National Museum of Natural Sciences Publications in Zoology, No. 4 Fauna of Sable Island and its Zoogeographic Affinities National Museums of Canada IF Or oP PCimmany ee SEP 8~ i979 \ RECEIVED y, rok OF SUNY te 1970 | | | a | Digitized by the Internet Archive in 2011 with funding from | California Academy of Sciences Library | | http://www.archive.org/details/publicationsinzo41 nati FAUNA OF SABLE ISLAND AND ITS ZOOGEOGRAPHIC AFFINITIES Pa RI F RER TE CANADA National Museum of Natural Sciences Musée national des sciences naturelles Publications in Zoology, No. 4 Publications en zoologie, n° 4 Issued under the authority of The National Museums of Canada FAUNA OF SABLE ISLAND AND ITS ZOOGEOGRAPHIC AFFINITIES — A COMPENDIUM By H. F. Howden J. E. H. Martin E. L. Bousfield D. E. McAllister Ottawa 1970 © QUEEN’S PRINTER FOR CANADA OTTAWA, 1970 Catalogue No. NM95-10/4 Contents The Coleoptera, by H. F. Howden, 1 The Odonata and Orthoptera, by J.E.H. Martin, 31 The Amphipod and Isopod Crustaceans, by E. L. Bousfield, 34 The fresh and brackish water fishes, by D. E. McAllister, 38 Appendices I Sable Island Noctuoidea, by E. W. Rockburne and D. F. Hardwick, 44 Il The freshwater and terrestrial Annelida, by G. E. Gates and J.E. Moore, 45 List of Tables Paper No. 1: I Climatic data — Sable Island, 6 II Daily climatic data for survey periods, 7 III Distributions of Sable Island Coleoptera, 20 Paper No. 5: I Data on Sable Island collections, 39 List of Plates Paper No. 1: Fig. ltos, 5 List of Maps Paper No. 1: 1 Sable Island, 4 2 Emergent areas of Sable and Grand banks during the Classical Wisconsin glaciation, 25 SUMMARY The faunistic groups discussed for Sable Island, Nova Scotia, include Insecta, Coleoptera (125 species), Odonata (5 species), Orthoptera (1 species), Lepidoptera (Noctuoidea, 23 species); Crustacea, Amphipoda (7 species), Isopoda (5 species); Annelida, Oligochaeta (2 species), Hirudinea (3 species); and Vertebrata, Pisces (3 species). Most of these have not been previously recorded from the Island, and a few are new to Canada. Collection data and climatic information for the survey periods in 1966 and 1967 are given. Zoogeographic comparisons are made for terrestrial, freshwater, and littoral marine organisms with particular reference to the species of adjacent land areas. The assumption that the Sable Island area and adjacent banks served as refugia during the last Wisconsin glaciation seems reasonable on the basis of the data presented. RÉSUMÉ Parmi les groupes fauniques dont il est question dans la partie traitant de l’Île de Sable, en Nouvelle-Écosse, figurent les Insectes: Coléoptères (125 espèces), Odona- tes (5 espèces), Orthoptères (1 espèce), Lépidoptères (Noctuidés, 23 espèces); les Crustacés: Amphipodes (7 espèces), Isopodes (5 espèces); les Annélides: Oligochètes (2 espèces), Hirudinés (3 espèces); et les Vertébrés: Poissons (3 espèces). La plupart n'avaient pas encore été observés dans l’île, certains sont même nouveaux au Canada. L'auteur présente des notes sur la capture des spécimens, et expose les conditions atmosphériques qui ont régné au cours des expéditions de 1966 et 1967. Il fait des comparaisons, du point de vue de la zoogéographie, entre les organismes terrestres, les organismes d’eau douce et ceux du littoral marin, en mentionnant tout particulièrement les espèces des zones terrestres adjacentes. En se fondant sur les données exposées, il semble raisonnable de supposer que la zone de l’Île de Sable et des côtes voisines a servi de refuge pendant la dernière période glaciaire du Wisconsin. vi THE COLEOPTERA BY H. F. HOWDEN* INTRODUCTION Sable Island is a sandy, crescent-shaped island that lies near the edge of the continental shelf at 43°57'N and 59°55'W, approximately 150 miles east of Halifax, Nova Scotia. The island has been the subject of interest and speculation, not only because of its historical interest, but also with regard to its biological and geological characteristics. Numerours papers have been written about the historical aspects of the island, and lists of the plants, verterbrates, and molluscs have been published (Erskine 1954; Clarke 1968). Despite this, no detailed study of the insect fauna was made until parties from the Entomology Research Institute, the National Museum of Natural History, and the Geological Survey of Canada visited the island in June 1966 and in July and September 1967. The present paper deals with the 126 species of beetles found on the island, their niches, abundance, time of occurence, and their zoogeographic relationships with the Nearctic and Palearctic faunas. ACKNOWLEDGEMENTS The Sable Island survey was made possible by the cooperation of numerous groups and individuals, whose assistance is gratefully acknowledged. In 1966 transportation was arranged through the Department of Defence, with Lt. Comdr. D. A. Muncaster, Commanding Officer of Helicopter Utility Squadron 21, Shearwater, N.S., in charge of the operation. In July and September of 1967 Mobil Oil Canada, Limited, furnished transportation and both Mr. A. R. Nielsen, President and General Manager, and Mr. B. E. George, in charge of operations at Halifax, were most helpful. While the survey party was on Sable Island, the members of the Department of Transport assisted in many ways, and we are particularly grateful to Mr. and Mrs. F. Androschuk and Mr. and Mrs. N. Bell. The following personnel made up the survey parties. June 6-13, 1966 — A. H. Clarke, National Museum of Natural History; J. H. Rick, National Parks Branch, Department of Indian Affairs and Northern Development; H. F. Howden and W.R.M. Mason, Entomology Research Institute. July 1-18, 1967 — W. J. Brown, H. F. Howden, J.E.H. Martin, R. H. Mulvey, and D. M. Wood, Entomology Research Institute; R. J. Mott and J. Terasmae, Geolegical Survey of Canada, Department of Energy, Mines and Resources. September 11-15, 1967 —E. C. Becker, J.E.H. Martin, and W.R.M. Mason, Entomology Research Institute. Sable Island climatic data for 1966 and 1967 were compiled with the assistance of Mr. G. Kendall, Meteorological Branch, Department of Transport. The following persons assisted with the determinations of the Coleoptera for the groups noted: Anthicidae — F. Werner; Aquatic Coleoptera — W. J. Brown; *Biology Department, Carleton University, Ottawa, and Entomology Research Institute, Canada Department of Agriculture, Ottawa. Carabidae — H. Goulet, C. Lindroth, R. de Ruette; Chrysomelidae and Curculioni- dae — W. J. Brown; Elateridae — E. C. Becker; Histeridae — R. Wenzel; Staphylini- dae — J. M. Campbell, L. Herman, M. Sanderson, A. Smetana; other Coleop- tera — W. J. Brown. Lists of other groups are credited to those who compiled them. ACTIVITIES OF MAN ON SABLE ISLAND In any consideration of the fauna of Sable Island, it is necessary to consider the influence of man with respect to the disturbance of habitats by purposeful and ‘chance’ introductions of plants and animals. Attempts to settle the island began in the 16th century; Lanctot (1933) has given a very interesting account of the La Roche settlement of 1598, which was not, as popularly supposed, merely the abandonment of prisoners. Even at that time there were wild cattle on the island that had probably been left (according to references cited by Lanctot) in 1539 by the Baron de Leri. The La Roche venture ended in 1603. Subsequently various attempts to stock or settle the island were made by a number of groups. In 1738 Le Mercier is said (Erskine 1954) “to have stocked the island with cattle, and it is presumed that he brought horses from New England at the same time.” In 1801, the government of Nova Scotia established lifesaving stations on the island, and at least a few people have been resident since that time. Today the Department of Transport maintains a meteorological station on the island with a staff varying ‘rom 11 to 15 persons. There are no other permanent residents. The introduction of various animals undoubtedly has had its effect on the native fauna. Cattle, pigs, Norway rats, cats, rabbits, dogs, and horses have all been present in numbers at various times during man’s residency (Cameron 1965). Today only the horses remain. Other strictly terrestrial mammals, such as mice, are absent. Plants, purposely introduced, have not fared much better. In 1901 an effort was made to stabilize the blowing sands by planting 81,345 trees and shrubs. A detailed account of this project was given by Saunders (1902), along with a list of the species introduced. Most of these were brought as seedlings from northern France (68,755 evergreens of 25 species, and 12,590 trees and shrubs of 79 species). They were packed in 18 boxes, without soil as far as I can ascertain, and were in transit for six weeks. A smaller shipment of about 2,000 plants, including 1,000 willow cuttings, originated from mainland Canada, the willows, at least, being brought from Brandon, Manitoba. Initially, the plantings appeared to be successful, but salt spray carried by fall and winter gales gradually eliminated the trees. The cause of the damage was not recognized as such at the time. In 1952 Erskine (1954) found one 15-inch shrub of buckthorn (Rhamnus frangula), which he suspected was the sole remnant of the 1901 planting. In 1954 Erskine listed 105 vascular plants that he had personally collected on the island and that he considered “native”, and another 27 (20 per cent) that he considered “introduced.” What portion of the introductions were purposeful remains unknown. Chance importation of plants and animals undoubtedly has been an important factor, although less important, perhaps, than in the adjacent maritime provinces 4 of Canada. Lindroth (1957) has developed a very plausable ‘ballast theory’ to explain the large number of European plants and animals (insects) that are now found in the maritime provinces. In the 17th and 18th centuries ballast consisting of dirt and