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BULLETIN NO. 94

Freshwater Vertdbrates of the and Subarctic

BY

V. C. WYNNE-EDWAHDS

Aberdeen University, Scotland

PUBLISHED BY THE JirISHERIES RESEARCH BOARD OF UNDER THE CONTROL OF THE HON. THE MIN1[STER OF FISHERIES

"I TA.WA, 1952

'i�

L�JI ____ �� ______� BULLETIN NO. 94

Freshwater Vertebrates of the Arctic and Subarctic

BY

V. C. \VYNNE-EDWARDS Aberdeen Flliversity, Scotland

PUBLISHED BY THE FISHERIES RESEARCH BOARD OF CANADA UNDER THE CONTROL OF THE HON. THE MINISTER OF FISHERIES

OTT A \VA, 19.52 Printed in Canada by University of Toronto Press for the Fisheries Research Board of Canada CONTENTS

INTRODUCTION 1 PHYSICAL CONDITIONS 2 THE FRESHWATER VERTEBRATE FAUNA 4 (A) Lampreys and 5 (i ) Petromyzontidae- Lampreys 5 (ii) ACipenseridae- Sturgeons 5 (iii) - Salmon, trout and char 6 Salmon (Salmo and Oncorhynchus) 6 Trout and char (Salvelinus, CTistivomer, H ucho and Brachymystax ) 7 (iv) Coregonidae- Whitefish, ciscoes, etc. 9 Inconnu and nelma () 9 Tullibee, "herrings" and ciscoes (Leucichthys ) 10 Whitefish () 12 Round whitefish () 16 (v ) Thymallidae- Graylings 16 ( vi) Osmeridae- Smelts 17 ( vii ) Catostomidae- Suckers 18 ( viii ) -Minnows 18 (ix ) U mbridae- Alaska blackfish, etc. 19 (x) Esocidae- Pike 19 (xi) Percopsidae- Trout-perch 20 (xii) Percidae- Perch, pike-perch 20 (xiii ) Cottidae- Sculpins or bullheads 21 (xiv ) Gasterosteidae- Sticklebacks 22 (xv) -Ling, burbot or methy 23

( B) Amphibia 24 Northern wood frog (Rana Sylvatica cantabrigensis) 24 Northern tree-frog (PseudacTis nigrita septentTionalis ) 25 (C) lvIammalia 25 White whale or beluga (Delphinapterus leucas) 25 (Phoca hispida) 25 Harbour seal (Phoca vitlllina ) 26 CONCLUSION 26

REFERENCES 26 INTR ODU CTIO N

THE MAJORITY of vertebrates dealt with here are fishes. A few amphibia, but no aquatic reptiles, extend into subarctic ; and of aquatic , two species of seal and one cetacean Delphinapterus occur in arctic fresh waters. Semi-aquatic mammals, for example beaver Castor (circumpolar), muskrat Fiber (nearctic), water-rat Arvicola (palaearctic) , and otter Dutra (circum­ polar) , are excluded, together with numerous aquatic , many of them typically boreal and arctic, for example, loons (Gaviidae) , geese, swans and many ducks (Anatidae ). Roughly one-seventh of . the land area of the world drains into the Arctic and adjacent ice-bearing , and many of the larger rivers have their sources far to the south in the temperate belt of , and . Five of them are two thousand miles or more in length, namely the (2,700), Yenisei (3,300) , (2,800 ), Yukon (2,000) and Mackenzie (2,525); and in summer they carry collectively an incalculable volume of relatively warm, usually mud-laden water to or beyond the Arctic circle, forming nahll'al channels for the dispersal of the temperate-zone continental aquatic fauna and flora into high latitudes. Within the area of Pleistocene glaciation in North America and Europe there is also a series of great lakes, the more northerly including Athabaska, Great Slave and Great Bear Lakes in the Mackenzie system, Nettilling and Amadjuak in , Ladoga and Onega in Russian Karelia; and because of impedi­ ments to drainage by glacial drift and permafrost, the frequency of smaller lakes and ponds is without parallel in other parts of the world. Parts of these freshwater systems either were covered by continental ice at some stage of the Pleistocene, so that their pre-glacial fauna was exterminated, or endured a climate so severe that few or no freshwater vertebrates survived. The post-glacial period, up to the present day, has been characterized, therefore, by recolonization of surprising speed and extent; and realization of this confers upon the study of the distribution of the freshwater fauna a deeper significance. Recolonization has, of course, taken place from refugia little or not at all affectedby glaciation. In North America, for example, the Mississippi basin is the most important; and in ,Asia several of the great freshwater systems, including Lake Baikal on the Angara-Yenisei, and probably the Ob, Lena and others, eVidently sustain an important part of their pre-glacial faunas. The spread of fishesfrom one river system to another has occurred in two ways : either through the ' to enter the new river by its mouth; or through the headwaters,. where changes in the interior drainage, most commonly resulting from temporary block­ age by continental ice or glacial drift, have diverted the waters from one system into another.

1 The first method is generally possible only to fishes able to live either in the sea or in fresh water, that is "euryhaline" species; it appears to be the most frequent method of recolonization, and it has the result that these fishes are usually debarred from the upper waters of a system because they cannot ascend impassable waterfalls. The second method has been important where transitory dams of ice or boulder-clay have formed lakes draining now into one system, now into another, as has occurred so commonly in North America. The resulting alternation of isolation and communication is certainly connected with the origin of species­ s\v,umsin Salvelinus, Coregonus and Leucichthys (see below ) in some American and European lakes. As an agency of recolonization it has been successful to an unexpected degree; for example, it may have permitted the little trout-perch PCJ'copsis to pass, from the Great Lakes- �lississippi refugium, far to the north and west via the low-lying into the Mackenzie, and thence to its mouth, and over the (lowest) divide into the Yukon, where it was discovered by us in the Porcupine River at Old Crow, Y. T., in 1945. Percopsis may pOSSibly have had a second refugium during the Pleistocene in the far north-west, together with the only cyprinid now known in the Yukon, COHesius plumbeus, and several other , including the lake-trout Cristivomer, the round whitefish Prosopiurn cylindracewn, the northern sucker Catostornus catostomus, the black­ fishDallia, pike Esox lucius and burbot Lota lota. The northwestern distribution of Percopsis is restricted to a very narrow "route" which rather indicates present recolonization; in any case, however, whether this and the other species survived in one or in two isolated refugia, separated by two thousand miles, the repopu­ hUlon of the glaciated area has taken place with surprising rapidity. The most important families of arctic and subarctic freshwater fish contain many euryhaline species, for example, the sturgeons (Acipenser), salmon, trout and char (Salmonidae ), whitefish (Coregonidae ), inconnu (Stenodus ), smelts (Osmeridac ), sticklebacks (Gasterosteidae ), and sculpins (Cottidae ). The sea­ going SaZmo and Oncorhynchus, which enter fresh water only to , have distributions centred on particular , and may be divided into Atlantic and Pacific series; whereas the more strictly freshwater coregonines and others, which feed and grow freely in fresh water, belong rather to continental series, either palaearctic or nearctic as the case may be. It is interesting to find in the \!ackenzie and Yukon that, whereas the headwater colonists al=e of course American or nearctic types, several of the euryhaline al1adromous colonists are essentially palaearctic, including the inconnu and two or three coregonids.

PHYSICAL CONDITIONS All the peculiar features of arctic freshwater habitats may be attributed to the effects of low temperature. Few or no cold-blooded vertebrates can survive the arctic winter except in fluid water; and the prevailing low summer temperature tends to retard growth and development. This retardation is to some extent compensated by adaptation, permitting faster growth at low temperature: thus Miller (1947, pp. 39-41 ) found little difference between the growth-rate of pike Esox lucius in Great Bear Lake and its growth-rate in Lesser Slave Lake

