port Hope. This case represents a range extension into Canada from the south, probably during the present century. The Golden-winged Warbler (Vermivora chrysoptera) is another species showing evidence of northward range expansion. Mcilwraith (1886), speaking of its status during the latter part of the 19th century, stated that it had been observed only twice at Hamilton but that it was more common near London where Morden and Saunders (1882) called it regular and rather common. The first record (spring) at Guelph (Howitt, 1924) was well into the present century, as were records more remote from the 19th century centre of numbers. Currently, the species occurs sparsely to northern Simcoe County (Devitt, 1943) and fans out to Peterborough (Baillie, 1954) and Hastings counties, with evidence of breeding to near these limits. The spread beyond the 19th century limits approximates 150 miles northward. The Bobolink (Dolichonyx oryzivorus) was present, but scarce, along the north shore of Lake Ontario in the early part of the 19th century. Charles Fothergill (R.O.M., MS., Clendenan volume) states that the Bobolink came to inhabit a newly made farm in the Port Hope area in 1810. The fact that he personally did not meet with it in that region until his fourteenth summer there, in 1830, attests to its scarcity. It is true that the Bobolink could have existed in natural, wet meadows, and in floodlands of streams in the forest, beyond this latitude, but unquestionably it has expanded its range northward during the present century. Fleming ( 1901) states that it was one of the species moving northward in Muskoka District in 1897, and in Parry Sound District in 1899. It came to occupy the Port Arthur region in 1932 (Dear, 1940). In the east it is now established far beyond where we might have expected it a century ago. It nests at Chapleau (Baillie, 1950b), and it is 34

known to occur in summer, in numbers approximating those along the north shore of Lake Ontario in Fothergill's time, as far north as Missinabi, Strickland, Kapuskasing, and Matheson (R.O.M. specimens). This limit is some 400 miles north of the north shore of Lake Ontario. A similar history pertains to the Eastern Meadowlark (Sturnella magna, with particular reference to the nominate race) . It was a familiar bird of the meadows and pastures of the pioneer farmland along the north shore of Lake Ontario in 1827 according to Charles Fothergill (R.O.M., MSS; Clendenan and McGillivray volumes). It too could have occurred in limited numbers in natural clearings and burns of the eastern forest well north of the north shore of Lake Ontario. Fleming (1901) gives the date of first appearance in Muskoka as 1863. The species was recorded as very rare at North Bay in 1904 (Thompson, 1922), but by 1939 it was recorded as "abundant" at the same place (Ricker and Clarke, 1939). At Sault Ste. Marie, where it is still not particularly plentiful, it became established about 1900 (Snyder, et al., 1942). Currently, outlying summer occurrences extend north to Gogama and even to the Cochrane area (Baillie, 1955c). The range extension northward is estimated to approximate 200 miles during the present century. A still more striking case is that of the Cowbird (Molothrus ater, nominate race and M.a. artemisiae). Fothergill (R.O.M., MSS) did not know this species, and it can be taken as reasonably certain that it did not occur in the Port Hope region on the north shore of Lake Ontario between 1817 and 1840, the period of his residence there. It was present in the southern counties in 1886 (Mcllwraith, 1886). Fleming (1901) records its first appearance in Muskoka District in 1889, and the first Parry Sound District record is for 1899. Thompson (1922) saw only three Cowbirds during the whole season of 1904 at North Bay, but Ricker and Clarke (1939) refer to the species as frequent there twenty years later. In the region west of Lake Superior, there is no early history, but Dear ( 1940) remarks on its increase in recent years, which is significant. The range of summer occurrence has definitely expanded during the past 30 years beyond the places mentioned. There are records of widely scattered occurrences north to the Canadian Transcontinental Railway with remote occurrences north to Trout Lake, Lake Attiwapiskat and Moose Factory. Unquestionably the species has moved northward 200 miles or more during the present century. The Cardinal (Richmondena cardinalis, nominate race) is probably the classic case showing northward range expansion during the past half century. Moseley ( 1947) dwells particularly on its population increase in Ohio, where its occurrence was from early times peripheral. The American Ornithologists' Union Check-List of North American Birds for 1886 gives its range limits as "north to New Jersey and the Ohio valley ( casually farther). . . ." Mcllwraith ( 1886) records the species as casual along the southwestern border of southern Ontario. Actually the first record for the province was in 1849 at Chatham (Morden and Saunders, 1882). It was first established 35

as a breeding bird at Point Pelee in 1901. Then followed a succession of breeding records which indicates the northward trend and eastward spread: Chatham in 1909; London in 1915; Brantford in 1919; Toronto in 1922; Orillia and Port Hope in 1939; Owen Sound in 1942; and Tweed in 1953. Probably the period of greatest population expansion occurred in 1938. There are also occurrence records in such outlying places as Port Arthur, Sault Ste. Marie, Gogama, Kirkland Lake, and Ottawa. Obviously the species has moved northward 250 miles in 30 years. Numerically the Grasshopper Sparrow (Ammodramus savannarum, race A. s. practensis) was regarded by Mcilwraith (1886) as very rare and its range limited to the southern border during the latter part of the 19th century. That author mentions but two records of occurrence known to him up to 1886. During the early part of the 20th century there were only two records of occurrence at Toronto (Fleming, 1907), and during approximately the same period none had been reported as far north as Muskoka District. Admittedly the species is one readily overlooked, but the fact that it was identified, collected, and reported early would have focussed the attention of the very capable field observers of a generation ago upon it. It would therefore appear acceptable to interpret the steady chronological increase of reports applicable to a widening geographic range, as indicative of a change of status of the species and not entirely an increase and sharpening of perception of collectors and observers. Currently the Grasshopper Sparrow is found in summer north to Manitoulin Island and central Parry Sound District where it breeds, and it has occurred east to Ottawa (Lloyd, 1924). Record of four pairs observed during the summer near Fort William (Denis, 1953) may involve the western race (A. s. perpallidus). This occurrence is the most remote to date. It is estimated that a northward extension of range of this species during the past 50 years approximates 200 miles. There are a number of additional cases which suggest northward range extension but because of faulty history or because it is too early to be certain of a trend, they are not tallied with more definite cases. There is little doubt that the Indigo Bunting (Passerina cyanea) has extended its range northward, but because of a total lack of early faunal lists for the northern part of its present range, no indication of its status there for the 19th or early 20th century can be gained. It was present in the south during the time of Fothergill, 1817-1840 (R.O.M., MS, McGillivray volume). Mcilwraith (1886) states that it probably does not penetrate far north in Ontario. Currently the Indigo Bunting nests north to the Cochrane region in the east (Baillie, 1955c), but, it should be added, in small numbers beyond southern Ontario, i.e., Sault Ste. Marie to North Bay. The Dickcissel (Spiza americana) has on occasions pushed northward in southern Ontario ( and in southern Manitoba) and records of occurrence concern stations as far inland as Middlesex County, where it has nested (Saunders and Dale, 1933), the north shore of Lake Ontario (Snyder, et al., 1941) and even to the Ottawa area (A. G. K[ingston], 1895). But these invasions were apparently fleeting. 36