rubble, which largely originated from France and England, was offloaded onto land because of laws against dumping the ballast into the major harbours. The fact that the flora and fauna of the ballast were often not subject to immersion in salt water accounts, at least in part (according to Lindroth), for the successful establishment of so many of the European species. This theory must be modified, however, when applied to Sable Island. The sandy shores and shoal waters surrounding the island prevent large ships from approaching the shore. At one time small ships could enter one lagoon, but since the early 1700’s all supplies have been brought to the island by lighter or surf boats. Under these conditions it is doubtful that ballast was ever offloaded onto the island. On the other hand, the numerous shipwrecks (see chart published by Grosvenor 1965) must be considered a likely source for chance imports. Here, as pointed out to me by J. Rick, one should consider the way ships were wrecked. When ships went aground, they were broken up slowly because of the sandy bottom. This means that any ballast would be thoroughly saturated with salt water, so that seeds or insects reaching the island via the shipwrecks would need to have some tolerance for immersion in salt water. Another source of chance imports was the hay brought at various times from mainland Canada as feed for livestock. This latter source could well be the most important one for the establishment of certain families of insects on the island. The problem of imports will be mentioned again under the discussion. PHYSICAL FEATURES AND CLIMATE As its name implies, Sable Island is composed entirely of sand. The island extends along an east-west axis (Map 1) for roughly 20 miles, has a maximum width of 1 mile and a maximum elevation of approximately 100 feet. The south beach is wide, with a very gradual slope; the north beach is much narrower and more abrupt. The highest dunes, which are generally along the northern side, attain maximum height near the east end. Between the dunes, where they flank the northern and southern beaches, there are often depressions, frequently filled with water. These lakes are most prevalent near the western end, diminishing in frequency and permanence toward the eastern end. Salinity of the lakes varies depending on exposure; several are essentially fresh water. The changing topography of the island has been of interest for many years. Any strong wind causes a veritable sandstorm, and the erosion by wave action has been particularly pronounced on the west end of the island. Predictions of the disappearance of the island and estimates of erosion of the west end appear in numerous references. Erskine (1954) listed various estimates: 1633 — esti- mated size of the island, 40 miles in circumference and much longer than wide (in this case certainly not more than 20 miles long); 1814 — four miles of west end washed away; 1820 — four miles of west end washed away; 1821-1851 — eleven miles of west end washed away; 1881-1883 — three miles of west end washed away; 1930-1939 — three miles of west end washed away. 0006S 4 00S SS06S 100009 050 I \ en | SS os SaNV1 YALVA oi HSau4 40 VauV —, OO oe er sniniminshalen-htintn lle = mm tn nn, " |: CORTE . TS Le ~~~ Ke were" ARE NOILVIS LHOIT eee LIN LSM n sm lle ,000pp — L ,000pp purysy ajqes — | den PLATE I Figs. 1,2. Fossil peat washed up on the south shore near the East Light 3. ‘Blow out’ along north shore near the West Light 4,5. Freshwater lakes between West Light and meteorological station 6. Collecting along drift line of freshwater lake 7. Vegetation and drift near freshwater lake 8. Ephalus latimanus, a flightless tenebrionid beetle TABLE I Climatic Data — Sable Island ‘Total Average Preci- or stat: velocity pitetion of wind Maxi- mum velocity Minimum Months Temp. 1900 F F Inches Days January 52:5 17.0 5.76 5 February 52.0 7.0 5 3 March 48.5 15.5 8 April 52.5 32.5 6 May 57.8 34.0 7 June 69.0 40.0 14 July 75.0 49.0 18 August 73.0 51.0 6 September 70.0 47.0 8 October 66.0 37.0 5 November 60.5 27.0 8 December 49.0 20.0 3 Averages 1901 January 47.0 5.0 3.24 2227 58 14 9 February 45.5 19.0 3.21 21.9 45 9 4 March 47.0 19.0 4.04 20.2 56 12 11 April 54.0 34,0 2.36 19.4 60 4 17 May 57.0 34.0 4.97 11353 34 1 10 June 63.0 44.0 2.38 14.8 36 1 15 July 77.0 53.0 2.90 12.9 36 0 19 August 78.0 60.0 3.36 11.3 34 1 13 September 76.5 48.0 1.65 17.4 42 5 7 October 68.0 41.0 4.52 18.4 48 4 9 November 57.5 30.0 2.10 18.2 62 7 0 December — = — = = = — Averages 60.95 | 1966 January 45 20 4.88 2273 64 1.8 4 February 47 3 2.02 19.6 52 8 4 March 47 19 4.11 16.1 38 6 12 April 45 28 0.87 13.9 39 5 13 May 58 34 6.35 14.3 43 5 19 June 60 42 4.96 127 32 1 21 July 70 48 6.23 11.1 28 0 26 August 71 49 3.49 11.9 29 0 14 September 70 43 2.69 13.6 36 1 11 October 65 33 4.61 15.5 40 5 12 November 60 28 1.81 15.2 34 2 10 December 55 26 6.64 16.9 40 10 6 Averages SWS 4.06 152 1967 January 50 13 6.71 16.7 52 10 5 February 51 6 5.83 20.9 58 14 5 March 49 12 3.78 18.8 44 13 7 April 52 25 3.81 17.9 42 5 10 May 53 35 gi 15.0 42 6 21 June 63 37 2.59 12.1 36 1 18 July 12 51 1.76 10.6 24 0 30 August 76 56 528) 10.6 24 0 18 September 76 48 2215 13.1 36 2 3 October 68 37 1.64 14.0 34 2 4 November 63 28 6.13 17.7 46 11 5 December 52 24 3.59 18.6 47 11 1 Averages 60.4 | 31.0 | 4.19 15.5 TABLE II Daily climatic data for survey periods Days with Date Maximum Temp. Minimum Temp. Total Precipitation some fog inches .04 0 .02 0 col OhnmOm ei bi © le ei be bi bei bit ni Oe Ho ui bi ni The above estimates total 25 miles of the west end washed away or, in different terms, a minus 5 miles of island! As the 1633 size estimate differs little from the present size of the island, possible explanations are that the estimates are incorrect, that the west end was rebuilt, or that the east end has been added to as the west end disappeared. Another estimate of the island’s size is mentioned by Saunders (1902). In 1901 it was said to be “about 21 miles long” with “early surveys giving the length [misreading of circumference? | as about 40 miles.” If only the 1901 estimate and the 1930-39 erosion estimates are considered, the figures still do not coincide with the size of the island today. The answer seems to be partly that the estimates have been exaggerated and partly that the island is eroding on the west end and being added to at the east. This shift in position is illustrated by Grosvenor (1965). Radiocarbon dating seems to bear this out, since the oldest surface strata found on the island occur near the west end and 7 are less than 700 years old. Fossil peat, washing up onto the beaches (figs. 1,2), indicates that the age of the neighbouring shoal areas is nearly 7,000 years B. P. (Medioli et al. 1967). The shifting position of the island is due, in part, to some of the climatic factors. Wind velocity averages about 18 m.p.h., and gales (32 m.ph. or more) are frequent; this, coupled with wave action, accounts for most of the shifting. A detailed account of these factors and their effects is given by James and Stanley (1967). Table I lists the monthly maximum-minimum of temperature, precipita- tion, and wind velocity for the years 1901, 1902, 1966, and 1967. Table II gives the daily data for the periods spent on the island by the survey parties during 1966 and 1967. It is evident that Sable Island has a true maritime climate, warmer in the winter and cooler in the summer than the neighbouring mainland. During the summer, fogs are frequent because of the proximity of the Labrador Current and the Gulf Stream. The generally high humidity and the cool summer temperature undoubtedly influence insect activity. Many of the beetles seem to occur as adults over an unusually long period, and I suspect that this is characteristic of many coastal species. LIST OF THE SABLE ISLAND COLEOPTERA All the various factors mentioned in the preceding sections have undoubtedly had their effects on the relatively meagre fauna of the island. In the following list of species some comments, such as “‘probably imported in hay,” will refer to the foregoing sections. Comments on distributions will be included under the discussion of zoogeography. Each species in the list is numbered, and the number is used in tables for reference to the species. The lists of families and genera follow the arrangement in Leng (1920); species are listed alphabetically under the genera. Both an estimate of abundance (i.e., rare, uncommon, moderately abundant, and com- mon) and the number of specimens actually taken (placed in parentheses after the estimate) are given, since in some cases only a moderate number of specimens of a readily recognizable species was taken; conversely a considerable effort was made to collect some uncommon species. Notes on the month(s) in which the species was taken and the type of area (niche) where it occurred are included. The general distribution of each species is given when possible; confused taxonomy precluded this in 14 species. A particular effort was made to check distribution records in areas adjacent to Sable Island; i.e., Newfoundland, coastal Nova Scotia, Prince Edward Island, and the New England states with particular reference to Cape Cod. Records were compiled for each species by personally checking the data on specimens housed in the Canadian National Collection, the Museum of Comparative Zoology, Harvard, and the collection of Boston University. Additional records were obtained from the literature (when the taxonomy was not in doubt). Other records were furnished by the specialists listed in the acknowledgments. If a species ranges over a wide area, only the general outline of the range is given. Family CICINDELIDAE 1. Cicindela hirticollis Say. Common (61). June through September. On or near beach, particularly south shore; occasionally found inland near lake 10. i. margins. Range: Newfoundland and Nova Scotia west to British