2 FIGUHE 1. The Arctic and Subarctic regions. An approximate southern boundary of the'lrea is indicated by the Northern Limit of Cereals (shown here by the fine dotted line; after Larousse's Atlas, Hachette, Paris). 10° further south; the same appeared to be true of whitensh. Coregonus and tullibee Leucichthys, but not of lake trout Cristivomer. The physical effects of temperature are somewhat different in running-water and standing-water environments, and these are treated separately. (a ) Several of the great rivers, as mentioned already, gather much of their water in temperate latitudes between 50° and 600N., where annual precipitation is higher than it is in the Arctic. In summer the upper waters are relatively warm, and during the two to four weeks required to carry these warm tributary waters down to the they retain their heat to an important extent. The Mackenzie, for example, may be inferred to have summer maxima in the main channel of the delta at 68°N. of 15° to 18°C., whereas in large lakes in the same latitude the temperature of the open water may never exceed 10°e. at the surface. Nansen (1914, p. 164 ) records a water temperature of 9.9°e. as late as September 7 in the Yenisei at 67°43'N. The great rivers are all characterized by extreme muddiness or turbidity, with the exception of about one hundred miles of the Mackenzie immediately below its outlet from Great Slave Lake, which serves as a mud-trap, and the Angara which similarly discharges the waters of Lake Baikal (Nansen, 1914, p . .305) . Visibility in the water is commonly less than 0.5 m., and this makes the habitat unsuitable for grayling Thymallus and freshwater trout Salt;elinus etc., which by habit take much insect food from the surface. The rivers are frequently several miles wide, shallow, and full of bars and alluvial islands. They also re­ semble one another in having remarkably even pronles, without falls or serious rapids to form barriers to shipping or migratory fish in their lower 1,000 to 1,500 miles. The normal speed of flow seems to be about 3 miles per hour (5 km. /hr. ) . (The Yukon, however, is much faster, averaging more like 5-7 m.p.h., and the effort required for anadromous migration is correspondingly much greater.) Complete vertical mixing is achieved by slow majestic swirls and eddies, which mark the river's steady flow; though confluent waters, like the clear Mackenzie and the muddy Liard, may keep to their respective banks for scores of miles. The rivers are actually closed by ice for at least six months, and remain near O°e. and carry floating ice for several weeks longer, extending the winter period for the river fauna from late October to the beginning of June. Some of the smaller rivers in the may in fact be frozen from October to May in­ clusive (Preble, 1908, p. 49 ). In the T\Iackenzie and Yenisei the spring break-up begins in the upper tributaries, and proceeds downsti"eam with the Roods as much as 100 miles a day. Ice-jams may cause the water level suddenly to rise 5 or 10 m., backing the current upstream for many miles in the tributaries. A powerful increase in the scour of the river is reinforced by surging blocks of ice, reared up together in wild confusion. Several descriptions have been given of the spectacle, but none more vivid than that of Henry Seebohm (1901, pp. 325- 333 ) at Turukhansk in 1877; it cannot be otherwise than a primary limiting factor in river ecology. In the upper Yukon, and no doubt some other rivers, the spring high water comes after the break-up, since the principal sources are in high snowy mountains. In the permafrost of Transbaikalia, where exceSSively low winter tem- peratures prevail, How is said to cease almost entirely in certain rivers, which may in many places freeze from top to bottom (Nansen, 1914, p. 313). Bottom ice, to be seen, for example, in the Lachine rapids at Montreal, sheathing sub­ merged boulders and stones where the swift current prevents the surface from freezing, must have a destructive effect on the bottom fauna. It is well known at Bear River rapids between Fort Norman and Great Bear Lake (Preble, 1908, p. 46), and is probably a characteristic feature of places where the water remains open all winter. ( b ) The larger lakes are essentially oligotrophic, without thermal stratification and with oxygen contents near saturation at all depths down to 300 m. This is true of Great Bear Lake, in which plankton production is very meagre, Secchi's disc is visible to a maximum depth of 29 m. anel predaceous fish such as lake trout (Cristivomer) mllst take a significant part of their food in the form of ter­ restrial insects at the surface (details from Miller, 1947 ). Many subarctic lakes are of Forel's "temperate" type, in which the maximum surface temperature exceeds 4°C., for example, Great Slave, Baikal, Onega, Ladoga. Great Bear is on the borderline, since in the open water the temperature of 4 C is but little exceeded, though locally in bays anel shallows 15°C. or higher may be attained in hot weather. The bottom fauna and fish are naturally concentrated in summer at these points. Most tundra lakes are probably of the "polar" type, in which the surface neve I exceeds the temperature of maximum density. Soper (1928, p. 16) found Nettil­ ling Lake still filled with ice in early August; and highly polar lakes barely thaw before the ice reforms, so that there can be little or no circulation at any time. In all very large lakes open water occurs at temporary contraction cracks throughout the winter, though the ice may be 2 m. thick, and this permits the survival of air-breathing seals in Baikal, Nettilling, etc. (see pp. 25-26). Precipitation exceeds evaporation throughout the Arctic regions, anel there are consequently no inland drainage systems nor saline lakes. Small and large stagnant ponds (cf. Johansen, 1922) are characteristic of permafrost country, and in many areas they dot the tundra and plains in countless numbers. They are seldom habitable by fish unless they have outlets; and they may freeze to the bottom unless the depth exceeds 2 In. The Alaska blackfish (see p. 19 ) lives in this habitat, as well as in streams, and is widely believed to be able to survive imprisonment in ice, though on circumstantial evidence only. In some of the thousands of ponds in the Mackenzie delta there are whitefish (Coregonus ) and other fish, completely isolated except during a brief period at the height of spring floods, when the delta channels overflow their banks; some, indeed, may be isolated for several years at a time.

THE FRESHWA TER VERTEBRATE FAUN A The four classes of Vertebrata dealt with in this section are the Lampreys ( Cyclostomata), Bony Fishes (Osteichthyes), Amphibia and Mammalia. In the latter class, only the white whale and two species of hair seal are included, all

4 three being in fact i10rmally marine mammals, only exceptionally occurring in arctic or subarctic fresh waters. There are no "cartilaginous fishes" (Chondrich­ thes ) nor aquatic reptiles in these waters. The sequence followed is systematic.

(A). Lampreys and Fishes A number of families of freshwater fishes are highly typical of, though not wholly confined to, the arctic, subarctic and alpine regions. In particular, the related Salmonidae and Coregonidae together contain by far the largest number, and the most valuable, of northern freshwater fish, including salmon, trout. whitefishand tullibee, and absolutely predominate in arctic fresh waters. (i) PETROMYZONTIDAE-LAMPREYS Lampetm (Entosphenus) faponica is known throughout the arctic mainland from the eastward across Asia to the Yukon and �Iackenzie. It was described from the Yukon by Bean as Ammocoetes auI'CUS, and occurs at least as far up as Dawson, where we took a larva in 194.5; Richardson (1836) p. 294) found one adhering to an inconnu in Great Slave Lake ( "Petl'omyzon fluviatilis" ). Several "subspecies" are recognized by Berg (1932, p. 30; see also his distribution map, p. 789), including kesslel'i in the Dvina, Pechora and Ob, and septcntl'ionalis, recorded by him likewise from the Pechora and Ob and thence eastward to the Anadyr, but not in Baikal. Like the European L. fluviatilis, which they closely resemble, they descend to the sea, returning to spawn. No other lampreys are knoWll to occur in the Arctic. Nansen (1914, p. 137 ) says they are used as bait for sturgeon on the Yenisei. In the delta and lower part of the Yukon they run up from the sea in early spring, and are, or were formerly, taken by the Indians with dip-nets, through holes cut in the ice, for use as dog food; they are said to be exceedingly oily (Evermann and Goldsborollgh, 1907, p. 227). (ii ) ACIPENSERIDAE-STURGEONS Some types of sturgeon spend part of their lives in the sea, and enter rivers to spawn: these are therefore euryhaline anadromous migrants; others complete their whole life-hiS'tory in fresh water. One of the former, Acipcnscl' baeri, is or has been of considerable economic importance in the Ob (including especially its tributary the Irtish) and Yenisei. It ascends the rivers from the sea as soon as the ice is out in the spring, though some are caught through the ice in winter in the mouth of the Yenisei at 71 0 ll'N.

(Nansen, 1914, p. 107 ); it attains a weight of 100 kg., and exceptionally 200 kg .. (220 and 440 lb. ) . The smaller "sterlet", A. mthcntls, attaining about 1 m. and usually 6 to 6)� kg. (13-14 lb.). is a non-migratory species in the Ob and Yenisei; and forms re­ garded by Berg (1932, p. 59) as hybrids between this and A. baeri are found in the same rivers, and in Lake Baikal, the Lena and . In modern times the sterlet has also penetrated the Russian canals and become established in the Dvina and Neva systems, coming originally from the Volga, in which it is indigenous also.

.5 ' Sturgeons are not native to any arctic rivers within the' area....of Pleistocene glaciation in the : that is, west of the Urals; nor do they occur in the Anadyr, Yukon, Mackenzie or rivers eastward until we come to the Churchill and Nelson systems in and . Their absence from this quadrant of the Arctic (roughly l60°E. to 1l0o W.) can certainly be ascribed to lack of opportunity for post-glacial colonization, since there is little doubt that the rivers in question are as suitable for sturgeons as the Dvina has proved itself to be. (It should be noted here in parenthesis that the Pacific sturgeon A. trans­ montanus is found on both shores of the temperate North Pacific, north to the Copper River in southern Alaska (Everman and Goldsborough, 1907, p. 231).) The type found in subarctic Canada is another species allied to the sterlet, the rock sturgeon A. fulvescens; this over most of its range does not migrate to the sea. It occurs in the rivers entering the west side of Hudson and James Bay from the Churchill southward, in Manitoba and ; and perhaps also on the side. It has apparently come as a "headwater colonist" from the Great Lakes system.

(iii) SALMONIDAE-S ALMON, TROUT, AND CHAR

SALMON (Salmo and Oncorhynchus) The salmon are normally sea-run fish, entering rivers only to spawn and fasting while in them. There are two genera, Salmo and Oncorhynchus, the latter, com­ prising the "Pacific salmon", being found only in the North Pacific region of America and Asia. Salmo, on the other hand, if we include the steelheads and cut-throat trout (S. gairdneri and S. clarki ) of the American Pacific slope and the related cut-throat type trout of Kamchatka (S. mykiss ), occurs both in the North Pacificand North Atlantic regions. Salmon are of little importance in truly arctic waters, other than the Yukon and Anadyr. S. salar, the Atlantic salmon, extends northeastward in greatly diminished numbers to the mouth of the Pechora, and northwestward to south Greenland, Labrador, and (Leaf River) in which it occurs only occasionally, according to the latest information provided by M. J. Dunbar in litt., and Legendre and Rousseau (1949, p. 135 ). It is abundant in Newfoundland, Iceland and , including the Baltic. , S. trutta, the , both sea-run and land-locked, is present in east to the Urals. S. gairdneri, comprising the steelhead and rainbow group, occurs in southern Alaska, and enters the Yukon Territory at one point by the Alsek River and its tributaries. Of the six species of Oncorhynchus two are known to occur north of , though in numbers too small to be of economic importance. These are the humpback or pink salmon, O. gorbuscha, which has been taken fr0111 the mouths of the Mackenzie, Colville, Kolyma, Indigirka and Lena rivers; and the dog salmon, O. keta, from the Mackenzie, Kolyma and Lena: Dymond and Vladykov (1933, p. 3743) believed that an imperfect specimen of Oncorhynchus, taken in Great Bear Lake and examined by them, belonged to this; species. 6 In the Yukon and Anadyr five species are found, including the two just named, the very fine king or spring salmon O. tshmcytscha, the coho O. kisutch, and the sockeye or red salmon O. nerka. The Yukon is famous for its king salmon, which occasionally reach 100 lb. (45 kg.) or more in weight. This species penetrates to the uppermost tributaries of the river, including the Lewes, Teslin, vVhite-Donjek­ Kluane, and Porcupine; and in streams entering l\1arsh Lake above 'Whitehorse, Y.T., and at Atlin, E.c., king salmon reach the astounding distance of 1,800 miles from the sea. To achieve this they must svvim against the rapid Yukon current and travel, at the most conservative estimate, 6,000 miles through the water, entirely without food- a migration without parallel among fishes. The whole journey takes about twelve weeks, and of course none survives to return to the sea. King salmon are present in shallow coastal waters from January on. The run starts early in June in Norton Sound, reaches Dawson about June 28, and Selkirk about July 10. Eegular fisheries are conducted as far up as Dawson, employing 6- to 8-inch (15-20 cm.) gill-nets set in eddies, and also automatic "fish-wheels" operated by the river current. The salmon are fit for human food for as long as (me month after entering fresh water, after which the silver colour gives place to reel and the fleshloses its flavour. The dog salmon also ascends the Yukon above Dawson, and its tributary the White River as far as Kluane Lake. Its colour and flesh deteriorate almost im­ mediately on entry into fresh water, but the fish is still suitable for dog feed. The remaining species do not go far up the river, the sockeye being more abundant than the coho and humpback. In the smaller Alaskan rivers, for example the Kubak (Kowak) , the humpback is usually the most abundant species, running up in July. For a fuller account of Alaskan salmon see Evermann and Goldsborough (1907). Several species of Salmo or Oncorhynchus have produced landlocked varieties; there are at least two of these in northwestern America north 6f the 60th parallel, namely the (S. gairdneri irideus) and the "redfish" or "kokanee" (0. nerka var.), both of which occur in the Yukon Territory in the Alsek Eiver system.