The history of Henslow's Sparrow (Passerherbulus henslowii, nominate race) is not unlike that of the Grasshopper Sparrow although it was not recorded for the province until the close of the 19th century ( Saunders, 1898) at the mouth of the Thames River on Lake St. Clair, and at Samia. However, the fact that it could have occupied open, wet swales in the forest, whereas the Grasshopper Sparrow inhabits drier, grassy uplands which were once forested, and the impression that the habits and behaviour of Henslow's Sparrow causes it to be comparatively more obscure, makes the evidence of change in its case less convincing. Other species have changed status from fortuitous to casual, or even frequent, but without substantial evidence of establishment. They need only be mentioned-The White-eyed Vireo ( Vireo griseus, nominate race); the Blue-winged Warbler, (Vermivora pinus); the Kentucky Warbler (Oporornis formosus); and the Hooded Warbler ( Wilsonia citrina). Others which are definitely established in the province, but whose chronological history of numbers and distribution are too incomplete to demonstrate northward spread, include the Blue Jay ( Cyanocitta cristata) ; the Blue-gray Gnatcatcher (Polioptila caerulea); the Yellow-breasted Chat (/cteria virens); and the Field Sparrow (Spizella pusilla). Certainly a life-time of faunal work in the province leaves the impression that range changes in a northward direction may have taken place in the case of each of these species and perhaps others, such as the Catbird (Dumetella carolinensis), and Brown Thrasher (Toxostoma rufum). DISCUSSION

Among the first three categories of change, twelve species have been mentioned. Under EXTIRPATIONS there are two, the Passenger Pigeon and the Turkey. Under RANGE SHRINKAGES AND FLUCTUATIONS, six were particularly selected, the Ruffed Grouse and Redstart, the Spruce Grouse, Bob-white, Prairie Chicken, and Sandhill Crane. Under INTRODUCED EXOTICS we have the European Partridge, Common Pheasant, Starling, and House Sparrow. All of these cases represent significant modifications in Ontario's avifauna. Most, if not all, can be attributed to the direct or indirect works of man. The fourth category, RANGE EXPANSIONS, includes twenty-three cases. These have been classified according to directional trend in range expansion. Southward, there is none; Westward, two, the Black Duck and Black-backed Gull; Eastward, seven, Holboell's Grebe, Ring-necked Duck, Homed Lark, Western Meadowlark, Brewer's Blackbird, Evening Grosbeak, and Claycolored Sparrow; Northward, fifteen, Egret, Turkey Vulture, Mourning Dove, Black-billed Cuckoo, Tufted Titmouse, Bewick's Wren, Carolina Wren, Mockingbird, Prothonotary Warbler, Golden-winged Warbler, Bobolink, Eastern Meadowlark, Cowbird, Cardinal, and Grasshopper Sparrow-more than twice as many species as appear to be expanding range in any other direction. In order to approach even speculation as to the factors involved, it is 37

advisable first to remark briefly on certain attributes of birds in general. As a Class birds are notoriously mobile creatures. They have developed a considerable measure of independence from their environment through temperature control and by special adaptations of structure and habit. But in spite of these attributes, no species is universally distributed. Each is limited in its dispersal by barriers, seen and unseen, and a rather precise range pattern can be circumscribed for each at a given period in history. But these patterns change, in some cases markedly, in a relatively short time. Obviously, certain bird species are capable of living in regions in which they are not indigenous. The Starling and House Sparrow, once lifted by man over an ocean barrier and released in a congenial environment, multiplied and spread until other barriers inhibited further expansion. The Fieldfare (Turdus pilaris) carried from Europe over the north Atlantic by a storm, has become established in southern Greenland (Salomonsen, 1950). Spread there will be limited by factors of their new environment: availability of food; the presence of environmental associations consistent with innate habitat choice; climate, particularly during the critical reproductive period when conditions must be close to the native optimum (Twomey, 1936). All these limiting agencies remain in force even though a physical barrier is hurdled. It has been observed that the sedentary House Sparrow, for example, has spread north to James Bay but that many, perhaps all, die off in winter (Williams, 1920). Continued occupation by the species probably depends on re-occupation by new pioneers. It is pertinent here briefly to sketch the nature of birds with respect to the production of pioneers. Bird banding has established the rule that adult birds tend to return for nesting to the same place, often the precise place, where they nested previously, and young birds tend to return to the general region but not to the precise place of their origin. The concept that young birds are forced to scatter is reinforced by the fact that the majority of species, at least of land birds, are territorial in behaviour during the breeding season. It is apparent then that pioneers of unoccupied terrain including marginal areas are usually first-nesting or relatively young individuals, the latest models of gene combination of their kind. It is they who establish the tradition of occupancy (Snyder, 1948) and press against dispersal barriers, within and at the periphery of range, for theoretically this pressure is exerted in all directions, not in a particular direction. If we find multiple species pressing simultaneously into previously unoccupied peripheral terrain in a particular direction, we would scarcely expect that there has been multiple and simultaneous increase in tolerance of species to new conditions. Rather we would look for some weakening of barriers or amelioration of environment which facilitate spread in that particular direction. In making pseudo-plains where once there was forest, and in increasing food supply through agricultural practices, an invitation to certain campestrian forms to expand eastward is apparent. But such does not explain all eastward expansions. Modifications of the environment by man through deforestation 38

and agriculture undoubtedly have facilitated the spread of certain southern forms northward. Sixty-five percent of all cases of range expansion in the province are in that direction. Other factors would appear to be involved, and these may include change in climate, whether on a short or long term basis. LITERATURE CITED E., AND L. s. DEAR 1947. Brewer's Blackbird breeding in Ontario. Wilson Bull., vol. 59, no. 3, pp. 175-176.

ALLIN, A.

ANONYMOUS

1891. Meeting of the London ornithological section. Canad. Ent., vol. 23, no. 5, p. 109. BAILLIE, JAMES L. 1940. The summer distribution of the Evening Grosbeak. Canad. Fld. Nat., vol. 54, no. 2, pp. 15-25. 1947. Prairie Chickens in Ontario. Sylva, vol. 3, no. 1, pp. 49-51. 1950a. Region reports. Audubon Fld. Notes, vol. 4, no. 4, p. 240. 1950b. Region reports. Audubon Fld. Notes, vol. 4, no. 5, p. 274. 1951. Region reports. Audubon Fld. Notes, vol. 5, no. 1, p. 13. 1953. Regional reports. Audubon Fld. Notes, vol. 7, no. 5, pp. 306-307. 1954. Regional reports. Audubon Fld. Notes, vol. 8, no. 5, pp. 342-344. 1955a. Region reports. Audubon Fld. Notes, vol. 9, no. 3, pp. 254-256. 1955b. Regional reports. Audubon Fld. Notes, vol. 9, no. 4, pp. 328-329. 1955c. Regional reports. Audubon Fld. Notes, vol. 9, no. 5, pp. 375-377. BAILLIE, JAMES L., JR., AND PAUL HARRINGTON 1936-37. The distribution of breeding birds in Ontario. Trans. R. Canad. Inst., vol. 21, pt. 1, pp. 1-50, and pt. 2, pp. 199-283. BARROWS, WALTER B.

1889. The English Sparrow (Passer domesticus) in North America. Bull. U.S. Dep. Agric., no. 1, p. 17. 1912. Michigan bird life. Spec. Bull. Mich. Agric. Coll., p. 657. BENT, ARTHUR CLEVELAND

1923. Life histories of North American wildfowl. Bull. U.S. Nat. Mus., no. 126, p. 229. BONNYCASTLE, RICHARD H.

1841. The Canadas in 1841. London, Eng., vol. 1, p. 168. E. 1945. Ottawa bird records. Canad. Fld. Nat., vol. 59, no. 4, p. 150.

BoURGUIGNON, A.

CAMPBELL, LOUIS W.

1947. American Egrets nesting on West Sister Island in Lake Erie. Auk, vol. 64, no. 3, pp. 461-462. CLARKE, C. H. D. 1948. The Wild Turkey in Ontario. Sylva, vol. 4, no. 6, pp. 4-12. 1954. The Bob-white Quail in Ontario. Bull. Fed. Ont. Nat., no. 63, pp. 6-15. CLARKE, C. H. D., AND R. BRAFFETTE 1947. Ring-necked Pheasant investigation in Ontario 1946. Ont. Dept. Lands and Forests [Mimeo.], pp. 1-11. DALE, E. M. S. 1927. The Clay-coloured Sparrow at London, Ontario. Canad. Fld. Nat., vol. 41, no. 3, pp. 64-65. 39

L. s. 1940. Breeding birds of the region of Thunder Bay, Lake Superior, Ontario. Trans. R. Canad. Inst., vol. 23, pt. I, pp. 119-143.