Columbia and south in the East to Florida and Texas. . Cicindela tranquebarica Hbst. Common (71). June through September. Found in sparsely vegetated areas away from beach. Range: Newfoundland and Nova Scotia west to British Columbia and south in the East to South Carolina and Kansas. Family CARABIDAE . Carabus serratus Say. Rare (1). July. Under board inland near meteorolo- gical station. Range: Newfoundland and Nova Scotia west to British Columbia and south to Georgia in the East and to New Mexico and Oregon in the West. . Calosoma calidum Fab. Common (42). June through September. Occurs over inland portions of island. Active during day in well-vegetated areas. Range: Miquelon Islands, Nova Scotia, west to British Columbia, northward to Northwest Territories and south to Georgia and New Mexico. . Calosoma frigidum Kby. Rare (1). July. Taken at black light near West Light. Range: Nova Scotia and New Brunswick west to British Columbia and south to Georgia, Texas, Colorado, and Utah. . Dyschirius sphaericollis Say. Moderately abundant (21). June, July. On wet sand around lake margins, active mainly at night. Range: Newfoundland, Magdalen Islands, Quebec, west to Alberta; east of the Rocky Mountains, south to Florida. . Bembidion mimus Hayw. Uncommon (6). June through September. Under boards in moist localities inland. Range: Newfoundland, Nova Scotia, west to Manitoba and south to Illinois. . Bembidion s. sejunctum Csy. Common (80). June through September. Mainly under driftwood on beaches, particularly on north shore. Range: southern Labrador, Newfoundland, Magdalen Islands, Prince Edward Is- land, Nova Scotia, and Quebec; a disjunct population occurs from Alberta south to New Mexico. . Pterostichus mutus (Say). Common (61). June through September. Occurs in a variety of areas away from beaches, particularly near lakes. Range: Newfoundland, Quebec, west to British Columbia, and south through Massachusetts to Rhode Island. Amara aenea DeG. Uncommon (9). June, July. No habitat data. European import. Range: Newfoundland, Nova Scotia, Quebec, Ontario, and Mas- sachusetts; Europe, W. Asia. Amara familiaris Dft. Moderately abundant (22). June, July. In dune areas behind beaches. European import. Range: Newfoundland, Nova Scotia, New Brunswick, Quebec, Ontario, Massachusetts and, as a separate introduction, British Columbia; Europe; Asia, west to Mongolia. . Amara impuncticollis Say. Rare (1). July. No habitat data. Range: New- foundland, Nova Scotia, Quebec, Ontario, Manitoba, and Indiana. 13: 14. 18. 19 20. 21° 22° Amara quenseli Schonh. Common (36). July through September. Under boards and litter inland; most abundant in September. A Holarctic boreo- alpine species. Concerning Sable Island specimens, Lindroth (in litt.) states “these are unusually large and belong to the flat form (probably a modification) named ‘silvicola Zimm. in Europe.” Range: Labrador, Newfoundland, Nova Scotia west to British Columbia, north to Northwest Territories and Alaska; northern and central Europe; northern Asia east to Kamchatka. Amara rubrica (Hald.). Common (81). June through September. On surface of sand and under debris away from beaches; most abundant in September. Lindroth (in litt.) states “some are the largest specimens I have seen and therefore look strange.” Range: Quebec, Ontario, Massachusetts, and New Jersey to Texas and Colorado. . Amara torrida (Ill.) (= cylindrica LeC.). Common (161). June through September. Occurs widely over island except on beaches. A Holarctic species. Range: Labrador, Newfoundland, New Brunswick west to British Columbia and north to the Northwest Territories and Alaska; northern Eurasia. . Agonum cincticolle (Say). Common (74). June through September. Inland under debris. Range: Ontario, Massachusetts (Nahant), south to Florida, Texas, and west to Indiana. . Agonum gratiosum Mann. Rare (4). June, July. Under debris near fresh- water lakes. A northern species. Range: Labrador, Newfoundland, Prince Edward Island, Nova Scotia west to British Columbia, north to Northwest Territories and Alaska, south to Indiana and Oregon. Agonum placidum (Say). Common (49). June through September. Under debris near lake margins. Range: Labrador, Newfoundland, Magdalen Is- lands, Nova Scotia west to British Columbia, south through Massachusetts (Cape Cod), and central United States to Mexico. Agonum sordens (Kby.). Moderately abundant (30). June through September. Under driftwood on beaches. A northern species. Range: Labrador, New- foundland, Nova Scotia, west to British Columbia, north to Alaska, Yukon Territory, and Northwest Territories, south to Massachusetts and Oregon. Metabletus americanus (Dej.). Rare (3). June, July. Under boards. An apterous species. Range: Newfoundland, Nova Scotia, west to British Columbia, north to the Northwest Territories, south to Massachusetts, Indiana, Colorado, Arizona, and Washington. Cymindis cribricollis Dej. Rare (1). July. No habitat data. Range: Labrador, Newfoundland, Nova Scotia, west to British Columbia, north to the Northwest Territories, south to New York. Chlaenius pensylvanicus Say. Common (71). June through September. Found along lake margins and under boards in moist areas. Range: Newfoundland, Nova Scotia, west to British Columbia, south to Massachusetts (Cape Cod), New Jersey, Kentucky, Nebraska, Washington. 23. 24. 25: 26. 27. 28. 29° 30. Si. 32; 53. Harpalus affinis Schrk. Moderately abundant (19). June, July. Under boards, usually near lake margins. European import. Range: Newfoundland, Nova Scotia, west to Ontario, south to Florida and Kansas; as separate introduction in British Columbia and Washington; Europe; Asia east to Lena River, Siberia. Harpalus pleuriticus Kby. Rare (1). June. Under debris near lake margin. Range: Newfoundland, Nova Scotia, west to British Columbia, north to Alaska, Yukon Territory, Northwest Territories, south to Vermont. Harpalus rufipes (DeG.). Moderately abundant (16). June, July. Under boards inland. European import. Range: Newfoundland, Prince Edward Island, Nova Scotia, New Brunswick, Quebec; Europe; North Africa; Asia, east to Lenz River; Japan. Stenolophus fuliginosus Dej. Rare (5). July through September. Under debris inland. Range: Newfoundland, Nova Scotia, west to British Columbia, south to Rhode Island, Pennsylvania, and Indiana. Omophron labiatum (Fab.). Common (184). Numerous at night around fresh- water lake margins. Range: Sable Island form (colour pattern differs from New Jersey - Florida specimens) also occurs on Cape Cod, Massachusetts; southern form from New Jersey along coast to Florida and west to Texas. Omophron tessellatum Say. Common (69). July. Numerous at night around freshwater lake margins. Prince Edward Island, Nova Scotia, west to Alberta, south to Texas and Arizona. Family HALIPLIDAE Haliplus immaculicollis Harris. Common (130). June through September. In lakes near the west end. Range: Newfoundland, Nova Scotia, west to Alberta, south to Texas. Family DYTISCIDAE Bidessus affinis (Say). Common (66). June through September. In lakes near the West Light. Range: Quebec west to British Columbia, south to Massachusetts (Cape Cod), Texas, and California. Hygrotus impressopunctatus (Schaller). Common (40). June, July. In lakes near the West Light. A northern circumpolar species. Range: Newfoundland, Nova Scotia, west to British Columbia, north to Alaska, south to Massa- chusetts, New York, and Indiana; northern Europe; Siberia. Deronectes griseostriatus (DeG.). Moderately abundant (36). June through September. In lakes near the West Light. A northern circumpolar species. Range: Labrador west to Alaska (to tree line), Newfoundland, Nova Scotia, south to Massachusetts (Cape Cod), New Jersey, Iowa, Colorado, Arizona, and California; northern Europe; Siberia. Hydroporus notabilis LeC. (= arcticus Thomson, fide Brown). Rare (4). June through September. In lakes near the West Light. A northern circumpolar species. Range: Labrador, Newfoundland, Quebec, west to British Columbia, north to Alaska, south to Wisconsin and South Dakota; northern Eurasia. 11 34. 35? 36. Se 38. 39; 40. 41. 42. 43. 44. Hydroporus signatus Mann. Moderately abundant (31). June through September. In lakes near the West Light. Range: Labrador, Newfoundland, Nova Scotia west to Alaska, south to Massachusetts, New Jersey, and Illinois. Ilybius angustior (Gyll.). Uncommon (8). July through September. In lakes on west end of island, mostly in September. A northern circumpolar species. Range: Labrador, Newfoundland, west to Alaska and British Columbia; northern Eurasia. Ilybius biguttulus (Germ.). Common (79). July through September. In lakes near the West Light. Range: Nova Scotia, New Brunswick, Quebec, Ontario, Massachusetts, south to Virginia, west to Illinois. Colymbetes sculptilis Harr. Rare (5). July. In lakes near the West Light. Range: New Brunswick, Quebec, west to Saskatchewan, south to Massa- chusetts, New York, and Michigan. Family H YDROPHILIDAE Anacaena limbata (Fab.). Rare (4). June, July. In shallows of lakes near the West Light. European import. Range: Newfoundland, Quebec, Ontario, British Columbia, and Massachusetts (Cape Cod); Europe. Enochrus hamiltoni (Horn). Common (45). June, July. In lake shallows near the West Light. Range: eastern Canada, Massachusetts (Cape Cod area), New Jersey, west to Indiana. Cercyon haemorrhoidalis (Fab.). Uncommon (12). June, July. In horse dung. European import. Range: Newfoundland, Nova Scotia, Quebec, Ontario, British Columbia, south to Connecticut and Indiana; Europe. Cercyon near pygmaeus (111). Rare (1). July. Data uncertain, probably in horse dung. The specimen seems distinct from pygmaeus; the latter is European in origin, occuring in Newfoundland, south to Connecticut, and also in British Columbia. Cryptoleurum minutum (Fab.). Uncommon (9). June, July. In horse dung. European import. Range: Nova Scotia, Ontario, British Columbia, Massa- chusetts, Indiana; Europe. Family SILPHIDAE Silpha lapponica Hbst. Rare (4). September. On carrion. A Holarctic species, occurring in North America north to tree line, rare south of Canadian border. Range: Labrador west to Alaska, south to Pennsylvania, Iowa, Kansas, New Mexico, California; northern Eurasia. Family LEIODIDAE Leiodes assimilis LeC. Uncommon (9). July through September. Flying at dusk and found crawling in the early evening at base of sand dunes along the north shore. Range: New Brunswick, Quebec, Ontario, British Colum- bia, New Hampshire, and Colorado. 