TROUT AND CHAR (Salvelinus, Cristivomer, H Heho and Brachymystax) :\1any kinds of trout, which are confined to fresh water and thereby isolated into as many separate populations as there are watersheds, have produced con­ fusing series of local varieties and subspecies. They are to be found in almost all the northern and arctic streams in which the water is clear, and in lakes. The most widely distributed of arctic freshwater fish may be grouped under the general name of Arctic char, Salvelinus alpinus. Either as a sea-run fish, commonly kno\i\fIl as "salmon" or "sea-trout" around the shores of and northward, or as landlocked fish, or both, they occur throughout the Arctic mainland and on most of the larger islands. Their range embraces Iceland, Spitz­ bergen, Britain, Scandinavia, the whole of northern Asia and North America, and both coasts of Greenland; the islands of Ellesmere, Baffin, Novaya Zemlya and New are also included.

7 The sea-run char or "salmon" enter the rivers on both shores of Hudson Bav, on the west from the Churchill northward, and thence right along the Arctic coast to the Coppermine (and Anderson ?) River; they may ascend for distances exceeding 100 miles up the Thelon, Back and no doubt other rivers, the upper limit depending on the presence of impassable falls. On the east side of Hudson Bay they extend from Great Whale River (Vladykov, 193.'3, p. 19) to Ungava Bay, northern Labrador and Greenland. They are abundant in Southamp­ ton and Baffin Islands, and recorded from Somerset Island. Manning (1942, p. 128) believes that they come in from the sea in August and September, descending again the following spring as soon as the ice is out; and he thinks it improbable that any remain in the sea during the winter. Spawning takes place beneath the ice, probably in the early months of the year. In summer these big sea-char may be very numerous in river mouths and shallow coastal waters, and are taken in gill-nets, or in favourable places by spearing, as for example below Bloody Falls near the mouth of the Coppermine. The normal weight is 5 to 10 lb. (2-4 kg. ), exceptionally 20 lb. (9 kg.); the flesh colour is variable, but usually red or pink. \lost non-migratory char are smaller and even more widely distributed. They are very variable in colour and markings. Gunther (1877a, p. 294; 1877 b, p. 476) described two "species" from northern , one of them from latitude 82)�ON. Both were small: one was found mature at 8 inches (20 cm.), and the other at 12 inches (30 cm. ), with the roes in the latter starting to develop in August. Other species have been named from Quebec (S. alpinus marstoni ) and other parts of northeastern Canada. In the northwest, from the District of :'-Iackenzie to Alaska and southward in the mountains, as well as in the Aleutians, Kamchatka and the Anadyr, the type found is the "Dolly Varden" trout, known in Hussian as the "malma" (S. a. malma ). Some of these species, including the ' last, may run out to the sea. Many of the landlocked char are remarkable for their brilliant nuptial colours. They are fond of eating salmon eggs when they can get them; they do not very readily take the angler's fly, but may often be easily caught with a small spinner or baited hook. The speckled or brook trout (S. fontinalis ) is an entirely distinct species in tempcrate and subarctic eastern North America, attaining its northern limit, so far as is known, 24 km. north of Great vVhale lEver on the east shore of Hudson Bay (Vladykov, 1933, pp. 18-19) and at the outlet of Payne Lake in northern­ most Ungava (Legendre and Rousseau, 1949, p. 134); and in the on the west shore of Hudson Bay (Hinks, 19-13, p. 90). Several other species of Salvelinus have been described from Siberia and Europe. Arctic char fresh-run from the sea are superior to the best salmon as table fish. They are readily obtained, however, only in the summer months, the season of plenty. The Old and New Worlds each have an additional kind of trout of great size, competing with the king salmon for the claim of being the largest of the Salmonidae. The nearctic fish is the lakc trout Cristivomer namaycush, well known in temperate North America, and extending into the Subarctic. It is

S chiefly, but not entirely, a lake fish, never, so far as is known, descending to the sea; and it has a northern limit somewhat as follows : Alaska and Yukon Territory, in the Yukon basin, the Kowak and some of the Pacific coast watersheds; Mac­ kenzie basin including Great Bear and Great Slave Lakes; Coppermine including Dismal Lakes (Hanbury, 1904, p. 210 ) and Fort Enterprise (Richardson, 1836, p. 182); ? Back River; Dubawnt-Thelon watershed down to Baker Lake. There is a specimen from Victoria Island in the Itoyal Ontario �1useum of Zoology (Dymond, personal communication), and another from Southampton Island (Manning, 1942a, p. 23) . It is present in the interior of U ngava, north at leasUo 600N. (Payne Lake, George and Kogaluk Rivers: Legendre and Rousseau, 1949, p. 134). Lake trout attain a great size, fairly frequently exceeding 50 lb. (22Jf kg.) and exceptionally reaching nea.rly 100 lb. (45 kg.). They are especially abundant in Great Bear and Great Slave Lakes, and in the latter have made up a substantial part of the commercial fishery devel?ped since 1945 (Rawson, 1947, p. 58) . They are voracious, to a large extent preying on other fish; but in Great Bear Lake they also take insect food at the surface. The giant Siberian trout is the "taimen", Hucho taimen, whose range extends from the Ob eastwards to the Lena, and south to the Volga, Lake Baikal and the upper Amur (Berg, 1932, p. 793) . According to Berg (ibid., p. 187 ) it attains 65 to 130 lb. (30-60 kg.), and in the Yenisei, Khatanga and Pyasina, old records go up to 177 lb. (80 kg. ), almost twice the weight of the largest lake trout or king salmon. A smaller related form is the "len ok", BrachYIHystax lenok, which has a similar range in the Arctic except that it is found also in the Kolyma. These trout do not enter the sea, and are in consequence entirely confined to unglaciated watersheds; nevertheless the taimen makes migrations upstream to spawn, be­ having like a salmon in its persistent attempts to leap \vaterfalls.

(iv) COREGONIDAE-WHITEFISH, CISCOES, ETC.

INCONNU AND NELMA (Stenodus leucichthys) The inconnu or "conny" has a particularly interesting and well known distribu­ tion. It was first recorded in North America by Samuel Hearne at Great Slave Lake in 1772, and received its name from the fact that it was "unknown" to the French-Canadian voyageurs. It is now recognized as being a race (St. leucichthys mackenzii) of the wide;ranging northern fish found in the Volga and other Caspian rivers (St. l. leucichthys), where it is called "belorybitsa" ("whitefish" ). and in the great northern rivers of the U.S.S.R. where it is known as "nelma" ( St. l. nelma). The conny stands somewhat intermediate between the Salmonidae and the typical whitefishes. Its teeth are in fact numerous, but so small and densely packed as to be inconspicuous. It is an anadromous migrant, spawning chiefly in running water probably in late summer; and those individuals inhabiting the lower .500 to 700 miles of the rivers largely return to the sea in the late fall. They migrate upstream in June to August as far as the first real rapids, and they ar� 9 therefore unknown in the Slave River above Fort Smith. Many no doubt spawn in the Mackenzie itself, including even the delta, in which yearling stages 3 cm. in length have been found by us. There is a large ,immature population, for maturity is not reached (in the nelma at least ) until the eighth year, when the fish's length is about 20 inches (50 cm. ) . Inconnu readily take the hook. Their food consists of small fish, including whitefish and lake herring, sculpins, minnows and sticklebacks; when young they t�ke aquatic insects (Rawson, 1947, p. 60 ). They do well in muddy rivers. It is the general consensus of opinion that the conny population of Great Slave Lake and its larger tributaries is independent of that in the Mackenzie proper, and that there is little or no migration between Great Slave Lake and the sea. There is probably no part of the navigable system of the river, from Fort Smith to the sea, from which connies are always absent; but they are quite uncommon in the lower Liard and between Fort Simpson and Providence rapids. They occur in the Peel, but appear to be absent from Great Bear Lake, being kept from it by the Bear River rapids; there is indeed an old record of Simpson's (1843, p. 217) of one taken at Fort Confidence, but it must be accepted with reserve, since the species was not rediscovered during the recent fisheries ex­ ploration. Inconnu form an important part of the fall fishery above Providence, and at Hay, Buffalo and Rocher Rivers, all situated around Great Slave Lake; they are taken incidentally in whitefish and river-herring nets at the Ramparts, Arctic Red Hiver and in the delta. They are not esteemed as food in summer, either here or in Siberia, being usually oily and rank, but they are palatable when small and also in winter and spring; and at all times they are acceptable as dog feed. The eastern limit in North America is in the Anderson. Probably they enter some of the larger rivers along the coast west of the Mackenzie; they are known in the Kowak (Evermann and Goldsborough, 1907, p. 236); and are generally distributed in the Yukon and its tributaries, though they are by no means as abundant as in the Mackenzie and of very little economic value. Specimens have been obtained in Teslin Lake on the 60th parallel, and inconnu are caught in small numbers in Kluane Lake; they have been verbally reported to us on less reliable authority from Mayo Lake. In the U.S.S.R. the nelma is present in all the large rivers from the White Sea eastwards, including the Dvina, Pechora, �b, Yenisei, Khatanga, Lena, Indigirka, Kolyma and Anadyr, the last, like the Yukon, entering the . The nelma ascends as soon as the ice is out, and its distribution is similarly restricted by the first serious rapids, which exclude it from Lake Baikal, standing 1,500 feet above sea level. It attains a larger size than in the Mackenzie, 50 lb. being not uncommon, and 88 lb. (40 kg.) recorded, compared with a maximum of just over 63 lb. (28.6 kg.) in the Mackenzie (Dymond, 19,13, p. 223 ). rULLIBEE, "HERRINGS" AND CISCOES (Leucichthys) The of this genus and the next is even more confusing and difficult than that of the Salmonidae. The tullibee or "freshwater herring" are in general