DEAR,

[DENIS, KEITH]

1953. [Editorial note] Thunder Bay Fld. Nat. Club News Letter, vol. 7, no. 2, p. 8. DEVITT, 0. E. 1943. The birds of Simcoe County, Ontario. Trans. R. Canad. Inst., vol. 24, pt. 2, pp. 284-285. FLEMING, JAMES H.

1901. A list of the birds of the Districts of Parry Sound and Muskoka, Ontario. Auk, vol. 18, no. 1, pp. 33-45. 1907. Birds of Toronto, Canada. Auk, vol. 24, no. 1, p. 80. FORBUSH, EDWARD HOWE

1927. Birds of Massachusetts and other New England States. Mass. Dep. Agric., vol. 2, pp. 365-373. HOWITT, HENRY

1924. Golden-winged Warbler in Canada. Auk, vol. 41, no. 3, p. 614. JONES, S. PAUL

1938. Holboell's Grebe and American Brant in Wisconsin. Auk, vol. 55, no. 3, p. 666. K[INGSTON], A. G. 1895. A new bird for eastern Ontario. Ottawa Nat., vol. 9, no. 4, pp. 92-93. KLUGH,A. B. 1905. The birds of Wellington County, Ontario. Ont. nat. Sci. Bull., no. 1, p. 4. 1906. The Carolina Wren, a summer resident of Ontario. Auk, vol. 23, no. 1, p. 105. LINDSAY, ROBERT V. 1931. Carolina Wren breeding at Toronto. Canad. Fld. Nat., vol. 45, no. 8, pp. 205-206. LLOYD, HOYES

1924. The birds of Ottawa, 1923. Canad. Fld. Nat., vol. 38, no. 1, p. 10. C. 1888. Prothonotary Warbler in Ontario. Auk, vol. 5, no. 3, pp. 322-323.

MclLWRAITH, K.

MclLWRAITH, THOMAS

1886. The birds of Ontario. Hamilton Assoc., pp. 1-304. L. 1938. Ring-necked Duck breeding in eastern North America. Auk, vol. 55, no.3,pp.401-404.

MENDALL, HowARD

MITCHELL, MARGARET

H.

1935. The Passenger Pigeon in Ontario. Contr. R. Ont. Mus. Zool., no. 7, pp. 1-181. MORDEN, J. A., AND W. E. SAUNDERS 1882. List of the birds of western Ontario. Canad. Sports. and Nat., vol. 2, no. 11, pp. 183-187, and no. 12, pp. 192-194. MOSELEY, EDWIN LINCOLN

1947. Variation in the bird populations of the north-central states due to climatic and other changes. Auk, vol. 64, no. 1, pp. 15-35. MURRAY, ANDREW

1859. Contribution to the natural history of the Hudson Bay Companies territory, Part III-Aves. Proc. Edin. New Phil. Journ., n.s., vol. 9, no. 2, p. 225. 40

E. 1909. Some old time reminiscences of old Ontario. Rod and Gun in Can., vol. 11, no. 6, pp. 524-526. RICKER, WILLIAM E., AND C. H. D. CLARKE 1939. The birds of the vicinity of Lake Nipissing, Ontario. Contr. R. Ont. Mus. Zool., no. 16, pp. 1-25. ROBERTS, THOMAS A. 1932. The birds of Minnesota. Univ. Minn. Press, Minneapolis, vol. 1, pp. 1-681. ORR,JAS.

SALOMONSEN, FINN

1950. The immigration and breeding of the Fieldfare (Turdus pilaris) in Greenland. Proc. Xth Int. Ornith. Cong., Uppsala, pp. 515-526. SAUNDERS, w. E. 1888. First occurrence of Western Red-tail in Ontario. Auk, vol. 5, no. 2, p. 205. 1898. Henslow's Sparrow (Coturniclus Henslowi). Ottawa Nat., vol. 12, no.9,pp. 177-178. 1906. Birds new to Ontario. Ottawa Nat., vol. 19, no. 11, p. 206. 1910. Pinnated Grouse in southern Ontario. Auk, vol. 27, no. 1, p. 79. 1914. The Tufted Tit-a new record for Canada. Auk, vol. 31, no. 3, p. 402. 1935. Southern Ontario birds. Bird-Lore, vol. 37, no. 5, p. 326. SAUNDERS, W. E., AND E. M. s. DALE 1933. History and list of birds of Middlesex County, Ontario. Trans. R. Canad. Inst., vol. 19, pt. 2, pp. 161-215. SCHMIDT, KARL P.

1938. Herpetological evidence for post-glacial eastward extension of the steppe in North America. Ecology, vol. 19, no. 3, pp. 396-407. SHEPPARD, R. w., w. E. HURLBURT, AND G. H. DICKSON 1936. A preliminary list of the birds of Lincoln and Welland Counties. Canad. Fld. Nat., vol. 50, no. 8, pp. 137-138. SNYDER, L. L. 1938. A fauna} investigation of western Rainy River District, Ontario. Trans. R. Canad. Inst., vol. 22, pt. 1, pp. 157-213. 1941. The birds of Prince Edward County. Univ. Toronto Stud. biol., no. 48, p. 55. 1948. Tradition in bird life. Canad. Fld. Nat., vol. 62, no. 2, pp. 75-77. 1951. Ontario birds. Clarke, Irwin & Co. Ltd., Toronto, pp. 88-89. 1953. Summer birds of western Ontario. Trans. R. Canad. Inst., vol. 30, pt. 2, pp. 157-213. 1954. On the distribution of Colymbus grisegena holbollii. Can. Fld. Nat., vol. 68, no. 1, pp. 41-42. SNYDER, L. L., E. B. s. LOGIER, AND T. B. KURATA 1942. A faunal investigation of the Sault Ste. Marie region, Ontario. Trans. R. Canad. Inst., vol. 24, pt. 1, pp. 99-165. SNYDER, L. L., E. B. s. LOGIER, T. B. KURATA, F. A. URQUHART, AND J. F. BRIMLEY 1941. A faunal investigation of Prince Edward County, Ontario. Univ. Toronto Stud. biol., no. 48, p. 86. SPEIRS, J. MURRAY, GEORGE W. NORTH, AND JOHN A. CROSBY 1944. Holboell's Grebe nesting in southern Ontario. Wilson Bull., vol. 56, no.4,pp.206-208. TERRILL, LEWIS Mel.

1952. The Clay-colored Sparrow in southwestern Ontario. Canad. Fld. Nat., vol. 66, no. 5, pp. 145-146. 41

L. 1922. The birds of North Bay, Ontario, and vicinity in 1904. Canad. Fld. Nat., vol. 36, no. 9, pp. 161-168. TwoMEY, ARTHUR C. 1936. Climographic studies of certain introduced and migratory birds. Ecology, vol. 17, no. 1, pp. 122-132. THOMPSON, STUART

VIRTUE, ROBERT

, 1929. Mockingbird (Mimus po[yglottos). Canad. Fld. Nat., vol. 43, no. 5, p. llO.

WALKINSHAW, LAWRENCE

H.

1949. The Sandhill Cranes. Bull. Cranbrook Inst. Sci., no. 29, pp. 1-202. WHITE, JAMES S. 1935. Further notes on Brewer's Blackbird (Euphagus cyanocephalus) breeding in northeastern Illinois. Auk, vol. 52, no. 1, p. 94. WILLIAMS,

M. Y.

1920. Notes on the fauna of the· Moose River and the Mattagami and Abitibi tributaries. Canad. Fld. Nat., vol. 34, no. 7, p. 124.