45. 46. 47. 48. 49. 50. 51° 52: 23: 54. 59: 56. SE 58. Family STAPHYLINIDAE Bledius basalis LeC. Rare (2). July. Collected under dead sea gull near top of north-shore beach. Range: Newfoundland, south along coast to Florida. Bledius opaculus LeC. Common (41). June through September. Mostly along top of ocean beach under dead sea gulls. Range: Along eastern seacoast south to New Jersey. Stenus dissentiens Csy. Uncommon (10). June, July. Under dead grass and debris near lake margins. Range: Michigan, Wyoming, Idaho, Utah, Oregon, Washington, and British Columbia. Stenus gratiosus Csy. Common (57). June, July. Under dead grass and debris near lake margins. Range: New Hampshire, Massachusetts, Michigan. Stenus juno (Fab.) Uncommon (14). June, July. Under debris in moist areas away from beaches. A European - North American species regarded by Lindroth (1957) as indigenous to North America. Range: Newfound- land, west to Washington, south to Massachusetts (Cape Cod) and Texas. Euaesthetus n. sp. (? ). Uncommon (11). June. Under a dead porpoise near the top of the beach on the north shore. Range: Unknown. Paederus littorarius Grav. Common (110). June through September. Under dead grass and debris near lake margins. Range: Nova Scotia, Ontario, Manitoba, British Columbia, Northwest Territories, New Jersey, and Ten- nessee. Hypnogyra sp. no. 1. Common (63). June through September. Under debris and horse dung. Range: Unknown. Hypnogyra sp. no. 2. Uncommon (12). June through September. Under debris. Range: Unknown. Philonthus alumnus Er. Moderately abundant (28). June, July. Under debris and carrion. A southern species. Range: New York, south to Florida, west to Arizona and California. Philonthus couleensis Hatch. Uncommon (15). June, July. Under debris. Range: Newfoundland, Nova Scotia, Washington, and British Columbia. Philonthus cruentatus (Gmel.). Uncommon (17). June, July. Under debris and horse dung. European import. Smetana (1965) maps the North American distribution and considers the species an introduction. Range: Newfoundland south to New York, west to Michigan; a separate population occurs in British Columbia south to Oregon; Eurasia. Philonthus debilis (Grav.). Rare (3). June, July. No habitat data. European import. Range: Newfoundland, south to Florida, west to Indiana and Nebraska; also a western population in British Columbia, Wyoming, Idaho, and Oregon; Eurasia. Philonthus fuscipennis (Mann.). Moderately abundant (22). June, July. Under dead grass and debris around lake margins, also under horse dung. European import. Range: Newfoundland, Nova Scotia, Ontario, south to 13 59; 60. OL: 62. 63. 64. 65. 66. 67. 68. 69. 70. TA Virginia and Illinois; western population in British Columbia, Washington, and Oregon; Eurasia. Philonthus lomatus Er. Common (40). June through September. Under debris, horse dung, and carrion. Range: Newfoundland, Nova Scotia, west to British Columbia, south to Florida and Indiana. Philonthus politus (L.) Rare (1). September. Presumably under horse dung. European import. Range: Newfoundland, west to British Columbia, south to North Carolina and Oregon; Europe. Philonthus sordidus (Grav.). Common (40). June through September. Under debris and horse dung. European import. Range: Newfoundland, west to British Columbia, south to Indiana and California; Eurasia. Philonthus varians (Payk.). Rare (1). July. No habitat data. A cosmopolitan species according to Smetana (1965). Range: Newfoundland, west to British Columbia, north to Northwest Territories, south to New Jersey, Colorado, and Arizona; Eurasia. Philonthus varius (Gyll.) Common (84). June through September. Under debris, horse dung. European import. Range: Newfoundland, Nova Scotia, west to British Columbia, north to Northwest Territories, south to Illinois, and Oregon; Eurasia. Philonthus vulgatus Csy. Uncommon (11). June through September. Under debris and carrion. Range: Nova Scotia west to Alaska, south to New York and California. Ocypus ater (Grav.). Rare (4). July through September. No habitat data. Probably European import. Range: Newfoundland, west to British Colum- bia, south to New Jersey, Louisiana, and California; Europe. Creophilus maxillosus villosus (Grav.). Common (42). June to September. Mostly under horse dung and old carrion. North American specimens differ slightly from European ones. Range: Newfoundland, Nova Scotia, west and south over much of North America to Central America. Heterothops fusculus LeC. Rare (2). July through September. No habitat data. Range: Newfoundland, Michigan, west to California. Quedius mesomelinus (Marsch.). Rare (4). July. Taken in old starling nest. A cosmopolitan species according to Smetana (1965). Range: Labrador, Newfoundland, west to British Columbia and Alaska, south to New York, Iowa, and California. Conosoma pubescens Payk. (of Horn) (? = testaceum Fab.). Rare (1). No habitat data. Possibly European import. Taxonomy of North American specimens confused at present. Range: Unknown. Tachyporus jocosus Say. Common (112). June through September. In dead grass and litter around lake margins. Range: Connecticut, west to Michigan and Indiana, south to Florida. . Mycetoporus americanus Er. Rare (9). June, July. No habitat data. Range: Pennsylvania, south to Florida, west to Illinois. 2. 13. 74. 15. 76. HE 78. 19. 80. 81 82. 83. Atheta sp. Common (187). June, July. Under horse dung and litter. Range: Unknown. Gnypeta sp. Rare (6). July. No habitat data. Range: Unknown. Falagria dissecta Er. (or near). Rare (9). June through September. Under debris near lake margins. Range: southeastern Canada, south to New York, west to Colorado and New Mexico. Oxypoda sp. no. 1. Rare (2). July. No habitat data. Range: Unknown. Oxypoda sp. no. 2. Rare (5). June through September. No habitat data. Range: Unknown. Family HISTERIDAE Hister curtatus LeC. (= lacustris Csy.). Rare (2). July. Under carrion. Range: Michigan, South Dakota, Indiana, Tennessee, South Carolina. Hypocaccus fraternus (Say). Common (180), June through September. Many found crawling on surface of sand, also common under horse dung and carrion. Range: Quebec west to Saskatchewan, south to Massachusetts (Cape Cod area), Virginia, Mississippi, Texas, and New Mexico. Baeckmanniolus palmatus (Say). (= dimidiatipennis LeC.). Common (50). June, July. Many found crawling on surface of sand, also under horse dung and carrion. According to Wenzel (in litt.) palmatus is primarily a coastal species and is closely related to two European species, B. maritimus (Steph.) and B. dimidiatus (IL). Range: Quebec (? ), Maine, Massachusetts (Cape Code area), south to New Jersey. Family OEDEMERIDAE Nacerda melanura (L.). Moderately abundant (27). July. Taken under old driftwood at top of ocean beach. Larvae found in same situation in June. A cosmopolitan species, often common in coastal areas. Range: Nova Scotia, New Brunswick, west to British Columbia, southward; Europe. Family ANTHICIDAE . Malporus formicarius (Laf.). Common (221). June through September. In litter and dead grass around lake margins. Range: Ontario, Massachusetts (Cape Cod area), Rhode Island, west to Iowa. Sapintus pusillus (Laf.) (= festinans Csy.). Uncommon (15). In dead vegetation around lake margins. Range: Quebec, Massachusetts (Cape Cod area), south to Florida, west to Colorado. Family ELATERIDAE Negastrius delumbis (Horn). Common (183). June, July. Found in sand dune areas at base of marram-grass (Ammophila breviligulata) and under Cakile edentula at top of beach. A flightless, coastal sand dune species. Range: Prince Edward Island (Cavendish), Nova Scotia (Lawrencetown), 15 84. 85. 86. 87. 88. 89. 90. 91. 92: Maine (Kennebunk beach), Massachusetts (Truro), and New York (River- mead, Long Island). Dalopius pallidus Brown. Common (71). June through September. Taken sweeping herbaceous vegetation. Range: Newfoundland, Nova Scotia, west to Saskatchewan, south to Massachusetts (Cape Cod area) and Illinois. Agriotes fucosus (LeC.). Uncommon (18). June, July. Under boards in areas of mixed vegetation near the West Light. A species usually associated with well drained forest margins. Range: Newfoundland, south to Maine and New Hampshire, west to Alberta, South Dakota, and Nebraska. Family HETEROCERIDAE Neoheterocerus fatuus (Kies). Common (58). June, July. In shallows along lake margins. Largely a coastal species. Range: New Brunswick, south to New Jersey. Family EUCINETIDAE Eucinetus terminalis LeC. Uncommon (14). July through September. Most- ly under boards in cranberry (Vaccinium macrocarpon) bogs. Range: Ontario, west to Saskatchewan, south to Vermont, Indiana, and Illinois. Family BYRRHIDAE . Cytilus alternatus (Say) (= ? sericeus Forst. of Europe). Moderately abundant (27). June, July. Near lake margins, some under boards. Range: Newfoundland, Nova Scotia, west to British Columbia, north to tree line, south to Massachusetts, Pennsylvania, and Colorado. Family NITIDULIDAE Omosita colon (L.). Uncommon (13). June, July. On and under old carrion. Considered a European import, now essentially cosmopolitan. Range: Newfoundland, west to British Columbia, south through Massachusetts to the southern United States; Europe. Family CRYPTOPHAGIDAE Cryptophagus fallax Bal.