10 smaller species than the whitefish, with the lower jaw equal to or longer than the upper, so that the chin forms the anterior tip of the fish. It would serve no useful purpose here to treat the genus in great detail (for which see Dymond, 1943). Many freshwater herrings are principally lake fish, being rather seldom taken in rivers; but there are also a few which are essentially river fish. All are largely plankton-feeders, sifting small organisms from the water passing through their gills, by means of numerous long gill-rakers arranged in four pairs of curved combs at the sides of the gullet. Of the lake herrings, the most widespread in Canada is the "tullibee" Lcucichthys artcdi, which, in one or another of its many local forms, extends from New England and southern Quebec to Great Bear Lake (whence it is celebrated as the "Bear Lake heming"), and is said to spawn in July. In the this and similar fish are named "ciscoes". Tullibee occur in the watersheds entering the east side of Hudson Bay north to Great vVhale River and Richmond Gulf (Dymond, 1933, p. 8), and in all those entering the west side of Hudson Bay; at certain times they are found in brackish water at their mouths. They are found throughout the Thelon-Dubawnt and Back systems (Hanbury, 1904), and in all parts of the Mackenzie basin north of the 60th parallel at least. In many of the more temperate lakes to the southward two or more independent species of herring may live together in the same waters; and the one in Lake Athabaska is apparently L. zenthicus. Tullibees are not reported either from northern Ungava, Labrador and Newfoundland, or from the Yukon and Alaska. Rather similar to the hlllibee is the large river herring of the Mackenzie and lower Yukon, which probably occurs also in the intermediate rivers entering the Arctic Ocean and the mouth of the Coppermine. This is L. laurcttae, and it is probably the nearctic representative of the Siberian "omul", L. autumnalis. In the Mackenzie it has been found as far up as Camsell Bend; it forms the principal object of the summer fishery at the Ramparts, and is caught in numbers also at Arctic Red River and Aklavik. It attains a length of 12 to 18 inches (.30-45 cm.) and a weight of 3 lb. (1.4 kg.). It is migratory, being caught at Aklavik between freeze-up and Christmas presumably on its way out to the sea, and ascending again in early spring. In the Yukon it is common at the mouth, but does not come up as far as the Canadian border. It has also been recorded from the Meade, Kual'll, Nushagak and Naknek Rivers in Alaska (Evermann and Goldsborough, 1907, p. 235). In most of the same rivers there is found a smaller specics seldom attaining 12 inches (30 cm. ) and with a strongly projecting chin, formerly called L. pIlsillus, but shown by Dymond (1943, p. 208) to represent the Asiatic species L. 8(/1'­ dinella. This is also a migratory fish, of little economic value in the :Vlackenzie, partly because all but the largest individuals pass through the 3··inch nets, which are the smallest in common use in the river. Its distribution in the Mackenzie is the same as that of the previous species. In the Yukon it is abundant right up to the headwaters, and is best known at Carcross, Y.T., lying in immense schools in snmmer beneath the railroad bridge across the narrows between Lakes Bennett and Tagish, where children take it on a hook. It is the only Leucichthys certainly

11 known to occur in the interior of the Yukon Territory, and is probably the species reported from Mayo and Teslin Lakes. The last two species, or their near relatives, are very plentiful in the Siberian rivers. L. sardinella, referred to by Nansen ( 1914, p. 107) as the "seJd", has the wider distribution, from the Dvina to the Anadyr, and also in Novaya Zemlya and Kolguev Island; it is migratory, and does not reach Lake Baikal. The "omul" L. autumnalis is not found in the Dvina or Ob, according to Berg (1932), though he records it in the intermediate Pechora, and in the Yenisei including Baikal, the Lena and Kolyma. Siberia has also the "tugun" and "peled" (L. tugun and L. peled), so that four species of Leucichthys coexist in the Yenisei and Lena, as against two in the Yukon and Mackenzie. In northern and arctic Europe there are the various fish known as "vendace" in north England and adjacent parts of Scotland, as "pollan" in Ireland, "lakesild" in Norway and "smasik" in Sweden; all are forms of L. albula, and inhabit lakes, as do the tullibee. Spawning takes place in the late fall.

WHITEFISH (Coregonus) The whitefish are collectively the most valuable group of arotic fishes, and justly celebrated for their good flavour. Richardson (1836, pp. 195-6) says "though it is a rich, fat fish, instead of producing satiety it becomes daily more agreeable to the palate; and I know, from experience, that though deprived of bread and vegetables, one may live wholly on this fish formonths, or even years, without tiring." Whitefish are generally larger than lake herrings, often weighing 3 to 5 lb. (about 2 kg.); and the exceptional "jumbos" weigh very much more. Whitefish feed to a large extent on bottom organisms, including crustaceans and molluscs, rather than on plankton; the stomach is thick, and the gill-rakers correspondingly shorter than in Leucichthys. Some are inhabitants of large rivers, others of lakes. They are rather sluggish fish, but many of them are migratory and capable of entering brackish or salt water. In North America east of the Rockies, the most widespread species is Coregonu8 clupeafOTmis, the common whitefish, of which a number of subspecies have been described. The species is highly variable, and it is difficult to find characteristics of taxonomic value. Such obvious feahlres as the size of the eye and the shape of the back, which is frequently developed into a prominent hump behind the head, vary from one locality and even one individual to another. The common whitefish ranges from New England to Ungava (Great Whale River: Dymond, 1933, p. 3; Koksoak River: P. Orkin, personal communication) ; and from the Great Lakes northward to Baker Lake, the Thelon and Back Rivers (Hanbury, 1904, pp. 10, 40, 124, etc. ); it is present in Great Slave and Great Bear Lakes. A closely allied H not conspecific form is well known in the Mac­ kenzie and Yukon, and rivers and lakes of Alaska; it is known as the "humpback" or "crooked-back", and to the Indians at Arctic Red River as "teltsin" (C. nelsoni ).

12 The common whitefish falls into the same species-group as the universal palaearctic species C. lavaretus, known as "powan" in Scotland, "sik" in Sweden, and "sig" in the U.S.S.R. This is found in all the mainland watersheds of northern Europe and Asia, and is of considerable economic importance. The Mackenzie humpback most closely resembles the Siberian form C. lavaretus pidschian. There is a second species in the Mackenzie and Yukon, known there as the broad or true whitefish, and by various Indian names such as "khlugu-zhey" (Arctic Red River) and "tezareh" (Yukon Territory), and soientifically as Coregonus kennicotti. It differs from nelsoni in many prominent characteristics, most conspicuously in the larger maximum size (over 4 lb. or 2 kg.), the blunt snout and rounded head, the broad back, large adipose fin, pale-coloured eggs (orange-yellow in nelsoni ). The gill-rakers are short, less than half the height of the adipose fin, 21 to 24 in number. In nelsonii the gill-rakers are about equal to the height of the adipose fin, and number 25 to 28. The rim of the eye in kennicotti is very much farther from the supplemental bone of the maxilla than in nelsoni. There can be little doubt that the Mackenzie broad whitefish is the same species as the Siberian "chir" C. nasus, and it should be named C. nasus kennicotti. The differences which distinguish the chir from the sig are precisely those which have just been mentioned as distinguishing kennicotti and nelsoni: the chir has the characteristic blunt head, the wide space between orbit and maxilla, the short gill-rakers (19) 21 to 25 in number, and the more numerous scales of the broad whitefish, and similarly grows to a larger size than the sig. The broad whitefish spawns in the rivers in August; in the Mackenzie at Arctic Red River and elsewhere females outnumber. males by 12 or 15 to 1; the sex-ratio is Similarly unequal in the humpback whitefish. The remaining Siberian species, the "muksun" (C. muksun), has no counter­ part so far recognized in North America, having 44 to 65 gill-rakers, far in excess of the number found in any nearctic Coregonus. In addition to those mentioned already, there are some additional types peculiar to special districts, for example the "squanga" found in Squanga Lake, southern Yukon Territory, which is notable for its extraordinary development of rough warts, the so-called nuptial tubercles, from head to tail; for the fact that there are 26 rows of scales, counted diagonally down the side of the body, which is more than in any other described form; and for having 28 to 29 gill-rakers, the highest number of any nearctic Coregonus. Superficially, and in some fundamental characteristics, it comes closest to C. muksun aspius from Lake Onega in Russia. At Squanga Lake the common whitefish (nelsoni ) is also present, but the two are said to have different spawning beds. Spawning takes place in the late summer and fall, as late as November in some northern speoies. The lake whitefish spawn in shallow water, and the principal fisheries take place immediately before freeze-up and al�Q beneath the ice in winter. The river whitefish are often strongly migratory, and both of the Mackenzie-Yukon species enter the sea. The broad whitefish C. nasus kennicotti is not to be found above the first rapids, and flas not been taken above Camsell

13 o

em.

\ )L I

FIGUHE 2. CoregOntlS nelsoHi.