42

CHANGES IN THE MAMMALIAN FAUNA OF ONTARIO BY

RANDOLPH L. PETERSON

THE GEOGRAPHIC AREA now comprising the Province of Ontario has been the scene of drastic faunal changes during its geological history. While it is the primary purpose of this paper to examine some of the more recent changes that have taken place, i.e., within the scope of white man's existence here, personal studies with various mammal groups have forcibly impressed this writer with the contention that many of the noticeable changes in distribution have been correlated with a post-glacial repopulation of Ontario following a total coverage of ice during the past glacial period. Different species have followed different modifications of a basic pattern of re-occupation around the east and west ends of the Great Lakes system. Most of the range expansions of mammals that have taken place in Ontario involve a general spread northward. Other conspicuous expansions involve either a western spread of eastern forms, such as Sorex fumeus or Microtus chorotorrhinus ( although as yet we have been unable to · establish recent expansion trends), or the eastern spread of western forms such as Eutamias ,:ninimus or Canis latrans. In one sense both the western and eastern expansions might also be interpreted in part as secondary manifestations of the general northern range expansions in that they are expanding east or west after moving north beyond the Great Lakes system. It therefore seems logical to assume that all range expansions of native species can be associated with postglacial development. There has been no apparent southward spread of northern forms with the possible exception of the brushwolf, Canis latrans. Other cases of range expansions involve introductions. These will be dealt with separately. Range reductions present a complex array of causative factors. In certain cases actual extirpation and extinction have already resulted. Closely associated are the changes in the fauna which involve fluctuation in numbers, some of a short cyclic nature, others less rhythmic, while still others may result in a slow, fairly persistent decline which may ultimately result in range reduction, extirpation, and perhaps even extinction in some cases. Since all populations are subject to fluctuations in numbers, only a few examples will be singled out to illustrate some of the basic types of population changes that have taken place or are now in the process of taking place. Consideration, in the following order, will be given: (1) range expansions; (2) introductions; ( 3) range reductions; ( 4) population fluctuations. 43

li:ii:il GLACIAL ICE

-GRASSLAND l!!!!!!!!I GIGAS 111111111 SH IR ASI m.a,ANDEASONI ~AMERICANA

Fm. 1. The distribution of moose in North America during the greatest extent of Wisconsin glaciation (Fig. 4 from Peterson, 1955) .

. ··•,-.. __ J .. --···· EXTENT OF GLACIATION' ; . ---- LAURENTIDE ICE SHEET STAGES ·--·-COADILLERAN ICE SHEET ROUTES OF MOOSE DISPERSAL

Fm. 2. The pattern of post-glacial dispersal of North American moose (Fig. 5 from Peterson, 1955). 44

· Nor 955). about 1875 (Fig. 6 from . f moose m Fm. 3. -The -~=--:~~-:-;~or,t~h!,;~Ai~merica cc distribution o Peterson, 1

~::-:---"-...--7:

45

RANGE EXPANSIONS

Alces alces, MOOSE In the case of the moose it was quite fortunate that this large, conspicuous mammal was sufficiently impressive, when it invaded a new area, to be noted in the pages of history. This historical record, coupled with morphological studies of geographic variation, has corroborated the reconstruction of the pattern as well as its development, both in time and space, of the re-occupation of northern Ontario by moose. It is unlikely that the past history of distributional shifts of many smaller, less conspicuous mammals will be available to us in similar detail. During the Wisconsin stage of glaciation moose were divided into four areas of habitat (Fig. 1). As the glacial ice retreated, the central portion of Canada

NUOION

IAY

: ! I

t...,. ',

-

-- ... ......

NORTHERN OISTIBUTION OF MOOSE IN ONTARIO •·••·•••• A,,IOXIMATI UMln AIOUT IIH • - - - A,,ROICINAH UMlfl HOUT IIU .,._, .. ar,1110,uN.U( UNITS AIOUJ lflO

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Fm. 5. The spread of moose into Northern Ontario (Fig. 8 from Peterson, 1955). 46

became free of ice long before the more persistent Laurentian and Cordilleran ice sheets retreated (Fig. 2). This allowed an early expansion northwestward from the area west of the Great Lakes. By 1875 (Fig. 3) the distribution of moose had reached the extent where first generally encountered by white man. The next 75 years (Fig. 4) shows a remarkable expansion into most of the suitable habitat of northern Canada. Data from a wide variety of sources have all shown remarkable agreement in the reconstruction of the range expansion across the central portion of northern Ontario (Fig. 5). For a more detailed discussion of the postglacial dispersal and recent range expansion.. see Peterson (1955, pp. 14-18; 45-51). Odocoileus virginianus, WHITE-TAILED DEER The white-tailed deer has followed the same basic pattern of northern range expansion as the moose, although it was a few years behind the advance of the moose and has not extended its normal range quite as far northward. The various stages of this spread are shown in Figure 6 which has been modified from Cross (1937b). There is evidence that the white-tailed deer, unlike the moose, utilized the Sault Ste. Marie entrance from the upper peninsula of Michigan.

FIG. 6. The northern spread of the white-tailed deer in Ontario from its original range of about the year 1800 to its present distribution (modified from Cross, 1937b). 47

Lynx rufus, BOBCAT In a recent study (Peterson and Downing, 1952) it has been shown that the bobcat has recently extended its range noticeably in the western portion of the province and in the area east of Lake Superior, while there appears to have been a decrease in numbers in southern Ontario south of Lake Nipissing (Fig. 7). The strong invasion from the upper peninsula of Michigan into the Sault Ste. Marie region since 1945 presents an interesting variation from the pattern established for moose. The trend has been an eastern and northern spread from the west rather than a straight northern movement of the southern or eastern form from southern Ontario.

i

! i

i i

i

192'M.\ .----LEGEN,D-------

•

Spe.cimens in Collections.

o Specimens recorded in thf. . Litera.tur" and other records.

x Occurre.nce. doubte.cl, or omitted from extensive fauna.l lists.

Fm. 7. The distribution and spread of bobcats in Ontario (from Peterson and Downing, 1952, p. 16).

48

Canis latrans, BRUSH WOLF The spread of brush wolves was first an eastern movement paralleling that of the Lake Superior bobcat, but subsequently it has occupied all of the central portion of the province and has spread southward. Cross ( 1937a) has provided a map showing the distribution of 1900 (restricted to Rainy River and western Kenora Districts) and in 1936 when it was known to have extended well north (see Fig. 8) and south to a line roughly joining the south end of Lake Huron and Pembroke. Records since 1936 show clearly that the entire area of southern Ontario may be included in its range. Compared with the timber wolf (Canis lupus) which has long occupied most of Ontario in rather static numbers, the brush wolf numbers have shown rather strong and presumably cyclic fluctuations. Its recent entry into the agricultural areas of southern Ontario has been accompanied by a certain amount of hybridization with domestic dogs. The extent and effect of this hybridization is presently being investigated.