-Brn. (? ). Rare (2). July. Presumably in sparrow nests. Probably European import. Range: Maine (Mt. Desert Isle); British Isles, Northern Europe. Anchicera ochracea Zimm. Moderately abundant (34). July through Septem- ber. In straw and sparrow nests. Possibly brought in hay from mainland. Range: Ontario west to Manitoba, south to Maryland and Indiana. Family LATHRIDIIDAE Lathridius minutus (L.). Common (73). July through September. In straw and sparrow nests. Possibly brought in hay from mainland. European import, now a cosmopolitan species. Range: Newfoundland west to Alaska, south through most of the United States; Europe. 93. 94. 95, 96. 97: 98. oo. 100. 101. 102. Thes bergrothi (Reitt.). Rare (1). July. In straw. Possibly brought in hay from mainland. European import. Range: New Brunswick, Manitoba, Saskatche- wan, Washington; Europe. Microgramme ruficollis (Marsh.). Moderately abundant (34). July. In straw and chicken litter. Possibly brought in hay from mainland. European import. Range: Quebec, west to British Columbia, south to Massachusetts and Oregon; Europe. Melanophthalmia near cavicollis Mann. Uncommon (18). June through September. In dead vegetation around lake margins. Range: Unknown. Malanophthalmia near pusilla Mann. Common (47). June through September. In dead vegetation around lake margins. Range: Unknown. Family M YCETAEIDAE Mycetaea hirta (Marsh.). Rare (1). July. No habitat data. European import. Range: Quebec, Ontario, British Columbia, Massachusetts, south to Maryland, west to Michigan; Europe. Family COCCINELLIDAE Coccinella u. undecimpunctata L. Common (77). June through September. In mixed vegetation inland. European import. Range: Newfoundland, Prince Edward Island, Nova Scotia, west to Ontario, south to Massachusetts (Cape Cod area) and New York; Europe. Family TENEBRIONIDAE Blapstinus near metallicus (Fab.). Uncommon (17). June through September. Crawling on sand inland. Range: Unknown, but seemingly conspecific specimens examined from Cape Cod, Massachusetts, and coastal areas to the south. Ephalus latimanus (LeC.). Common (254). June through September. Com- monly found at night on the sand in the lower parts of the dune-grass formations. A flightless, coastal sand dune species. Range: Massachusetts (Cape Cod), south to New Jersey. Family PTINIDAE Ptinus fur (L.). Uncommon (15). July. Feeding on insect fragments in heated quonset hut at West Light. The only species found on the island that occurred only in heated buildings. European import, essentially cosmopolitan. Range: Nova Scotia, west to British Columbia and southward; Europe. Family SCARABAEIDAE Onthophagus nuchicornis (L.). Common (47). June, July. Only in horse dung. European import. Range: Newfoundland, Nova Scotia, west to Ontario, south through Massachusetts (Cape Cod area) to Maryland; occurring as separate population from British Columbia to Idaho; Europe. i, 103. 104. 105. 106. 107. 108. 109. 110. 18 Aegialia arenaria (Fab.). Common (289). June through September. Mainly in dune, marram-grass areas; often seen crawling on surface. A flightless species. Considered a European import, but I have some reservations. Darlington (1927) first reported the species from North America (Ispwich, Mass.) It has not been reported from other areas. The Cape Cod - Sable Island distribution is not a typical import pattern, hence my reservations. Range: Massachusetts; Europe. Aegialia spissipes LeC. (= ? rufa Fab. of Europe). Uncommon (10). July. Found crawling on sand at base of high dunes along beach between 6 and 7 p.m. Landin (1960) has questioned whether spissipes LeC. and rufa Fab. are distinct. So little material is presently available that the question seems difficult to resolve. If spissipes is the same as the European species, rufa, its rarity on both continents and its wide North American range make any explanation of its distribution difficult, particularly if one argues that it is a European import. At present, considering its range and the presence of other related species, I consider spissipes native to North America. Range: Miquelon Islands, south to Massachusetts (Cape Cod area), west to Michigan. Aphodius fimetarius (L.). Common (33). June, July. In horse dung. European import. Range: Newfoundland, Nova Scotia, west to British Columbia, south through much of the United States; Europe. Aphodius subterraneus (L.). Common (65). June, July. In horse dung. European import. Range: Nova Scotia, New Brunswick, Quebec, and New Jersey; Europe. Serica tristis LeC. Moderately abundant (38). June through September. Adults flew in daytime, feeding mainly on leaves of Rubis. Most common in area of meteorological station on west end of island. Range: Prince Edward Island, Nova Scotia west to Manitoba, south to Massachusetts (Cape Cod area), and New York. Phyllophaga drakii (Kby.). Common (68). June through September. Adults most active in early evening, feeding on several herbaceous plants including Rubis. Larvae found around roots of marram-grass (Ammophila brevili- gulata). Range: Nova Scotia, New Brunswick, west to Wisconsin, south through Massachusetts (Boston area) to Georgia. Family CHRYSOMELIDAE Phaedon near oviformis LeC. Common (182). June, July. Around lake margins, most common in early June. Range: Unknown, but seemingly conspecific specimens seen from Quebec (Fall collection), and from Mas- sachusetts (Woods Hole and Nantucket). Pyrrhalta sablensis Brown (1969). Moderately abundant (85). June, July. Adults and larva found feeding on cranberry (Vaccinium macrocarpon), seemingly restricted to one small area near a small freshwater lake just east of the West Light. The lake, which had a salinity of 25 parts per million, was less saline than any of the surrounding lakes. Range: Sable Island. 111: 112. 113. 114. 5: 116. 117. 118. 119. 120. 121: Altica potentillae Brown. Moderately abundant (32). June, July. Majority taken feeding on Potentilla anserina. Range: Nova Scotia, west to British Columbia, south to Massachusetts (Nantucket). Mantura floridana Crotch. Moderately abundant (35). June, July. Majority taken feeding on Rumex crispa. Range: Newfoundland, Nova Scotia, west to Manitoba, south through Massachusetts to Florida and Texas. Phyllotreta chalybeipennis (Crotch.). Common (120). June, July. Feeding on Cakile edentula growing near top of ocean beach. Range: Massachusetts (Nantucket and Marion), south along coast to Florida. Brachyrhinus ovatus (L.). Moderately abundant (29). June, July. Mostly under boards in grass-herbaceous areas. European import. Range: Newfound- land, Nova Scotia, west to British Columbia, south to Massachusetts (Cape Cod area), New Jersey, Iowa, and Colorado; Europe. Family CURCULIONIDAE Brachyrhinus sulcatus (Fab.). Common (47). June through September. Mostly under boards in mixed herbaceous areas. European import. Range: Newfoundland, Nova Scotia, west to British Columbia, south to Massa- chusetts (Cape Cod area) and North Carolina in east, California in west; Europe. Sitona scissifrons Say (a complex of forms). Common (42). June through September. Majority taken sweeping beach pea (Lathyrus japonicus) in dune areas. Range: Nova Scotia, New Brunswick, south to Massachusetts (Cape Cod area), and west to (at least) Manitoba and Iowa. Hypera nigrirostris (Fab.). Rare (1). July. No habitat data. European import. Range: Newfoundland, Nova Scotia, west to Ontario and Wisconsin, south to Maryland; also, as separate population, in British Columbia; Europe. Hypera punctata (Fab.). Rare (2). July. Sweeping herbaceous vegetation near lakes. European import. Range: Newfoundland, Nova Scotia, west to British Columbia, south to Maryland, Texas, and California; Europe. Hyperodes sp. Common (135). June, July. Most taken sweeping herbaceous vegetation, particularly in areas near lakes. Range: Unknown because of confused taxonomy; seemingly conspecific specimens seen from Prince Edward Island, Nova Scotia, and Massachusetts (Cape Cod area). Notaris bimaculatus (Fab.). Moderately abundant (32). June through September. Mostly found crawling on sand in beach-dune area. Differs slightly from majority of mainland specimens in appearing to be less robust. A northern species. Range: Nova Scotia, New Brunswick, west to Alberta, north to Northwest Territories and Alaska. Rhynchaenus pallicornis (Say). Common (105). June through September. Taken sweeping herbaceous vegetation near lakes. Range: Nova Scotia, New 19 1222 123: 124. 125: 126. Brunswick, west to British Columbia, south to Massachusetts (Cape Cod area) and Kansas. Ceutorhynchus hamiltoni Dietz. Rare (1). July. On Cakile edentula. Range: Newfoundland, Prince Edward Island, Nova Scotia, New Brunswick, Quebec, Massachusetts (Cape Cod area), and Rhode Island. Phytobius near sulcicollis Fahr. Rare (1). July. No habitat data. Range: Unknown (of sulcicollis—Ontario, Quebec, south to Maryland). Anthonomus signatus Say. Common (56). June through September. On herbaceous vegetation near lakes. Range: Prince Edward Island, Nova Scotia, west to British Columbia, south to Massachusetts (Cape Cod area) and North Carolina. Rhinoncus castor (Fab.). Uncommon (10). June, July. Mostly on Rumex. European import. Range: Newfoundland, Prince Edward Island, Nova Scotia, west to British Columbia, south through Massachusetts (Cape Cod area) to Florida; Europe. Sphenophorus cariosus (Oliv.). Moderately abundant (20). June, July. Crawling on ground in grass areas near lakes. Range: Massachusetts (Cape Cod area), south along coast to Florida and Louisiana. TABLE III Distributions of Sable Island Coleoptera Hudsonian- Eastern a Canadian Transcontinental 17, 19, 21, 24, 34, 88 6 Zone Circumpolar LSS 3132539.35645 Ul Primarily Coastal 83, 110, 122 3 Canadian Eastern 8, 37, 85, 104 4 Zone Transcontinental 44, 55, 120 3 Circumpolar = = Introduction 25, 38(? ), 63, 90, 103(? ), 106 6 Canadian- Coastal 45, 86, 100 3 Transition Eastern 6, 7, 12, 14, 36, 39, 48, 74, 77, 78, 79, 84, 18 Zone 87, 91, 107, 108, 112, 116 Transcontinental 1, 2, 3, 4,5, 9,.18; 20, 22, 26, 28; 29) 30; 2 47, 51, 59, 64, 67, 111, 121, 124 Circumpolar 49(? ), 62(? ), 66, 68, 80(?) 