In C. nelsoni the snout is sharp, the brow concave, the infra-orbital bar very IH'TOW. In C. kCllllicotti the snout is blunt, the brow broad and convex, the infra-orbitals wider; the . margin of the pre-opercular is more sharply angled, and there are other differences in the shape and proportions of the opercular bones. Below are shown the anterior left gills, external view. In nelsolli the gill-rakers are long and 2.- o! ! CTn.

lcm.

FIGURE 3. COl'egonus kennicotti. sharply serrated, 25 or 26 in number in this specimen; in kennicotti they are much shOlter, scarcely serrate un the margins, and 21 in number. Both drawings are from specimens collected in the Mackenzie at Arctic Red River, N.vV.T., August 12, 1944. For comparison the two fish are reduced to the same apparent siz'�, though .the specimen of nelsoni is actually about two-thirds the length of the kennicotti. Bend in the main Mackenzie River. The upstream run takes place in July and August, the broad whitefish coming somewhat el\rlier than the humpback; and the �return may be in mid-winter, since there was formerly an important winter fisheryfor broad whitefish about Minto and Pelly Crossing in the Yukon Territory. Whitefish are usually caught in gill-nets set in shallow water; in rivers the nets must be short and set in eddies close to the banks; in Siberia the seine net is commonly used. Very occasionally whitefish will take the angler's fly. They are frequently cured by air-drying and smoking in wooded country. The fish is cut along both sides of the spine, and a single fillet prepared from the top side with­ out cutting into the belly; it remains attached to the spine at the tail; so that the prepared fish may be hung over a pole with the bones on one side and the flesh on the other; the head and entrails are removed. Salmon fillets are also dried and smoked by the Pacific coast Indians. The fish loses70 per cent of its weight by evaporation and is a valuable food, for dogs particularly, during winter travel. Fish is a staple food of sledge dogs throughout the north, and has been mucl1 used for feed in fox and mink ranches. A dog will consume up to 1,000 lb. of fish per annum.

ROUND WHITEFISH (Prosopium ) The round whitefish is.more slender in shape than the Coregonus speoies, with a small mouth, blunt snout, and fewer shorter gill-rakers (up to 16 only). It inhabits rivers and lakes. Prosopium cylindraceum is very widespread in the Arctic and Subarctic, both in Siberia (Yenisei excluding Baikal, also Lena, Kolyma, Anadyr and Kamchatka) and in North America (Alaska, Yukon, Mackenzie and eastwards to the Copper­ mine, Bathurst Inlet and Churchill River, Great Lakes, central and southern Ontario, Quebec and Ungava north to the K,oksoak (fide P. Orkin » . It may enter salt water at the mouths of rivers. The round whitefish from the Anadyr, Kam­ chatka and North America is P. cylindraceum quqdrilaterale, first described by Richardson from Fort Enterprise. It lis seldom sufficiently concentrated to be the object of a special fishery, although it is one of the most universal of northern freshwater fish. It spawns in the fall. Other species include the Rocky Mountain whitefish ( P. williamsoni ), not yet certainly known north of British Columbia; and a rare and interesting little fish known as P. coulteri, so far detected only in a few isolated places in 'Washing­ ton, British Columbia, southern Alaska (near Chignik and Lake Aleknagik) , and the Yukon Territory (Sockeye Lake on the upper Alsek, 60030'N., 137°38'W.) ; these are all in the Pacificslope drainage. The young stages of Prosopium species have "parr-marks", that is, a row of uniform large rounded patches of dark pigment like finger marks, on the sides of the body. In this they resemble the salmon-like fishes and grayling, but differ from other Coregonidae.

(v) TIrrMALLIDAE-GRAYLINGS The "bluefish" of and Alaska is the nearctic representative of the widespread species Thymallus arcticus, which in Asia extends from

16 Kamchatka and the Anadyr west to the Urals. In Canada the Arctic grayling extends southeastwards to the Churchill and its tributaries. It is present in the Thelon, Back, Coppermine and Anderson systems, and abundant in clear streams and shallow lakes draining into the Mackenzie and Yukon and their upper tribu­ taries; also throughout western and northern Alaska, south to the Nushagak River. It is not known to occur in rivers entering the Pacific proper, excepting the Alsek, which is also unique among Pacific coast watersheds in containing the round whitefish. Bluefish are distinguished by the magnificent , which is of great size, with blue spots on a dark ground and an edging of red. They may be caught on an artificial fly more readily than any other fish known to us, and attain a length of 16 inches (40 cm.) and a weight of about H4 lb. (0.8 kg.). Characteristically they are found in schools in clear water, and avoid the large turbid rivers; in the Mackenzie itself they may be taken at several pOints, such as Wrigley at the outlet of Great Slave Lake, and along the right bank at Simpson, where the water is clear; occasionally they penetrate milky glacier streams and are then pale in colour. Grayling apparently spawn in early spring, running out of the small creeks soon thereafter in May; Preble (1908, p. 512 ) describes an Indian grayling trap used at this season near Fort Simpson. They are good eating only if cooked as soon as caught.

(vi) OSMERIDAE-SMELTS Smelts are small fish usually less than 12 inches (30 cm.) in length, but rich in fat, sweet-tasting and extremely nutritious, notwithstanding their characteristic and somewhat disagreeable odour. Many of them are marine, spawning among the breakers on beaches or in shallow water, for example the well-known capelin Mallotus villosus of the coasts of the arctic North Atlantic, North Pacific and Bening Sea, and the "eulachon" or candle-fish of the Pacific coasts. Smelts have an adipose fin, and are silvery fish with a projecting lower jaw; teeth are developed and often conspicuous. One species (or perhaps superspecies ) spawning in fresh water is circumpolar, namely Osmerus eperlanus. It enters the sea freely, though it may be entirely landlocked in some waters. Typical eperlanus occurs throughout and the Baltic countries to the Murman coast. In Siberia, including Kamchatka, in Alaska and northwest Canada, east to the Mackenzie, a closely similar type is found, which may reasonably be placed in the same species, as O. e. dentex. It ascends the Mackenzie to Arctic Red River, and is common in the delta. (Berg does not record it from the Lena and Kolyma. ) Similarly the eastern forin is O. e. mordax, occurring in the Great Lakes-St. Lawrence region. It is scarcely subarctic at any point of its known range, which reaches Comeau Bay on the North Shore of the Gulf of St. Lawrence, Lake St. John and Rimouski. A second species, the pond smelt Hypomesus olidus, is a little-known an­ adromous fish, entering the rivers on both shores of Bering Sea, and the Mac­ kenzie; it has been taken by us in the Peel above McPherson, actually over the border of the Yukon Territory.

17 The chief value of smelts is as food for many kinds of predacious fish, seals and birds. ( vii) CATOSTOMIDAE-SUCKERS The common sucker Catostornus cornmersonni extends fmm the Great Lakes to Hudson Bay and the :\1ackenzie, as far down as Good Hope; but it is not known to occur in any waters in the . It is usually called the "grey sucker" in the northwest. The northern sucker C. catostomus is much more widely distributed; its northern range includes George River in Ungava (Legendre and Bousseau, 1949, p. 1.34), and the Albany, Nelson, Churchill and Thelon watersheds. In the last it is recorded in Artillery Lake (Critchell-Bullock, 1931, p ..33), head of Hanbury HiveI' (Hanbury, 1904, p. 41) and in the same river below Dickson's Canyon (Hornby, 1934, p. 109). It is also found in Great Bear Lake, the :Mackenzie to the delta. the whole Yukon watershed, the Alsek-Desedeash, and Alaska generally. Across Bering Strait it has been found in the Kolyma, Indigirka and Yana, and reported from the Anadyr. This species is the "red sucker" of the , though the same llame is used for other species in temperate parts of the . Suckers spawn in spring. They are not of much economic value, though by no means to be despised as food, in spite of the numerous small bones.

( viii ) CYPRINIDAE-:\hN KOWS Other than Coucsius, no cyprinids are known to occur in Ungava north of the St. Lawrence drainage, nor even in the latter east of Comean Bay (English Hiver). At least six species occur in arctic and subarctic America west of Hudson Bay. Pfrille (Phoxinus) ncogaca, Fine-scale minnow. Heported from Fort Severn, N. Ontario; Charlie Lake, Peace Hiver district, B.C.; and taken by us at Fort Good Hope, District of :Mackenzie. Rhinichthys cataractac, Long-nose dace. :\Iackenzie basin : at the mouth of the North Nahanni, at Good Hope, and in the Peel River at 66°.32'N., 137°0.5'\'\1. The last station is in the Yukon Territory. Otherwise known from the Atlantic to the Pacific, chiefly in the St. Lawrence drainage and thence south of the inter­ national boundary; but reaching at least the Severn and other rivers in northern Onthrio which enter James Bay (Radforth, 1944, p . 38. ). Coucsius plumbcus (including C. dissimilis and C. greeni ), Northern chub. :\Iackenzie basin : generally distributed down to the delta, but not found in Great Bear Lake. Yukon basin : below Five Finger rapids; Donjek HiveI'; Porcu­ pine River at Old Crow. This is the only cyprinid in the Yukon; it must occur in Alaska, though never reported. It is found in all provinces of Canada from New Brunswick to British Columbia; north to the Severn in Ontario (Radforth, 1944, p. 44), and the Indian House Lake region of the George River system, Ungava (Legendre and Rousseau, 1949, p. 134). Platygobio gracilis, Flat-head chub. A large "minnow", also frequenting the "\1ackenzie, but never reported below Good Hope. It extends southward to the

18 three prairie provinces. Originally described by Hichardson from Carlton House on the Saskatchewan. Notropis hudsonitts, Spot-tail shiner. St. Lawrence and Great Lakes north­ westward to the Saskatchewan-Nelson system, and Hayes Hiver above Yark Factory. �lackenzie watershed: Lesser Slave Lake, Lake Athabaska, and the main river down to Good Hope. N otropis atherinoides, Emerald shiner. Distributed as the previous species, but known north of 600N. only from \lills Lake and Simpson in the Mackenzie. In northern Europe and Siberia a larger number of cyprinids occur, including the roach Rutiltts rutilus, dace Leuciseus let/eiseus, ide L. idu8, minnow Phoxinlls phoxinus, tench Tinea tinea, gudgeon Cobia gobio, Crucian carp Camssills earassius. Several of these are fish large enough to be of economic value.