1936

-......,

',

,,,

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I

,I

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Fm. 8. The spread of brush wolves in Ontario (modified from Cross, 1937a). 49

Urocyon cinereoargenteus, GRAY FOX The gray fox provides an example of a species that occupied southern Ontario for a period up to the coming of white man ( as evidenced by material collected at early Indian midden sites) but disappeared for almost 300 years and has recently (circa 1930-40) made another invasion (Downing, 1946). In a recent review of the latest records of occurrence (Peterson, Standfield, McEwen, and Brooks, 1953) it was shown that "at the present time three populations of the gray fox are apparently extending their ranges into Ontario, U. c. borealis around the east end of Lake Ontario, U. c. ocythous in the Rainy River region west of Lake Superior, and a third population, probably U. c. cinereoargenteus, around the east end of Lake Erie." Didelphis marsupialis, OPOSSUM The opossum provides another example of a species which made more than one invasion into Ontario, but in this case it has been restricted to the most southern latitudes of the province. Peterson and Downing ( 1956) have reviewed the records of its occurrence in Canada. The first record known was a report in 1858. The next mention of the opossum occurred in 1875. No further records were available until 1892-1900 when several animals were observed. Subsequent records were not available until 1934 when a single animal was taken in Kent County. Beginning in 1947, records of the opossum have become increasingly more common, and its range has been extended farther north. Of the mammals considered, it would appear that the opossum is undoubtedly the one most affected by low winter temperatures as a limiting factor in its northern distribution. Its low metabolism and naked tail and ears and poorly furred feet would not allow this species to become readily adapted to our colder climates. OTHER SPECIES

There is little doubt that several other species have extended their range in Ontario within the last 300 years, but unfortunately data do not exist to substantiate early limits of distribution. Most small mammals escaped early notice and even today their northern limits of distribution are poorly known. In a few isolated cases, such as that of the western chipmunk, Eutamias minimus (Peterson, 1953) there has been an apparent eastward and southward movement in recent years, although this may be an increase in knowledge of distribution rather than a real recent dispersal. On the whole, however, its pattern follows that of the brush wolf, and there appears no valid reason to suspect that it will not continue to extend farther eastward. While the evidence is not yet available, it would appear logical that real expansions of range westward of eastern forms such as Sorex fumeus and Microtus chrotorrhinus have taken place and will continue in the future since there are no limiting factors apparent that would prevent them from doing so. Several species reach their northern limits of distribution within the province. Extension of range northward on broad fronts will probably proceed 50

rather slowly and in an oscillating fashion. Once the distributional front extends beyond the level of the upper Great Lakes (Lake Superior) opportunity is afforded for the species in question to expand more rapidly east or west. It is perhaps significant that at least two purely western species ( Canis latrans, and Eutamias minimus) have extended well east of Lake Superior while eastern species such as Sorex fumeus and Microtus chrotorrhinus have not yet extended west of Lake Superior. On the other hand such species as Condylura cristata and Blarina brevicauda have been able to proceed west south of the Great Lakes to join northwestern movements from the east, thereby encircling the entire Great Lakes system. In a few other instances there is a certain amount of evidence, though not yet conclusive, that expansion is still in progress. The grey squirrel Sciurus carolinensis was formerly quite rare as far north as Algonquin Provincial Park but recently has become more common. The raccoon was formerly unknown north of Lake Superior (Miller, 1897) but seems to have extended well north of its former range of about 1850. INTRODUCTIONS

Lepus europaeus, EUROPEAN HARE In 1912 nine European hares (L. e. hybridus) escaped near Brantford, Brant County, Ontario, and became established and have spread out to occupy practically all of southern Ontario south of the Precambrian Shield (Reynolds, 1955; Peterson and Reynolds, 1954). By 1930 it had occupied all of southwestern Ontario west of Durham County. It reached Prince Edward County by 1936. Since 1948 at least three records have been reported north of the Precambrian Shield although it seems likely that its range will remain largely restricted to the south of this area. Releases ofthis hare were also made in the Port Arthur-Fort William area on the north shore of Lake Superior in 1942, 1943, and 1945 (Allin, 1950). Survivors were observed up until 1949 but none was seen afterwards. Sciurus niger, FOX SQUIRREL Fox Squirrels (S. n. rufiventer) were introduced to Pelee Island, Lake Erie, about 1895, from southern Ohio (Lloyd, 1925). This population became established and is now commonly encountered on the island (Downing, 1948). Subsequent releases from Pelee Island have been made on Point Pelee (circa, 1950) but the fate of these releases is as yet unknown. Rattus rattus, HOUSE RAT This species has been definitely recorded only on one occasion from a shipment of plants from France to Strathroy, Ontario, in 1927. Specimens in this museum collected from this introduction include both the roof rat (R. r. alexandrinus) and the black rat (R. r. rattus). As far as is known, no established populations presently survive in Ontario. 51

Rattus norvegicus, NORWAY RAT OR BROWN RAT This species, perhaps along with the house mouse (Mus musculus), was probably the first exotic species to reach Ontario. No records are available to establish the original point or points of introductions and subsequent spread. In 1830 it was thought not to have spread farther than the water-front in the Toronto region (Fleming, 1913). In all probability there have been repeated re-introductions to supplement the earlier established populations. At present it is quite common throughout southern Ontario north to a level of North Bay and Sault Ste. Marie, and again west of Lake Superior from the lakehead to Kenora. North of these points the established populations are noticeably restricted to the isolated centres of human habitation providing suitable shelter and food supply. Northern records of occurrence are largely restricted to the level of the northernmost trans-Canada railway line. Undoubtedly introductions are frequent farther north, particularly in the coastal parts of Hudson and James bays, but apparently such introductions survive only temporarily, and as yet no known established populations exist there. Mus musculus, HOUSE MOUSE The introduction and spread of this species closely approximates that of the brown rat with the exception that it apparently has been able to survive more successfully in the more northern areas. In general, its northern limits are similar to those of the brown rat with the exception that it is known to have extended farther up the railway toward Moosonee. Specimens are available from Fraserdale and Smoky Falls. Myocaster coypus, NUTRIA OR COYPU This South American hystricomorph rodent was apparently first introduced as breeding stock for fur farms in Canada about 1927 (Hodgson, 1949), although only a few animals existed in Ontario until after World War II. Several fur breeders across Ontario then became interested in this species and many were imported from various sources. As results of escapes or releases, nutria have since been caught in the wild state from such widespread points as Rainy River District, Thunder Bay District, Lake Erie District, and southern Algoma District (Downing and Lumsden, MS). The last constituted the first wild caught specimen received by the Royal Ontario Museum ( 1944). At present it is not known whether or not this species has survived an entire year or produced young in the wild state in Ontario. Cervus canadensis, WAPITI Attempts have been made to introduce the western form of Wapiti into many areas of Ontario with either limited or no success to date. A partial list of local points of releases includes Algonquin Provincial Park, NipigonOnaman Crown Game Preserve, Chapleau Crown Game Preserve, Pembroke, Burwash, Peterborough, etc. The known distribution of existing herds is shown in Figure 9. 52

RANGE REDUCTIONS

Cervus canadensis, WAPITI The eastern race of the Wapiti, C. c. canadensis, formerly extended throughout most of southern Ontario south of Lake Nipissing (Fig. 9). Although Seton (1927) reports on a specimen reportedly taken near North Bay in 1893, the Wapiti was quickly reduced in numbers following the arrival of white man, and by 1750 most remaining animals were restricted to the Ottawa Valley where they persisted until about 1850. By 1875 the animal was thought to have been extirpated from Ontario and it shortly after became extinct south of the border.

FIG. 9. The known distribution of existing herds of introduced Wapiti with the limits of distribution of the original native Wapiti shown for about 1800. They were already much reduced by that date and became extinct only 50 years later.