5 Introduction 10, 11, 23, 40, 42, 56, 57, 58, 60, 61, 25 65, 89, 92, 93, 94, 97, 98, 101, 102, 105, 114, 115, 117, 118, 125 Primarily Coastal 27, 46, 99, 113, 126 5 Austral Eastern 16, 54, 70, 71, 81, 82 6 Region Introduction ~ — Total 112 (126) ZOOGEOGRAPHIC RELATIONSHIPS The previous section includes the known distribution of each beetle species, with particular reference to eastern Canada and the northeastern United States. At best the distributions are incomplete, for in many cases the species either are poorly known or have been poorly collected. However, the general distributional picture given is fairly reliable for 112 of the 126 species. In attempting to chart these 112 distributions, some type of north-south grouping Was necessary, and I elected to use various combinations of Merriam’s classification of North America, i.e., Boreal Region with the Arctic, Hudsonian, and Canadian zones; Austral Region with the Transition, Upper and Lower Austral zones. Generally, various combinations of zones are used, as few species are limited to one zone or, in a few cases, to one region. Even then, species are listed as occurring primarily in one or more zones when occasionally recorded from other zones or regions. The east-west grouping within the various divisions in the chart seems self-explanatory, “‘coastal” referring to the East Coast only. In Table III, species are referred to by the number used in the list, and if there is any doubt that a species is an import, the number is followed by a question mark. The east-west divisions have been omitted from the table when not pertinent. In a few cases they have been included to emphasize the fact that none of the species fall within a particular category. If one views the distributions outlined above as they relate to Sable Islarid and if one considers the climate and location of the island, it becomes evident that there is an unusual mixture of faunas. The island represents essentially the southern limits of the ranges of the species listed under the Hudsonian-Canadian zones. Conversely most of the species listed as Austral have their northern limits on the island. A number of the latter, Omophron labiatum (27), Philonthus alumnus (54), Phyllotreta chalybeipennis (113), Sphenophorus cariosus (126), and in addition, Hister curtatus (77), Cryptophagus fallax (90), Ephalus latimanus (100), and Aegialia arenaria (103) have not been previously reported from Canada. Part of the explanation for this undoubtedly is the maritime climate: the relatively warm winter allowing the survival of some southern forms, the cool summers making possible the survival of northern forms. Interpretation of the other faunal elements necessitates comments on the general distributions of the various species. It is not surprising that the Hudsonian-Canadian species are either transcontinental or circumpolar, since these are the usual patterns for much of the northern biota. The small number of species considered as occurring primarily in the Canadian zone is an indication that few are restricted to this zone. This may be attributable to the effects of the Pleistocene glaciations. A large portion of the zone was glaciated, with the resulting extinction of relatively sedentary species. Survival had to be in peripheral areas in Alaska, in coastal refugia, or in areas to the south. The large number of species occurring today in the Canadian-Transition zone is indicative of reinvasion from the south. This seems to support the idea that if there was a distinct beetle fauna occurring primarily in the Canadian zone before the last glaciation, it was largely eliminated (Howden 1969). The coastal and eastern species considered as belonging to the Canadian zone survived either just south of the glacial limits or in an offshore refugium. The possibility of the latter will be considered subsequently. Species listed as 21 introductions (? ) might be circumpolar, and, conversely, species listed for the Canadian-Transition zone as circumpolar (? ) might be introductions. The latter, including those listed as introductions, have been included in the zone on the basis of both their distribution in North America and in the Palearctic region. In the Canadian-Transition zone, 86 per cent of the transcontinental species occurring on Sable Island are predators. Normally species associated with the coniferous forest constitute the conspicuous transcontinental elements, but since trees are lacking on Sable Island and few plants native to the island have a transcontinental range, it is not surprising that the ubiquitous North American species occurring on Sable Island are largely predaceous. In general, predatory species often have a wide food selectivity, and, of necessity, are more motile than their prey. The same statements are true, but to a lesser extent, for the Eastern Canadian - Transition zone species; of these, 61 per cent are predators. The species considered to be European imports on Sable Island comprise 24 per cent of the total beetle fauna (only 20 per cent of the vascular plants are European, according to Erskine (1954)). This is not surprising if one considers the large number of imported species occurring in adjacent areas of the Canadian maritimes (Lindroth 1957). Some of the species may have come, via shipwreck, direct from Europe. If many arrived in this manner, and if one considers the number of shipwrecks (well over 200), it seems odd that all the European species that occur on Sable Island are found also in adjacent coastal localities of North America. Other aspects such as the ballast theory, have been discussed under the heading “Activities of Man on Sable Island”. There does seem to be a strong possibility that many of the European species that first became established on the Canadian mainland were then brought to Sable Island with livestock or in hay, straw or produce. In the chart there are no imports listed as occurring on Sable Island for the Austral region. A number of imports exist there, but none have become established on Sable Island. Several species, particularly Cryptophagus fallax (90) and Aegialia arenaria (103), that are considered introductions, have rather unusual restricted ranges in North America, not unlike the restricted coastal ranges of a few native species. I wonder, as have Darlington (1927) and others, about the possibility of some species having relict amphi-Atlantic distributions. If, at times of lowered sea levels, exposed portions of the continental shelf could serve as refugia, there could also be the possibility that a few species have natural amphi-Atlantic distributions. At present it seems best not to speculate on the question of amphi-Atlantic distributions. How- ever, the question of offshore refugia during the Pleistocene in the vicinity of Sable Island is worthy of consideration. THE SABLE ISLAND AREA — A PLEISTOCENE REFUGIUM? Certain elements of the flora and fauna surrounding the Gulf of St. Lawrence have long puzzled biologists. Many beetles that occur in the Maritime region attain their maximum size on Newfoundland. Lindroth (1963) discussed this phenomenon as seen in some ground beetles (Carabidae) and concluded that “the presence of populations deviating in size from what is normal within the species should not be used as an argument in favor of longtime isolation and ‘subspecies’ 22 should not be fourded on differences in size alone.” Despite this, Lindroth (1963: 107) does consider that some carabids may have survived the last glaciation on or near Newfoundland. Munroe (1956) cited a number of papers pertaining to the controversy about a refugium in the area and concluded that, while theories on migrating biotas may be largely correct, they do not, “however, account for the general occurrence of endemics and disjuncts in the Gulf region, at low levels as well as high both as localized forms and as radiants in Labrador and elsewhere in the Northeast. An eastern refugium separate from the general southern glacial-margin strip is required.” The validity of the above is contingent upon the geological history of the Sable Island and Grand Banks areas. Were they emergent during intervals of lower sea levels and were they unglaciated? Could they have served as refugia during the Pleistocene or, more particularly, during the last (Classical Wisconsin) glaciation? Theories concerning the age of the Sable Island formation and its origin are as confusing and controversial as some of the historical aspects. There can be no doubt that the present emergent area, i.e., Sable Island, has shifted. This makes carbon dating of surface features nearly useless. Cameron (1965) and James and Stanley (1967) discussed the shifting nature of the island. Cameron (1965) also suggested that it was formed from outwashings of glacial till, speculating that currents caused a 1,400-foot dome of sand to be formed over ‘‘bedrock” and that the age of the emergent area is no more than 6,000 years. Erskine (1954) discussed a similar hypothesis, but in less detail. In a recent analysis of the geology of the area Stanley and Cok (1967) presented evidence and ideas concerning the extent of the Wisconsin glaciation and concerning unglaciated areas that were probably exposed during periods of lowered sea level. Their findings coincide to a considerable degree with unpublished data accumulated by Drs. V. K. Prest and J. Terasmae of the Geological Survey of Canada. In general the Sable Island Banks represent an old formation, being, according to Stanley and Cok, “essentially erosional in origin and consisting of sedimentary deposits of late Mesozoic to Tertiary age.’ Near and on Sable Island there is a thin overlay of Pleistocene age, attaining on Sable Island an approximate depth of no more than 200 feet (Terasmae, pers. comm.). As shown on Map 2 and also by Stanley and Cok, a considerable portion of the Sable Island bank was exposed and unglaciated during the last maximum. The question remains, could and did this area serve as a refugium for plant and animal life? From work done by Lindroth (1965) and others, there seems little doubt that a considerable flora and fauna can survive adjacent to a glaciated area. Undoubt- edly much depends on climatic factors, and I am not qualified to speculate on the possible Pleistocene climate of the Sable Banks. The position of the Gulf Stream during that time would be very important. With the emergent areas of the Grand Banks possibly deflecting the Labrador Current, the Gulf Stream may have been even closer to the Sable Island area than its present distance of 50 miles. If true, a narrow coastal strip may have had a relatively mild climate. Good biological evidence for a refugium is scanty. Fossil peat from 8,000 to 10,000 years old (C14 dated) occurs in the area, having washed up from presently submerged areas. This suggests that a larger vegetated area certainly 23 existed during those times. Further to the south, evidence derived from ele- phant teeth (Whitmore ef al. 1967) and from fossil peat (Emery et al. 1967) indicates that there was undoubtedly a considerable biota present on exposed areas of the continental shelf during the last glacial maximum. Sable Island, if the geological evidence is correct, is the last exposed remnant of a much larger area. However, because of its shifting location, the length of time that the biota has been in residence remains problematical. Some extinction of the fauna has undoubtedly occurred with the reduction of the land area, and further disruption has resulted from the activities of man by his importation of plants and animals. How long the native elements have been established is a matter for speculation, since no fossils are available. The only vertebrate animal possibly indicative of an extended period of residence is the Ipswich Sparrow, and this subject is controversial. Since more species of insects can live in a small area, they seem more useful than vertebrates as indicators of a possible refugium, at least for Sable Island. Of the beetles, only one species, Pyrrhalta sablensis (110), is seemingly endemic to the island. The species is closely related to a few mainland forms but is undoubtedly distinct on the species level. Since the species is winged and since there is seemingly no way to judge the time needed for divergence, its presence is not necessarily indicative of a long residence. The flightless click beetle Negastrius delumbis (83) occurs in coastal sand dune areas from Prince Edward Island, south along the coast of Nova Scotia, then to Cape Cod, Massachusetts, and coastal Rhode Island. Most, if not all, of the mainland areas in which the click beetle now occurs were glaciated. Its spotty and relatively limited distribution and its inability to fly make the present range difficult to explain, unless one considers survival during the Pleistocene in areas presently submerged. The same can be said for the tenebrionid Ephalus latimanus (100). This large flightless species (fig. 8) iscommon on Sable Island and occurs in dune areas to the south from Cape Cod to New Jersey. Another species with the same northern coastal type of distribution is the winged weevil Ceutorhynchus hamiltoni (122), which occurs on Cakile along ocean beaches from Newfoundland south to Rhode Island. Two other groups also warrant consideration. One contains northern forms, occurring largely in Canada, with Sable Island now representing, essentially, the southern end of their distributions. The weevil Notaris bimaculatus (120) belongs to Holarctic, northern genus, with Sable Island representing primarily one of the more southern portions of its range. Specimens from Sable Island and the adjacent maritime regions are more slender than more inland forms, and there is the possibil- ity of some confusion in their taxonomy. Other species that belong in this ‘northern’ group are Bembidion sejunctum (8), Colymbetes sculptilis (37), Agriotes fucosus (85), Aegialia spissipes (104), Serica tristis (107), and some Staphylinidae. Both Agriotes fucosus and Serica tristis are species usually found along woodland margins: their presence on Sable Island could be indicative of an adaption to coastal conditions as the emergent areas gradually reduced in size. Phyllophaga drakii (108) is another species with a more southern range, usually associated with open woodland areas. 24 os 009 089 <0? À Hid3q 008 —---~ d!8 000'2 * 00021 i 43402 321 WnWIxyH 77777777) NY320 DILNVILY SP À ! | INOL2I#30384 oS? L 1 0 NAOL3LLOTAYHD 4 c seg Fa Ky ¢ + (Era € 1Y2IMSNNUS MAN u Seas _ > NO1aNDIW \ eee \ esc! sanvasi in ‘ Nava WW ES 4 GNYIONNOAAIN 1238300 00S 5 F0 1 Ay a 00S T : 055 009 {069 SL UOTJEIOEIS UISUO9SIM [BOISSETD oY} SULINp syURg pUeID pue aJqeS JO svore JUS — 7 dep 25 The last group contains southern (Austral) coastal species, with the northern end of their range on Sable Island. These are Omophron labiatum (27), Bledius opaculus (46), Blapstinus near metallicus (99), Phyllotreta chalybeipennis (113), and Sphenophorus cariosus (126). Although all of these may be capable of flight, none, except perhaps the Phyllotreta, can be considered strong fliers. None of the species have been found on the adjacent mainland of Nova Scotia, the Cape Cod area being, in most cases, the northern point of their mainland distribution. In Omophron labiatum, specimens from Sable Island and Cape Cod consistently differ in colour pattern from more southern specimens; no other differences have been noted. Since it is well known that the ranges of both northern and southern forms often extend further south or north along the coast, these elements are to be expected and, in themselves, throw little light on the age of the island. However, when considered in conjunction with the presence of flightless species and the present isolation of the island (more than 90 miles from the nearest point of land), they can best be explained by the theory of dispersal over large, emergent, coastal areas during times of lowered sea level. The importance of dispersal by wind or currents (Darlington 1938) is nearly impossible to assess. Major storms in the Sable Island area occur most frequently during the fall and winter, when there is little insect activity. Because of this, I consider wind transport of the flightless or heavy-bodied insects as less likely in the Sable Island area, particularly over the water gaps as they now exist, than it might be in areas further to the south. The Sable Island Banks and adjacent areas during the Pleistocene probably resembled physiographically the present, somewhat dissected, coastal areas stretching from New Jersey to South Carolina. Faunal dispersal along this coast appears to be relatively easy, with a few areas, such as Chesapeake Bay, constituting barriers for some species. The derivation of the native Sable Island beetle fauna can, conceptually, be best explained by their having dispersed during a time when there were large areas exposed with only relatively minor water barriers interrupting north-south migrations. In considering the fauna of Sable Island, its possible origins, and its methods of dispersal, comparisons have been largely made with adjacent areas of the mainland, in relation to both the native and the imported elements. A comparison of the Sable Island fauna with that of another essentially similar island is the next desirable step. Unfortunately, there is no really suitable island for such a comparison. The most similar, at least in climatic respects, is Iceland, where most, if not all, of the fauna seems to be postglacial in origin. According to Larsson and Gigja (1959), not more than 160 species of beetles breed in Iceland. Of these, probably 25 or more cannot survive outside heated buildings. Of the 126 species on Sable Island, all but Ptinus fur breed outdoors; thus the two islands have essentially the same number of free-lving species. Larsson and Gigja believe that 81, or 50 per cent, of the 160 species living on Iceland are imports. Of the 125 free-living species, 56 or 42 per cent, are imports. However, they consider as indigenous to Iceland 12 species that are imports in North America; this suggests that their percentage of imports is much too low. Of the free-living species, only 8 species are common to both Sable Island and Iceland, and these are surely imports to both. 26 Essentially, Iceland has nearly twice the number (or percentage) of imported free-living species as are found on Sable Island. Some of this difference between the two islands is certainly due to their proximity to different faunal regions; the rest might be the result of different glacial histories, ecological factors, differences in types of imports, or other unrecognized factors. These, unfortunately, seem to exclude speculation on the relative ages of the two faunas. However, if the variables can be considered to be roughly equal, then it might be argued that Sable Island has the older fauna. Iceland has an area of approximately 30,000 square miles, over 100 times the size of Sable Island. Climatically the two are rather similar. The weather data for Vestmannaeyjar, Iceland, are as follows: mean temperatures: January, 34.5°F; April, 39.6°F; July, 52.6°F; October, 42.1°F; record extremes: 71°F maximum, -6°F minimum. Rainfall averages 52.9 inches per year. These data are very similar to those for Sable Island as given in Table I. If one utilizes the theories of MacArthur and Wilson (1967), Iceland is about 5 times as far from its faunal source area (for beetles) as is Sable Island, but it has 100 times the area of Sable Island. Therefore, if time is not a factor, it might be expected to support a larger fauna. Since both islands support 125 free-living species, the time available for faunal invasion may be a factor that explains at least part of this