(ix) U MBRIDAE-ALASKA BLACKFISH, ETC. The Alaska blackfish, Dallia peetomZis, is one of the most curious and inter­ esting of northern fish. Its range is circumscribed, being chiefly in the low grounds and coastal regions of north\vest Alaska, from the southward, and also in St. Lawrence Island and a small area of the Chukotski Peninsula adjacent to the latter on the Asiatic side of Bering Strait. \Vhere found it exists in extraordinary numbers, and "is probably the most abundant of all the fishes which occur in the fresh and brackish waters of the northern part of Alaska" (Turner, 1886, p. 101). It has been reported in the Yukon from the delta, Andreafski, and at the junction of the Tanana; from the Kushkokwim and Nushagak Hivers; and around Norton Sound generally. The blackfish attains a length of only 8 inches (20 cm. ), and is exceedingly fat. It inhabits especially the countless small shallow ponds of the tundra, and even sphagnum "which at times seem to contain but sufficient water to more than moisten the skin of the fish" (ibid. ). Turner thought this mossy cover­ ing might help to protect them from the winter's cold. They spawn in June and July, and sometimes block the outlet streams of the ponds completely, so immense are their numbers. The Indians, in Turner's time, removed "tons and tons" of them daily. Undoubtedly blackfish are exceptionally adapted to withstand freezing. They are exceedingly tenacious to life; and, after being caught and left ,in wicker baskets to freeze, they may remain alive for weeks, and revive on thawing or when swallowed by dogs, although they have been frozen so hard that they must be chopped out with an axe or divided into pieccs with a club. Lucien .vI. Turner is still almost our sole authority on the natural history of Dallia, which certainly merits further study. The generic name was bestowed ill honour of Dr. W. H. DaIl, pioneer naturalist of the Alaska Telegraph Expedition, 1866, and collector of the original specimens of the blackfish.

(x) ESOCIDAE-PIKE The or "jackfish" Esox lucius is circumpolar. In the nearctic region its known northern limit extends into Ungava as far as 56°N. in the George Hiver (Legendre and Housseau, 1949, p. 1.34 ), though it has not been reported

19 from any rivers of the Labrador coast beyond English River at Baie Comeau; thence into the whole of Ontario and Manitoba, including the Albany, Severn, Nelson and Churchill Rivers. In the Northwest Territories it has not been found in any of the rivers of Keewatin, nor in the Back nor Coppermine watersheds, though it occurs in the Anderson (Preble, 1908, p. 513). It is present and often abundant throughout the lowland rivers and lakes of the Mackenzie system, including Great Bear Lake, where it attains a weight of at least 16 lb. (7�4 kg.) and is rather scarce and local. In the Yukon watershed it is generally distributed and usually common; it is present also in the Kowak River in Alaska (Evermann and Goldsborough, 1907, p. 273). Esox lucius does not occur in the Pacific coast drarinages of North America or Asia, sou:th of 600N. (that is, south of the Beripg Sea). In eastern Asia the pike is absent from Kamchatka except at the base of the peninsula (Vivinskoi River on the east coast and Penshina River entering Okhotsk sea); but it is generally distributed northward and westward, in the Anadyr, Kolyma and all the northern rivers of Siberia and Europe, excepting those on the Atlantic slope of Norway. It has never been reported from arctic or subarctic islands, but only from the mainland, for it is a stenohaline species unable to survive long in the. sea. The distribution maps published by Berg (1933 p. 606, fig. 558), and amended by Hinks (1942, p. 92 ), are evidently in­ correct regarding the nearctic northern limit.

(xi) PERCOPSIDAE-TROUT-PERCH This endemic American family contains only two species, one of which is Percop,sis omiscomaycus the trout-perch, a small and adaptable fish of no econ­ omic value, often frequenting sluggish muddy waters. Its flesh is semi-trans­ parent, and it is marked externally with large black spots; it combines in a highly anomalous manner the head and scale characteristics of a perch with the adipose fin of the salmonids. It grows to 6 inches (15 cm. ). It has a wide range, from the Mississippi valley and the Hudson River north to Lake St. John, James Bay, the Albany, Severn and Nelson systems; thence to the Mackenzie basin, in which we found it down to Good Hope (66°N.); and Miller (1947, p. 37) reported it in Great Bear Lake. It extends farther, having been found in 1945 at'Old Crow on the Porcupine River (67°40'N.), which flows to the Yukon; it should be looked for in Alaska.

(xii ) PERCIDAE-PERCH, PIKE-PERCH The American pike-perch ·Stizostedion (=Lttcioperca ) vitreum, more com­ monly known as the "dore", "wall-eyed pike," "walleye" or "pickerel," extends northward to Lakes Mistassini and Chibogamau and the waters draining into James Bay in western Quebec; to the whole of Ontario and Manitoba, as far as the Churchill (Vladykov, 1933, p. 25); and the Mackenzie watershed down to Great Bear Lake and the Mackenzie delta. Records have not been found for any of the rivers of Keewatin.

20 This is a valuable commercial species in YIanitoba, and very widely known as a sporting fish. Its normal size is 2 to 3 lb. (about 1 kg.), but it may exceed 12 lb. (5 kg. ). It is chiefly a lake fish, though generally distributed in the main Mackenzie River. The related palaearctic species do not extend north of the rivers entering the Baltic, or east of the Volga-Casp�an system. The American perch Perea flaveseens does not reach as far to the north as the dore, and has little claim to be included here. In Ontario and Manitoba it in­ habits the James and Hudson Bay rivers north to the Saskachewan-Nelson; it has been found in Lesser Slave Lake and Lake Athabaska in Alberta, but not, so far, in any part of the Northwest Territories. The Old World perch Perea fluviatilis, on the other hand, is generally dis­ tributed in temperate and arctic Europe and Asia, east to the Kolyma. The related "pope" or "ruff" Aeerina eernua is very Similarly distributed, but has no close relative in North America.

(xiii ) COTTIDAE-SCULPINS OR BULLHEADS These are small fishes, some< marine and others freshwater, of no economic importance but very characteristic of cold northern seas, rivers and lakes. All are bottom-feeders, and some of the littoral marine species may enter the mouths of rivers. An example is the long-horned sculpin Myoxoeephalus quadrieornis, found in geographically varying forms on all the Arctic coasts, that of Norway excepted, but lincluding the Baltic and Hudson Bay; and in the mouths of many of the rivers. The several freshwater forms of this fish retain throughout life some of their juvenile characteristics, lacking for example the four rugose spines on the head, to which the name quadrieornis refers; they are found in Wetter, Wener, Ladoga, Onega and other lakes in northern Europe, where they are regarded as relicts of the "Yoldia Sea." The so-called "Triglopsis thompsoni" of the Great Lakes seems to be an analogous derivative (cf. Vladykov, 1933, p. 32) . Of the ordinary freshwater bullheads it is convenient to distinguish those which are prickly-skinned all over from those which are mostly smooth or slimy. The former include Cottus sibirieus and C. rieei. C. sibirieus is found in the Ob, Yenisei, Lena and Kolyma (Berg, 1933, p. 699 ); it closely resembles C. rieei in the long curved preopercular spine, and the broad flat head which gives the name "spoonhead muddler" to the American species. Cottus rieei has the follow­ ing northern range : Great Lakes northward in Ontario to James Bay and Fort Severn (Radforth, 1944, p. 44); the Saskatchewan system in the Prairie Provinces (but not the Nelson?); Mackenzie watershed, in Lesser Slave Lake and Lake Athabaska (Dymond, unpublished ), in the main Mackenzie River to the delta, and in the Peel River. It has not been found in Great Bear Lake (nor yet in Great Slave, where it must be present ), and it is unrecorded from the Yukon system. On the Pacific coast it reaches some rivers in southeastern Alaska, for example in the Ketchikan area (Evermann and Goldsborough, 1907; p. 306) .

21 The smooth bullheads more or less resemble Cottus gobio, a palaearctic species practically universal in cold clear streams in northern Europe and Asia. C. cognatus, the most widespread freshwater bullhead in North America, is scarcely distinguishable from C. gobio; it was described by Richardson (1836, p. 40) from Great Bear Lake, and extends from there and the Hanbury River (Dymond, unpublished ) to Quebec and . Smooth bullheads are found in all parts of the Yukon system and Alaska, and they must for the present be included in C. cognatus; indeed all the diverse American material lumped together under this name could probably be equally well grouped into the superspecies C. gobio. Under this heading mention must be made of the unique endemic fauna of Lake Baikal, which comprises not only a ilUmber of peculiar amphipod crustacea and unique molluscs, one of which is a nudibranch, but also two endemic families of fishes related to the bullheads, namely the Cottocomephoridae and the Comephoridae. These contain about 16 species between them. There is abundant evidence that Lake Baikal is a very ancient body of fresh water, and the origin of the oldest part of the peculiar fauna dates from early tertiary times.