Antler fragments from Northern Light Lake and Pickerel Lake in western Ontario (Cahn, 1937) seem to be the only available evidence of the original Wapiti in Ontario west of Lake Superior. Evidently it was extirpated long before it disappeared south of the border in Minnesota where a few animals remained until after 1900. There was a small band reported to have been observed in the Northwest Angle of Minnesota as late as 1932 (Roberts, 53

1945). In all probability the Wapiti formerly present in northwestern Ontano was the Manitoba Wapiti (C. c. manitobensis) rather than the eastern race as has been suggested by some. Rangifer arcticus, BARREN GROUND CARIBOU The Barren Ground Caribou formerly migrated along the Hudson Bay coast as far east as Cape Henrietta Maria. In 1948 the Royal Ontario Museum field party to the latter area secured one set of Barren Ground Caribou type antlers attached to the base of the skull, along with several Woodland Caribou antlers. The Hudson's Bay Company factor at Moose Factory informed this writer that they were in that area regularly until some twenty years previously ( circa 1928). Tyrrell (1913) reported summer occurrence of this species west of Fort Severn in 1912. Banfield (1954) also quotes additional evidence for the former occurrence of this species along the Hudson Bay coast of Ontario. Apparently there have been no records of occurrence in recent years. Rangifer caribou, WOODLAND CARIBOU A recent summary of this species in Ontario has been presented by de Vos and Peterson (1951). Woodland caribou apparently did not occur south of Lake Nipissing during the period of white man in this area. There has been a fairly steady retreat during the period 1800 to 1900 north to a line roughly joining Michipicoten on the east shore of Lake Superior to Lake Abitibi (Fig. 10). They have disappeared also from the Rainy River and southwestern Thunder Bay Districts with only an occasional sighting of these animals occurring in southern Kenora District. Isolated populations remain on the islands of Lake Superior ( Slate Islands, St. Ignace Islands) and of Lake Nipigon. Apparent increases have taken place in the Patricia portion of Kenora District in recent years. Felis concolor, COUGAR The Cougar was extirpated from Ontario soon after 1850. Seton (1927) gives several accounts of occurrences of this mammal in southern Ontario between 1800 and 1853. It was apparently restricted to the area south of a line joining southern Georgian Bay and Renfrew although a few questionable reports of occurrences farther north do exist. It is not known when this species disappeared from the province, but it appears likely none were left by 1900. Lynx canadensis, CANADA LYNX This species formerly occurred, although sparingly, over most of southern Ontario north of the Carolinian Zone north of Lake Erie. Its distribution has been pushed northward to the level of Algonquin Provincial Park, although occasional animals still wander farther south. A general decline in numbers has been evident in recent years, and it cannot be considered very 54

abundant anywhere in Ontario. It seems possible that the recent spread of the bobcat may have the effect of eventually extirpating the lynx northward. This seems particularly true in the Rainy River region west of Lake Superior ( see Fluctuations in Populations) .

I

\

' ' .... .....

FIG. 10. The retreat of the woodland caribou from its original range.

Gulo luscus, WOLVERINE The history of the southern distributional limits of the wolverine in Ontario is shrouded in mystery. Seton (1927) could find no authentic record south of Lake Nipissing. The only evidence yet available for this area is a mounted specimen in the Royal Ontario Museum which is reported to have been taken in Grey County in 1890. Cahn (1937) estimated that it disappeared from the Ontario Provincial Park region about 1900. It has become exceedingly rare throughout the province with the only known occurrence restricted to the northwestern section of the Patricia portion of Kenora; less than a dozen pelts per year have reached the fur trade from that area since 1928 although a few are used locally. OTHER SPECIES

Several other species, particularly the larger mammals, have retreated from the more thickly populated areas along the southern portions of Ontario. In 55

some cases distributions have been reduced to isolated occurrences, making it difficult to actually fix the present normal range. Some of the species that obviously have decreased in numbers and/or range along their southern occurrences in the province are: Lepus americanus, Castor canadensis, Erethizan dorsatum, Canis lupus, Ursus americanus, Martes americana, Martes pennanti, Lutra canadensis, Lynx rufus, and Odocoiles virginianus. In the case of the badger (Taxidea taxus) there are two small isolated "remnant populations," one in southern Ontario north of Lake Erie (Bartlett, 1955) and one in western Rainy River District. It is quite likely that the badger was never present in a much wider area, and it is yet unknown whether the effects of human habitations are reducing their range still further or perhaps are having the opposite effect in opening up the area further and improving the badger's chance for survival here. FLUCTUATIONS IN POPULATIONS

Although all animals are subject to constant fluctuations in relative abundance, some trends seem evident when some measures of these fluctuations are available. The classic examples of the cyclic fluctuations of the snowshoe hare (Lepus americanus) and the Canada Lynx (Lynx canadensis) are well known. In the case of the snowshoe hare, each recovery peak of abundance seems to return to something approximating its former level. The Lynx, on the other hand, has continued its cyclic periodicity, but has shown a rather constant decrease over the past thirty years as shown by the following table showing the number of pelts marketed from Ontario: TABLE

I. Recent trends in the production of Canada Lynx pelts in Ontario.

Year

1920-21 1926-27 1930-31 1934-35 1940-41 1944-45 1950-52

High

4,568 2,611 986

Low

630 871 429 215

It is quite obvious that if this trend continues the Lynx will soon be a rare mammal facing extirpation in Ontario. Perhaps, in fact, it may have already lost much of its former range, particularly along the south. A more encouraging recent trend may be cited in the case of the beaver (Castor canadensis). From 1919-20 to 1935 there was a drastic reduction from over 100,000 pelts sold to less than 9,000 in 1935. Following a three-year closure of trapping seasons the number of pelts sold increased from 16,934 in 1939-40 to over 100,000 again in 1951. The reversal in the trend follow56

ing the closed season appears to have been a direct reflection of conservation measures, particularly the establishment of the registered trap line system across the province. Having a specialized, restricted habitat and well marked signs of its occurrence, the beaver is subject to easy eradication by overtrapping. The Arctic fox (Alopex lagopus) has never been particularly abundant along the southern fringe of its distribution along the coasts of Hudson and James bays. Fur production figures show rather erratic fluctuations with less than 1,000 pelts sold since 1923 and less than 300 since 1936. Occasionally individual animals wander far inland, with wild caught animals having been taken as far south as Ritchie Township in Cochrane District and Township 1OB in Sudbury District. While the possibility remains that these two cases are the result of escaping from fur farms, no records of fur farms anywhere near these points have been found. Most other fur-bearing species here recorded have shown considerable fluctuation, but no long range trends. Fisher (Martes pennanti) and marten (Martes americana) showed a general decline during the period 1930-1945 but have recently shown upward trends. The recent status of these two species in Ontario has been recorded by de Vos ( 1954). Unfortunately, data are not available to determine the long range trends, if any, in most small mammals. Studies should be carried out to determine the long range trends in the fluctuations of all mammals. These trends may well produce a forecast of changes which ultimately may produce either increases or decreases in normal range. The activities of man in our modern civilization will play an increasingly important role in influencing the trends in changes in our fauna of the future. REFERENCES ALLIN,A. E.

1950. European hare introduced into the District of Thunder Bay, Ontario. Canad. Fld. Nat., vol. 64, no. 3, pp. 122-124. BANFIELD, A. W. F. 1954. Preliminary investigation of the Barren Ground Caribou. Canad. Wildl. Serv., Wildl. Mgt. Bull., ser. 1, no. lOA, pp. 1-79. BARTLETT, C. 0. 1955. Badgers in Kent and Elgin Counties, Ontario. Canad. Fld. Nat., vol. 69, no. 1, pp. 12-13. CAHN, A. R. 1937. The mammals of the Quetico Provincial Park of Ontario. J. Mammal., vol. 18, no. 1, pp. 19-30. CROSS, E. C. 1937a. Wolf! Wolf! Rod and Gun in Can., vol. 38, no. 8, pp. 18-19; 32-33. 1937b. The white-tailed deer of Ontario. Rod and Gun in Can., vol. 38, no. 9, pp. 14-15; 32. DE Vos, A. 1954. Ecology and management of fisher and marten in Ontario. Tech. Bull. Ont. Dept. Lands and Forests, pp. 1-90. 57

Vos, A., AND R. L. PETERSON 1951. A review of the status of woodland caribou (Rangifer caribou) in Ontario. J. Mammal., vol. 32, no. 3, pp. 329-337. DOWNING, s. C. 1946. The history of the gray fox in Ontario. Canad. Fld. Nat., vol. 60, no. 2, pp. 45-46. 1948. A provisional check-list of the mammals of Ontario. Misc. Pub. R. Ont. Mus. Zool., no. 2, pp. 1-11. DOWNING, S. C., AND H. LUMSDEN The nutria in Ontario. (MS) DE

FLEMING, J. H.

1913. Mammals. in The natural history of the Toronto Region, Ontario, Canada. Canad. Inst., Toronto, pp. 206-211. HODGSON, ROBERT G. 1949. Farming nutria for profit. Fur Trade J. Can., Toronto, pp. 1-110. LLOYD, HOYES

1925. The acclimatization of the fox squirrel at Pelee Island, Ontario. Canad. Fld. Nat., vol. 32, no. 6, p. 138. MILLER, G. s. 1897. Notes on the mammals of Ontario. Proc. Bost. Soc. Nat. Hist., vol. 28, no. 1, pp. 1-44. PETERSON, R. L. 1953. Notes on the eastern distribution of Eutamias minimus. Contr. R. Ont. Mus. Zool. Palaeont., no. 37, pp. 1-4. 1955. North American Moose. Univ. Toronto Press, pp. 1-280. PETERSON, R. L., ANDS. C. DOWNING 1952. Notes on the bobcats (Lynx rufus) of eastern North America with the description of a new race. Contr. R. Ont. Mus. Zool. Palaeont., no. 33, pp. 1-23. 1956. Distributional records of the opossum in Ontario. J. Mammal., vol. 37,no.3,pp.431-435. PETERSON, R. L., AND J. K. REYNOLDS 1954. Taxonomic status of the European hare in Ontario. Contr. R. Ont. Mus. Zool. Palaeont., no. 38, pp. 1-7. PETERSON, R. L., R. 0. STANDFIELD, E. H. McEWEN, AND A. C. BROOKS 1953. Early records of the red and the gray fox in Ontario. J. Mammal., vol. 34, no. 1, pp. 126-127. REYNOLDS, J. K. 1955. Distribution and populations of the European hare in southern Ontario. Canad. Fld. Nat., vol. 69, no. 1, pp. 14-20. ROBERTS, THOMAS S.

1945. The vanished mammals of Minnesota. in The Mammals of Minnesota by Swanson, G., T. Surber and T. S. Roberts. Minn. Dep. Cons. Tech. Bull., no. 2, pp. 7-19.

SETON, E.T.

1927. Lives of game animals. 4 vols., Doubleday Doran, Garden City, N. Y.

TYRRELL, J.B.

1913. Hudson Bay exploring expedition 1912. Ont. Dept. Mines Ann. Rept., vol. 22, no. 1, 161-209.

58

CHANGES IN THE CLIMATE OF ONTARIO BY

M. K. THOMAS*

1. INTRODUCTION ONTARIO, a province larger than most countries of the world, has climates which vary widely from one season to the next. In winter, northern Ontario frequently experiences periods which are as bitterly cold as any that truly Arctic climates can offer, while in summer, periods of hot, humid, tropical weather can and do occur in southern Ontario. With such contrasts between winter and summer across the province, it might seem that any climatic trends or fluctuations would be largely masked. It will be shown, however, that there have been changes in the climate of Ontario, large changes or variations from prehistoric times scaling down to smaller changes or fluctuations during the past century. There are several possible causes of these variations and fluctuationschanges in the earth's crust, variations of solar activity, increased amounts of carbon dioxide in the atmosphere are, perhaps, the most important of these. Because of the complicated and complex processes of weather and climate there are reasonable physical arguments to support most alleged causes. Whatever the causes people in all walks of life have become interested in our climatic changes. In this study no attempt has been made to assess the worth of the various theories of climatic change. Rather, the pertinent facts have been gathered together to show the changes that have actually taken place in the climate of Ontario. 2. EARLY CLIMATES Pre-Pleistocene

Before examining the instrumental record of the weather during the past century, it might be well to describe some of the early pre-historic climates of Ontario. Geologists, through the study of plant fossils, tell us that our climate has alternated over the millions of years between warm or "genial" climates and cold or "glacial" climates. During the warm periods, which have prevailed 90 per cent of the time, average temperatures were probably several degrees above the Ontario summer temperatures of today. Salt beds were laid down in the dry, warm periods, and during the warm, moist periods luxurious vegetation flourished which is responsible for today's oil and coal fields. Glacial climates, when glaciers or ice sheets thousands of feet in depth *Published with the permission of the Director, Meteorological Branch, Dept. of Transport, Toronto. 59

covered large sections of the earth, including Ontario, have recurred · at intervals of approximately 250 million years.

Pleistocene During the present era we are still in the most recent ice age, the Pleistocene. Geologists have evidence that there have been four major glacial expansions on a world-wide scale during this age. No one can say, however, if we are only in a slightly warmer interval, or if our climate will steadily warm to a "genial" climate some millions of years hence. During the more active phases of the Pleistocene period, water and ice erosion and sediment deposits shaped much of today's surface physical geography of Ontario. The Wisconsin glaciation was the most recent in the Pleistocene and its ice fields retreated from parts of Ontario less than 10 thousand years ago. Dillon (1956) estimates that at the maximum of this Wisconsin glaciation, summer temperatures over the Ontario ice fields may have been as high as the low 40's and that the average annual precipitation must have been less than 10 inches. Post Glacial In contrast to the pre-Pleistocene climates where bed rock fossils are the identifiers, post glacial climates may be studied by the analysis of pollen deposited in bogs, by varve analysis in sedimentary deposits, and finally by dendrochronology or tree ring analysis. A limited number of such investigations in this part of the continent indicate that as the climate slowly warmed, a tundra vegetation and then the various forest zones moved northward over Ontario. In a relatively short time, between 4000 and 1000 B.c., a "climatic optimum" was reached when southern Ontario was probably covt:red with a rich deciduous forest and northern Ontario with a spruce-fir forest. The presence of this forest cover suggests to Hare (1951) that mid-summer temperatures in Ontario at that time were probably 4 degrees higher than at present and that the annual precipitation was also higher than today's. Following the warm, humid, climatic optimum, conditions became much drier and drought resistant vegetation moved into Ontario from the southwest. About 500 B.c. there was a rapid and drastic change to a wetter and cooler climate. This change is well documented in Europe and there is vegetation evidence to suggest that it took place in this region also. In contrast to this remarkable change the climatic variations during the Christian Era should be classed as minor. Historical There was a warm, dry period in the early Christian centuries when a Mayan civilization flourished in what is now a Central American jungle, and when the Norse established Greenland colonies. But then in the 13th century the climate swung back towards colder conditions in the medieval wet 60

period. These changes, which probably affected Ontario to a certain degree, were, however, slight in comparison to some of the early changes-in fact, Miller ( 195 3) points out, "There are no records of weather after the Christian Era which might not apply to a single, perhaps abnormal, year of the present time." Reports and writings of early explorers, missionaries, and soldiers in Ontario indicate that the weather in the 17th and 18th centuries was much the same as it is today. Writings of Champlain in 1615 and by the Jesuits at Fort Ste. Marie near Midland between 1639 and 1649 mention everyday weather occurrences that might appear in a 20th century diary. To the best of our knowledge there were no instrumental weather records maintained by the French during this period. One of the first meteorological articles (Stirling, 1763) published concerning Ontario appeared in the Philosophical Transactions of 1763 and discusses a particularly dark day at Fort Detroit when smoke (apparently) from forest fires caused almost total darkness all day. Settlement of what is now the province of Ontario did not begin appreciably until the last decade of the 18th century. There are no instrumental records now available that date back to that decade but we have learned a few climatic facts from diaries. For example, while there were no excessively cold winters in the first quarter of the 19th century in Ontario, there were some extremely cool summers. In particular, 1816 is marked as a year almost without a summer. 3.