(xiv) GASTEROSTEIDAE-STICKLEBACKS

Two species of sticklebacks, Gasterosteus aculeatus and Pllngitius pungitius, are found in the Arctic. Both are euryhaline, occurring in salt, brackish or fresh water; but, being small fish and not migratory, individually they stay for the most part in one or other of these environments; and conspicuous differences are developed in G. aculeatus between marine and freshwater races. Though small, sticklebacks are sometimes so numerous, and so universal in their combined distribution, that they are probably the most important forage fish for the large predatory trout and char. The three-spined stickleback G. aculeatus has an interrupted Atlantic-Pacific distribution. It has been found in rivers entering the head of James Bay (Rad­ forth, 1944, p. 93 ), at Payne Lake, Ungava (Legendre and Rousseau, 1949, p. 134), at (Bean, 1882, p. 128) and Tayane (Dymond, unpublished) in Hudson Bay. From Baffin Island (Halkett, 192R, p. 117), Hudson Strait (Vladykov, 1933, p. 22 ) and south Greenland it extends commonly southward in the Atlantic coast drainage of Canada. It occurs in Iceland and Europe generally, east to the Urals. In the North Pacific region it is present from the Anadyr and Kamchatka to Japan; and on the American side from the Pribilofs and Dutch Harbor (Aleutian islands ) south to California. The nine-spined stickleback P. pungitius is more widely distributed. In North America it is present in the Great Lakes, Newfoundland, Labrador, Ungava, BaffinIsland (Manning, 1942, p. 129; Halkett, 1928, p. 117; at least 800 feet above sea level at : Soper, 1934, p. 131 ); the Churchill, other rivers of Manitoba:, and those entering James Bay; in the Mackenzie system, including the Slave River, North Nahanni and the delta, besides Great Bear Lake, whence it has been known since the time of Richardson (1836, p. 57 ). It has not been found in the Yukon Territory, but has been recorded from the lower Yukon River

22 at Andreafski and st. Michael; and from many other Alaskan rivers north to Point Barrow; also from the Pribilof and Aleutian Islands (Evermann and Goldsborough, 907, p. 273)). In A�ia it is recorded from every northern watershed, including those of Kamchatka and Bering Sea, though absent from Lake Baikal. It is generally distributed likewise throughout the waters of Northern Europe, and Greenland, but absent from Iceland.

(xv) GADIDAE-LING, BURBOT OR METRY Lota lata is the circumpolar freshwater ling, formerly known in the fur country of Canada as the "methy," a name, or by the French-Canadian name of "loche." In Britain it is the "burbot."

It 1� usually stated to "exist in every river _ and lake from Canada to the northern extremity of the continent" (Richardson, 1836, p. 248; similar references by other authors) . However, there seems to be only one published reference each for Keewatin and D ngava. Its northern limit, so far as known to us, is restated as follows. In Quebec it occurs east to Lake St. John and north to Chimo, Dngava Bay (Kendall, 1909) ; the latter record is based on L. M. Turner's collection, and Dr. M. J. Dunbar informs me that Turner's unpublished manuscript says of Lata maculosa : "The burbot is quite common in the lakes of the Ungava district; in fact more common than the number of specimens obtained would indicate." It is present throughout Ontario and Manitoba north to the Severn (Radforth, 1944, p. 32) and Churchill (Vladykov, 1933, p. 42) . Critchell-Bullock (1931, p. 34) caught one in the Hanbury-Thelon system during his journey to Hudson Bay, making the Irish comment that it is "almost absent." It is found in the Coppermine (Preble, 1908, p. 515, quoting Thomas Simpson's Narrative, 1843, p. 267), Great Bear Lake, and throughout the Mackenzie and Yukon systems. Other Alaskan rivers from which it has been recorded include the Kowak and Nushagak. It occurs south­ ward on the Pacific coast to the . In the palaearctic region it is found in south Britain, all the Baltic countries (but not Norway), and from the White Sea eastwards it is recorded by Berg (1933, pp. 784-5) from every northern watershed in Russia and Asia as far as the Kolyma and Anadyr. The Old World subspecies is L. lata lata; that in the Kolyma, Anadyr, in Alaska and the Mackenzie is L. lata leptura; and the common American form is L. lata maculosa (see Hubbs and Schultz, 1941, pp. 17-27; but Macken�ie and Yukon specimens do not appear to be intermediate between lcptura and maculosa, as anticipated by these authors). Freshwater ling occasionally attain a length of over 4 feet (120 cm.), and a weight of 75 lb. (34 kg.), but are always insipid in flavour and not esteemed as food: The liver is large and rich in oil. They are voracious in habits, and occur both in lakes and in large and small streams. They spawn in mid-winter under the "ice. Other species of Gadidae sometimes enter brackish and rivers, in-

23 eluding Microgadus tomcod (St. Lawrence and Labrador ), navaga ( = gracilis 2 (Mackenzie, Alaska, eastern Siberia ), and Gadus ogac (eastern Arctic and Greenland ). The tomcod is landlocked in Lake St. John, Quebec (Legendre and Lagueux, 1948, p. 157) .

(B) Amphibia In North America two species of frogs extend into the subarctic region of permafrost. These amphibians must hibernate in water of sufficient depth to escape being frozen in; and in summer the water must be warm enough for long enough to allow the development of the egg and tadpole stages. As far as can be ascertained, all the northernmost species metamorphose in their natal summer, never hibernating as tadpoles. Very interesting work by Moore (summarized 1949, pp. 315-338 ) has revealed important phYSiological differences between different American species of Rana in the l'elatiOl;} between water-temperature and the development-rate of the embryo. The species which extends farthest north, namely the northern wood frog R. sylvatica, has the highest rate of development at any given temperature; it has also the lowest "temperature coefficient", indicating that temperature-changes have the least effect in modifying the rate of development; and it can tolerate lmver minimum temperatures than other species. It may be observed that frogs extend farthest north in regions where the summer is warmest, and that their limit roughly coincides with the 50°F. (lO°e. ) July isotherm, rather than with a mean annual temperahlre of O°c., as has sometimes been stated; it appears to be the summer rather than the winter temperature which is limiting. Other known adaptations of northern races of frogs include larger egg-size, submerged egg-masses which cannot therefore be frozen in surface ice, and a marked tendency to shortening of the hind legs. A considerable number of Amphibia stop short in the region of James Bay in the east, and the 60th parallel in and Alaska, including Triturus viridcscens, T. gral1ulosus, Bufo amcl'icanus, B. hemiophrys, B. boreas, Rana pipiens and R. pretiosa. At least two species extend considerably further north than this: their northern limits are defined below.

NORTHERK \VOODF ROG (Rana syluaticacantab l'igensis ) Southern Alaska, Fort Yukon (Cope, 1889, p. 437 ); Selkirk and Lake Lebarge, Y.T. (personal observation ); down the Mackenzie valley to Fort Norman and 67°.30'N. at Old Fort Good Hope (Preble, 1908, p. 501 ); probably Anderson River (ibid. ); Great Bear Lake, collected by Richardson (Boulenger, 1882, p. 46 ); Churchill and York Factory on Hudson Bay (Preble, 1902 .. p. 133). In Ungava frogs of this species occur north to Fort Chimo at the mouth of the Koksoak River (Hildebrand, 1949, p. 168); the Eskimos there refrain from harming them, adhering to the widespread primitive belief that frogs have an influence on the weather. Hantzsch (1930 p. 195 ) also saw an unidentified frog on the Ungava coast. Packard reported "R. septentrionalis" from Okkak, Labrador.

24 There are no records of frogs on the east shore of Hudson Bay, however, north of East Main. The northern limit outlined here considerably exceeds the range shown in Moore (1949, p. 317).

NORTHERN SWAMP TREE-FROG (Pseudacris nigrita septentrionalis ). In the Mackenzie region well to the northward of Great Slave Lake, namely at Forts Simpson and Norman (Preble, 1908, p. 502 ); Wrigley, and perhaps the Franklin Mountains to the eastward, where "Tilliams (1933, p. 31 ) mentions hearing "pipers" in early July; Great Bear Lake, the typc locality of septcntri­ onalis (Boulenger, 1882, p. ,33.5 and plate 23 ), which is based on specimens collected by Richardson. Otherwise in the Prairie Provinces north to Norway House and York Factory, l\Janitoba (Preble, 1902, p. I.34). In the Old World the common frog Rana temporaria has the highest northern range, extending to the whole of Scandinavia, Finland and the U.S.S.R., to well above the Arctic circle. It ascends to 9,000 feet (.3,000 m.) above sea level in the Alps and Pyrenees, and to 4,000 feet (1,200 111. ) in the Dovrefeld in Norway, in both cases close to the snow-line. Several other species, including R. arvalis and Bufo vulgaris, ' just reach the Arctic circle. No Amphibia are native to Iceland, Greenland or Newfoundland, though introduced frogs have long been established in the last-named.

(C) Mammalia

WHITE WHALE OR BELUGA (Delphinaptcrus leu cas ) The white whale scarcely merits inclusion here. Like some other porpoises it often frequents estuaries, and in pursuit of fish it may penetrate a considerable distance into fresh water on temporary visits. The type locality of the species is the mouth of the Ob River. Nansen (1914, p. 148) saw it in the Yenisei at 69°43'N., more than 100 miles above the head of Yenisei Bay, It is very abundant in summer off the mouth of the l\fackenzie, where it was seen by the early explorers. Pors,ild (1945, p. 21) says that some ascend the larger branches of the delta, and they are reported to have been seen as far up as Point Separation at its head. Preble (1908, p. 128 ) mcntions a report of two seen in the Peel at Fort McPherson, 20 miles above thc delta and not much short of 150 miles from the open sea. Anderson (1937, p. 101 ) says it comes into the mouth of the Yukon, and Preble (1902, p. 40 ) that it enters the Churchill and Nelson. RINGED SEAL (Phoca hispida ) Like the beluga, the ringed and harbour seals are inclined to make excursions into fresh water, and often turn up far inland in large rivers. In both these species, however, there are populations partly or wholly isolated from the sea in inland waters. Ringed seals (Ph. hispida caspica ) occur in the Caspian and Aral Seas, which are, of course, salt; there is also a race (Ph. hispida sibirica ) in the freshwater Lake Baikal. Two other landlocked forms inhabit

25 Lake Saima in Finland (Ph. hispida saimensis ), and Lakes Ladoga and Onega in Hussia (Ph. hispida ladogensis ). In Arctic America there is a permanent population of ringed seals in NeU!illing Lake, Baffin Island, named Ph. hispida soperi by Anderson (1942, p. 27 ). This form is identical with that found on the adjacent coast of , and there may possibly be some sporadic movement between the sea and the lake, a distance of 52 miles, through the Koukdjuak Hiver. Manning saw a seal two miles up the river (Soper, 1944, pp. 238-9); he also found ringed seals in Lake Bennett, Melville Peninsula, in summer (l\ilanning, 1943, p. 104). HARBOUR SEAL (Phoca vitulina ) Sutton and Hamilton (1932, p. 37) report harbour seals visiting freshwater lakes in Southhampton Island; and Preble (1902, p. 71) saw one several miles above York Factory in the Nelson. They often ascend the St. Lawrence to Montreal and occasionally Ottawa; and Seton (1912, p. 56) mentions that he had been told of the presence of small seals "in the old days" in Lake Ashkeek, 50 miles northeast of Temiscamingue, which drains by the Kippewa to the Ottawa Hiver. It has been known since A. P. Low's traverse of 1896 that there were land­ locked seals in the Upper and Lower Seal Lakes in Ungava, 90 miles inland from Hichmond Gulf on the east coast of Hudson Bay. These had been assumed to be ringed seals, but Doutt (1942) discovered them in fact to be a dark-coloured race of harbour seal, named by him Ph. vitulina mellonae.