ONTARIO WEATHER RECORDS

History The instrumental record of weather observations in Ontario dates from 1840 when the British Government established a Magnetic and Meteorological Observatory at Toronto. There were some earlier records kept by individuals, such as Rev. Dade at Toronto, but the details of the observations are not known and the continuity of the records was not maintained. At the Observatory in Toronto, however, weather observations have been taken every day for well over 100 years at almost the same location. These records constitute one of the longest continuous series of observations on the continent of North America. During the decades of the 1840's and 1850's weather observations were undertaken by private citizens at Stratford, St. Mary's, Hamilton, and Montreal. Data from these stations were published in the Canadian Journal but these stations closed after a few years. During the 1860's an ambitious plan to have all Upper Canada Grammar Schools become meteorological observing stations failed, although a few years of incomplete records were obtained from such places as Barrie, Belleville, Goderich, Pembroke, Simcoe, Peterborough, and Windsor. 61

The real impetus for establishing and maintaining weather reporting stations was the establishment by the Federal Government of the Meteorological Service of Canada in 1871. During the following decade small cooperative observing stations were set up in many cities, towns, and rural areas of Ontario. Continuous records have been maintained at a few of these locations until the present day. Also, during the 1870's the Meteorological Service established a synoptic weather observing and reporting network to supply daily telegraphic weather messages for forecasting purposes. Weather data from a few of these early stations-Kingston, Parry Sound, and Southampton-are also continuous to the present time, although most of these stations now function solely as climatological stations. In 1873 the Meteorological Service began to publish an Annual Report which contained detailed climatological data from all observing stations. Such data sources are available for each year up to the present time although since 1916 the climatological data have appeared in a separate publication called the Monthly Record of Meteorological Observations. Over the years the number of stations in Ontario has grown steadily, by 1956 they number 325. With so many observing stations and the growing feeling of the value of continuous, homogeneous observations, tomorrow's student of climatic change should have abundant data at hand. Homogeneity When searching climatological data for evidence of climatic change, serious attention should be given to the homogeneity of the record. Many apparent climatic changes are often traced back to a change in the methods of observing, to a change in the instruments used, or a relocation of the instrument site. The Meteorological Service attempts to meet these problems by standardizing equipment and methods and by making each observing site as representative of the surrounding country as possible. Mitchell ( 1953) in his study of homogeneity has stated that mean temperature changes of as much as rn°F. may be caused by minor site relocations, and as much as 1° by such factors as local growth of foliage around shelters, change in thermometer calibration, or changes in shelter design. Changes of up to 2 ° may be caused by local building and pavement construction. While artificial temperature changes may be either "up" or "down," the usual break or discontinuity in precipitation records is caused by a poor exposure giving a smaller non-representative precipitation "catch." These apparent precipitation changes are usually brought about by leaking, faulty equipment, or on account of the shielding effect of new buildings or vegetation of recent growth. The data used in this study have been examined rather closely for homogeneity although rigorous checks have not been applied. The obvious non-homogeneous data have not been used; doubtful data have likewise been put aside. It is possible that for individual stations, errors of as high as 1 to 2°F. in temperature may be inherent. 62

City Effect

In urban areas there is a very real effect which has raised temperatures in a few of the larger cities perhaps 1° in the past century. This is called the "city effect" and is caused both by the increase in artificially produced heat and by the increase in city industrial atmospheric pollution which acts as a blanket over the city. This effect is most noticeable on winter nights but it does apply in general to all seasons of the year. Besides directly affecting the temperature records the city effect also tends to lower snowfall values. Since much of our snowfall takes place at temperatures close to the freezing point, a slight increase in temperature over the city will cause snow to melt to rain before striking the ground. 4.

TEMPERATURE

Regime Of all the meteorological elements, temperature is, perhaps, the most convenient to use in climatic trend studies. Temperatures are observed in a uniform manner in the macroclimatic layer inside a thermometer shelter at a height of four feet above a level, grassy surface and are usually representative over an appreciable area. Also people are primarily interested in temperature and so perhaps find it easier to understand than any of the other elements. Consequently, more climatic trend work has been based on temperature than on any other element. In Ontario, temperatures have been officially recorded as high as 109°F. and as low as - 73°F. During the summer, mean temperatures in the order of 70° are expected in southern Ontario and near 60° in the northern limits of settlement. In winter, when average temperatures of zero or lower are expected over most of northern Ontario, southern temperatures are usually above 20°. Values of mean annual temperature, that is the average temperature over the twelve months of the year, are not very meaningful but through their use, the relative warmth of various years may be compared rather simply. Regional

In a recent temperature change survey, Longley ( 1953, 1954) combined the annual temperature records from several stations in each region of Canada and plotted ten-year moving means of annual temperature. Figure 1, which is after Longley, shows the moving means for the Ontario districts brought up to 1955. The values for each district are based on the records of three or more stations and are credited to the final year in each decade. The moving means were obtained by averaging the first ten years of record, then adding in the eleventh year while subtracting the first year and obtaining a new average and so on until the final decade is reached. The curves for all districts indicate a cold period in the decade of the 1880's, which was common not only across Canada but throughout much of 63

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the Northern Hemisphere. Warming was rapid in Ontario in the 1890's, and in southern Ontario only minor fluctuations were apparent for the following thirty years. In northeastern Ontario, after the first decade of the century, a decreasing trend was established that lasted until the decades ending in the 1940's. During the past 15 years the annual temperature trend has been upwards in the northeastern part of the province but the level presently reached is no higher than that of 50 to 60 years ago. Northwestern Ontario ( included with Manitoba) shows decreased temperature values during the past decade. This is in common with a slightly decreasing trend throughout all of Western Canada which began during the late 1940's. In southern Ontario, the three districts show high temperatures recorded in the decades ending in the late 1930's. Following lower values for a few years the increasing trend took over and at the end of 1955 the ten-year moving means are at a maximum in southern Ontario. The current year of 1956 has been a relatively normal year and speculation is aroused as to whether or not the warming up trend has been halted for the time being. Seasonal Since the annual temperature has increased throughout the province it is now of interest to examine the different seasonal temperatures. Fig. 2 shows moving means of temperature for each season at Toronto. Winter is taken as December of the previous year, January and February; Spring consists of March, April and May; etc. It can be seen that the trend has been upward in all seasons. Winter, although the most variable has not warmed up much more over the past 100 years than summer, which has fluctuated the least. Features of these curves are the succession of very hot summers in the 1930's and warm winters in both the 1930's and again in the 1950's. As a result of an exceptionally cool spring in 1956 the short-term trend in that season is towards cooler temperatures. Over all, the annual curve shows an increasing trend during the most recent years, similar to earlier temperature increases which occurred in the 1860's, the 1890's and the 1930's. Toronto temperatures have, of course, undergone an additional city-effect increase as described in the section above. The writer has estimated that this amounts to, perhaps, 1 ° per century. Without taking this into account the rates of increase per century are shown in Table 1. Deducting the probable city effect the rates of increase are shown in the neighbourhood of 3 ° per century for all seasons in the Toronto area. It should be noted that the Toronto temperatures used here have been reduced to a common standard. In the past, the mean temperature for a day has been obtained by various methods such as averaging temperatures at certain fixed hours, averaging the 24-hourly temperatures, or by averaging the daily maximum and minimum temperatures. These Toronto data have been adjusted to conform with the present system of obtaining the average of the 24-hourly observations. Temperature data from other Ontario stations, mentioned below, have been obtained by averaging the daily maximum and 65

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