CONCL US ION In conclusion the writer wishes to acknowledge his debt to the great work of L. S. Berg on the freshwater fishes of the U.S.S.R, a debt regrettably lessened by the diffictIlties of the Hussian language, and failure to secure in Britain a copy of the existing translation. The book itself is superbly illustrated. In such a survey as this article provides there are sure to be errors, and there are known to be omissions. The northern ranges of the various species have been drawn up with as much care as possible, and every North American locality given is either original, or, much more frequently, taken from the available literature. Acknowledgement of the source is made in the text where the in­ formation provided is deemed not to be common knowledge. Certainly the preparation of the article has been remarkably instructive, and has shown how loosely and inaccurately the North American ranges of many freshwater verte­ brates have heretofore been defined.

REFERENCES ANDERSON, R. M. Mammals and birds of the Western Arctic District, Northwest Territories, Canada. In Canada's western northland, pp. 97-122. Ottawa, 1937. Two new seals from Arctic Canada with key to the Canadian forms of hair seals (Family Phocidae ). Ann. Rep. Provancher Soc. (Quebec ), 1942, 23-34, 1942. BEAN, B T. H. Notes on some fishes from Hudson's a.y . Proc. U. S. Nat. Mus. for 1881, pp. 127-129, 1882. 26 BERG, L. S. Les poissons des eaux douces de rU.RS.S. et des pays limitrophes. 3rd ed., 889 pp. Leningrad, Part I, 1932; Part 2, 1933. (In Russian. ) BOULENGER, G. A. Catalogue of the Batrachia Salientia. London: British Museum, 1882. COPE, E. D. The Batrachia of North America. Bull. U. S. Nat. Mus., No. 34, 1889. CRITCHELL-BuLLOCK, J. C. An expedition to sub-arctic Canada. Canadian Field-Naturalist, 45, 33-34 (Fish ), 1931. DOUTT, J. K. A review of the genus Phuca. Ann. Carnegie Mus. (Pittsburgh ), 29, 61-125, 1942. DYMOND, J. R Notes on the distribution of Salmo salal" and Salvelinus alpinus in north­ . Canadian Field-Naturalist, 46, 185, 19.'3 2. The Coregonine fishes of Hudson and James Bays. Contr. Canadian Biul. Fish., N. S. 8, :3-12, 1933. Pacific salmon in the Arctic Ocean. Proc. 6th Pacific Sci. Congress, p. 435, 1940. Atlantic salmon in Ungava Bay. Canadian Field-Naturalist 55, 19-20, 1941. The Coregonine fishes of northwestern Canada. Trans. Royal Canadian Inst. 24, 171-231, 1943. Note of specimen-localities in the Northwest Tenitories of fish, other than Coregonidae, in the collections of the Royal Ontario ;vluseum of Zoology (unpublished ). (Kindly furnished by Professor Dymond in 1944. ) A list of the freshwater fishes of Canada east of the . Royal Ontario Mus. Zool., Misc. Publ., No. I, 1-36, 1947. DYl\IOND, J. R, and V. D. VLADYKOV. The distribution and relationship of the salmonoid fishes of North America and North Asia. Proc. 5th Pacific Sci. Congress, pp. 3741-3750, 1934. EVERMANN, B. W., and E. L. GOLDSBOROUGH. The fishes of Alaska. Bull. U. S. Bur. Fish., 26, 219-360, 1907. GUNTHER, A. Account of the fishes collected by Capt. Feilden between 78° and 83°N. lat., during the Arctic Expedition 1875-6. Proc. Zool. Soc. London, 1877, 293-295, 1877a. Report on a collection of fishes made by .II'lr. C. Hart during the late Arctic Expedition. Proc. Zool. Soc. London, 1877, 47.5-477, 1877b. HALKETT, A. Check list of the fishes of the Dominion of Canada and Newfoundland. Ottawa, 1913. Notes on a collection of fish from Baffin Island. Bull. Nat. Mus. Canada, No. 53, 117-118, 1928. HANBURY, D. T. SpOlt and travel in the Northland of Canada. London, 1904. HANTZSCH, B. Contributions to the knowledge of extreme northeastern Labrador. Canadian Field-Naturalist, 45, 195 (Amphibia ), 1931. (Transl. by .M. B. A. Anderson. ) HARPER, F. Amphibians and reptiles of the Athabaska and Great Slave Lake regions. Canadian Field-Naturalist, 45, 68-70, 1931. HILDEBRAND, H. Notes on Rana sylratica in the Labrador peninsula. Copeia, 1949, 168-172, 1949. HINKS, D. The fisbes of Manitoba. : Manitoba Dept. of Mincs and Resources, 1943. HORNBY, J. vVildlife in the area, N. VV. T., Canada. Canadian Field-Nat1tralist, 48, 105-111, 1934. HUBBS, C. L., and L. P. SCHULTZ. Contributions to the ichthyology of Alaska, with descriptions of two new fishes. Occ. Papers Mus. Zool., Unit;. Michigan, No. 431, 1941. JOHANSEN, F. The crustacean life of some Arctic lagoons, lakes and ponds. Rep. Canadian Arctic Expedition, 1913-1 918, 7(N), 3-31, 1922. KENDALL, W. C. The fishes of Labrador. Proc. Portland Soc. Nat. Hist., 2 ( 8 ) , 207-244, 1909. LEGENDRE, V., and l J. ROUSSEAU. La distribution de quelques-uns de nos poissons dans Ie Quebec Arctique. Annales de l'Ass. Can.-Fran!;. pout' i'Avancement dcs Sciences, 15, 133-135, 1949. LEGENDRE, V., and R. LAGUEUX. The tomcod (Microgadus tomcod ) as a permanent fresh­ water resident of Lake St. John, P. Q. Canadian Field-Naturalist, 62, 157, 1948. "-1ANNING, T. H. Remarks on the phYSiography, Eskimo and mammals of Southampton Island. Canadian Geogr. J., 24, 17-33, 1942a. Notes on some fish of the eastern Canadian Arctic. Canadian Field-Naturalist, 56, 128-129, 1942b. Notes on the coastal district of the eastern barren grounds and Melville Peninsula from Igloolik to Cape Fullerton. Canadian Geogr. J., 26, 84-105, 1943.

27 �lrLLEH, R. B. l\orth 'Ncst Canadian fisheries surveys in 1944-194,5, Chap, 4 (Great Bear Lake ). Bull. Fish. Res, Bd, Canada, No, 72, 31-44, 1947. :VlooRE, J, A, Patterns of evolution in the genus Rana. In Genetics, palaeontology, and evolution, pp. 315-,3·38. Princeton, 1949. �IUNROE, E. G, Notes on the fish of the interior of the Labrador peninsula. Arctic, 2, 165-173, 1949. �ANSEN, F. Through Siberia, the land of the future. London, 1914. POHSILD, A. E. Mammals of the Mackenzie delta, Canadian .Field-Naturalist, 59, 4-22, HJ 4,5. PUEBLE, E. A. A biological investigation of the Hudson Bay region. N. Amer, Faullll, No. "" U, S. Dept, Agric" 1902, A biological investigation of the Athabaska-Mackenzie region. N. Amer. Fauna, No. 27. U, S. Dcpt. Agric" 1908, RADFOHTH, L Some considerations on the distribution of fishes in Ontario, ContI'. Royal Ontario Mus, Zool., No. 25, 1-116, 1944. RAWSON, D, S. North 'Nest Canadian fisheries surveys in 1944-194,5. Chap. 5, Great Slave Lake. Bull, Fish. Res. Bd, Canada, No, 72, 45-68, 1947, HrcHAHDsON, J, Fauna boreali-Americana, Part III (The fish), London, 1836, SEEBOHM, H. The birds of Siberia. London, 1901. SETO?>1, E. T. The Arctic prairies. London, 1912. SIMPSON, T. Narrative of the discoveries on the north coast of America. London. 184:3 SOPEH, J, D. A faunal investigation of sonthern Baffin Island, Bull. Nat, Mus, Canada, 1\'0. 53, 1928. Fishes. In Canada's eastern Arctic, pp, 131-1,32. Ottawa : Dept. ,ofthe Intcrior, 19.'34. The mammals of southern Baffin Island, N.vV,T., Canada. J. ., 25, 221-2,54, 194·1. SUTTON, G. M., and W. J. HAMILTON, Jr. The mammals of Southampton Island. Mem. Carnegie Mus, (Pittsburgh ), 12, 1-111, 19.'32. TummR, L. M. Contributions to the natural history of Alaska, Part IV (Fishes ), Arctic Series, Signal Sen)ice, Wash., No. 2, 87-11.'3, 1886. VLADYKOV, V. D, Fishes from the Hudson Bay region (except the Coregonidae ). ContI'. Canadian Bioi. Fish., 8, 13-61, 193.'3. \\'JLLIAMS, M. Y. Biological notes, covering parts of the Peace, Liard, l\'lackenzie and Great Bear River systems. Canadian Field-Naturalist, 45, 23-.'31, 193.'3. 'VY?>1NE-EDWARDS, V. C. North ''\Test Canadian fisheries survevs in 1944-1945, Chap. '2 (The Yukon Territory ); Chap. 3 (The 'dackenzie River ). B!:l!. Fish. Res. Bel. Canac/u, No. 72, 6-30, 1947.

28