discrepancy. Iceland was largely glaciated, and Sable Island seemingly was not, so the above comparison at least does not refute the idea of faunal survival on Sable Island during the Pleistocene. CONCLUSIONS The presence of endemic species and the seemingly unusual distributions of some animals and plants occurring in the Canadian maritimes and around the Gulf of St. Lawrence have caused numerous workers interested in biogeography to theorize on the possibility of an offshore refugium during the last glacial maximum. Recent geological work lends credence to this biological theorizing. From the geological standpoint there is considerable evidence that large areas in the vicinity of Sable Island were exposed and unglaciated during the Classical Wisconsin Maximum. Unfortunately the geological evidence: does not at present give any indication of the composition of the Pleistocene biota of the area or even of whether there was any. The biological evidence is still inconclusive, but the information available seems to support, or at least does not refute, the theory of offshore refugia. Of the 126 species of Coleoptera on Sable Island, about 12 per cent have distributions most easily explained by invoking the concept of offshore refugia. Only one species, Pyrrhalta sablensis, is endemic. Explaining the presence of others, such as the flightless Negastrius delumbis and Ephalus latimanus, by chance dispersal over the present water barriers is difficult; their dispersal over relatively narrow water barriers seemingly is a more logical hypothesis. The assumption that the Sable Island area and adjacent banks served as refugia during the last Wisconsin Maximum seems reasonable on the basis of present data. During glacial expansion, offshore areas became exposed and available for invasion from the mainland. Water barriers were greatly reduced, and north-south movement along the newly exposed shore-lines perhaps accounted for the present 27 odd mixture of northern and southern species now present on Sable Island. During glacial recession, reinvasion of the newly exposed areas of mainland initially must have been relatively easy, becoming less so with rising sea levels. Reduction in the size of Sable Island and inundation of other areas have undoubtedly caused the extinction of much of the biota. However, a small fraction of the fauna does seem to support the concept of the area serving as a Pleistocene refugium. REFERENCES BROWN, W. J. (1969). A new species of Pyrrhalta from Sable Island (Coleoptera; Chrysomelidae). Can. Ent. 101: 109. CAMERON, H. L. (1965). The Shifting Sands of Sable Island. Geographical Review 55: 463-476, DARLINGTON, P. J., Jr. (1927). Aegialia arenaria Muls. in New England, with local records for other species. Psyche 34: 98-99. CLARKE, À. H., Jr. (1968). Recent and Post-Pleistocene Mollusca from Sable Island, Nova Scotia. Annual Reports for 1967 of the Amer. Malacological Union pp. 11-12. EMERY, K.O.,R. L. WIGLEY, A. S. BARTLETT, M. RUBIN, and E. S. BARGHOORN (1967). Freshwater peat on the Continental Shelf. Science 158: 1301-1307. ERSKINE, J.S. (1954). The Ecology of Sable Island, 1952. Proc. N.S. Institute of Science 23: 120-145. GROSVENOR, M.B. (1965). Safe landing on Sable, isle of 500 shipwrecks. National Geographic 128: 398-431. HOWDEN, H. F. (1969). Effects of the Pleistecene on North American insects. Ann. Rev. Ent. 14: 39-56. JAMES, N. P. and D. J. STANLEY (1967). Sediment transport on Sable Island, Nova Scotia. Smithsonian Misc. Coll. 152 (7): 1-13. LANCTOT, G. Fe (1933). L’establissement du Marquis de la Roche a l'Ile de Sable. Canadian Historical Association Annual Report, 1933, pp. 33-42. LANDIN, BENGTH-OLOF. (1960). Lamellicorn beetles from Newfoundland and Nova Scotia. Opus. Ent. 25: 129-137. LARSSON, S. G. and G. GIGJA (1959). Goleoptera 1. Synopsis. Jn The Zoology of Iceland. Vol. 3, part 46a. Ejnar Munksgaard, Copenhagen and Reykjavik. 218 pp. LENG, C. W. (1920). Catalogue of the Coleoptera of America, North of Mexico. John D. Sherman, Jr., Mount Vernon, N.Y. 470 pp. LINDROTH, C. H. (1957). The faunal connections between Europe and North America. John Wiley & Sons, Inc., N.Y. Almavist & Wiksell, Stockholm 344 pp. (1963). The faunal history of Newfoundland. Opus. Ent. Sup. 23. 112 pp. (1965). Skaftafell, Iceland, a living glacial refugium. Oikos Sup. 6. 142 pp. MACARTHUR, R. H. and E. O. WILSON (1967). The Theory of Island Biogeography. Princeton Univ. Press, Princeton, New Jersey. 203 pp. MEDIOLI, F., D. J. STANLEY, and N. JAMES (1967). The physical influence of a Paleosol on the morphology and preservation of Sable Island, off the coast of Nova Scotia. Proceedings VII Congress on Quaternary Soils 9: 246-259. MUNROE, E. (1956). Canada as an environment for insect life. Can. Ent. 88: 372-476. SAUNDERS, W. (1902). Experiments in tree planting on Sable Island. Jn Report of the Director. Experimental Farms Reports for 1901. 1-2 Edward VII, sessional paper No. 16: 62-77. 29 SMETANA, A. (1965). Staphylinini und Quediini (Col., Staphylinidae) Von Newfoundland, Sudost-Labrador und Nova Scotia. Acta. Ent. Fennica 20: 1-60. STANLEY, D. J. and A. E. COK (1967). Sediment transport by ice on the Nova Scotial Shelf. Trans. Nat. Symposium, Marine tech. Soc. pp. 109-125. WHITMORE, F. C., K. O. EMERY, B. H. S. COOKE, and D. J. P. SWIFT (1967). Elephant teeth from the Atlantic Continental Shelf. Science 158: 1477-1481. 30 THE ODONATA AND ORTHOPTERA By J.E.H. MARTIN * ODONATA During the summer of 1967 five species were collected and found breeding on the island. Nymphs of all species except Sympetrum corruptum (Hagen) were obtained, and teneral adults of this species were collected on the margin of the freshwater lake near the West Light. ZYGOPTERA Family COENAGRIIDAE Enallagma civile (Hagen) Adults: 216 159, 1-18 July 1967; 156 119, 11-18 September 1967. Nymphs: Numerous nymphs collected on the above dates. Range: Arizona to Florida, south to W. Indies and Colombia, north in Canada to Nova Scotia and New Brunswick, southern and western Ontario, Manitoba, and Saskatchewan. This species has not been collected in Quebec, and Walker (1953: 208) suggests it has entered the Maritime Provinces from the New England states. Nymphs were abundant in all the freshwater ponds and lakes. Ischnura verticalis (Say) Adults: 886 699, 1-18 July 1967; 26 19, 13 September 1967. Nymphs: Numerous nymphs collected on the above dates. Range: An eastern species, Newfoundland to Manitoba, south to South Carolina and Texas. Walker (1953: 260) states that this species is “rather a feeble flyer and seldom, if ever, ventures over the open water of a pond or stream.” This was the most abundant species on the island, nymphs being taken in all freshwater ponds and lakes. ANISOPTERA Family AESHNIDAE Anax junius (Drury) Adults: 29, emerged Ottawa, Ontario, 1 August 1967, nymphs collected 15 July 1967. Nymphs’ 36 69, collected same date. Range: A wide-ranging migratory species; occurring throughout North America, Alaska to Panama, West Indies, Hawaiian Islands, and west coast of Asia. Nymphs were collected from only one pond located near the Meteorological Station. *Entomology Research Institute, Canada Department of Agriculture, Ottawa. 31 Family LIBELLULIDAE Sympetrum corruptum (Hagen) Adults: 29, 15-16 July 1967. Nymphs: Not observed. Range: Southwest Ontario to British Columbia, south to Baja California and Mexico, east to Florida, and north to New York. This species has not been collected in Quebec or the Maritime Provinces and was uncommon on the island. Sympetrum rubicundulum (Say) Adults: 5d, 11-15 July 1967. Nymphs: Many, on the above dates. Range: Eastern Canada, south to Virginia, west to Utah, and north to South Dakota. Nymphs were numerous in all freshwater ponds and lakes. ORTHOPTERA The only orthopteroid insect collected during the three visits to the island in 1966 and 1967 was Melanoplus sanguinipes (Fab.). This species, reported by Piers (1896: 216) as Melanoplus atlantis (Riley), is probably the first insect recorded from Sable Island. One male and three females taken on September 23, 1894, were received by Piers from the Marine and Fisheries Department, which reported that these insects had become “a frightful scourge” on the island. Piers, from information received from the superintendent, stated that locusts had not been observed on the island prior to 1891, but because of exceedingly dry summers they attained devastating numbers by 1895. Garden crops were destroyed, and damage reached such proportions that it was necessary to import quantities of hay to feed the ponies. Piers (1918: 293) made further reference to the outbreak and from correspondence with the superintendent of the island reported that on June 11, 1896, millions of young locusts appeared; fortunately, a cold wet period followed, which presumably destroyed the young insects. Piers stated locusts had not been seen on the island since 1896 and suggested that in 1891 they had flown to the island, aided by the prevailing southwest wind. The possibility that the species did in fact exist unnoticed on the island prior to 1891 and following 1896 cannot be excluded. Family ACRIDIDAE Melanoplus sanguinipes (Fabricius) Adults: 206 209, 11-15 September 1967. Nymphs: Numerous, 1-8 July 1967. Range: A wide-ranging species in Canada, United States, and Alaska. The species is migratory at times; extensive flights have been reported by Corkins (1922: 1-4) who observed swarms from an airplane at altitudes of 500-800 feet with a few individuals at 1,650 feet. REFERENCES CORKINS, C. L. (1922). Notes on the migration of Melanoplus atlantis Riley in northern North Dakota in 1920. Can. Ent., Vol. 54, 1-4. PIERS, H. (1896). Preliminary notes on the Orthoptera of Nova Scotia. Trans. N.S. Inst. Sci., Vol. 9, 208-218. (1918). The Orthoptera of Nova Scotia; with descriptions of the species and notes on their occurrences and habits. Proc. and Trans. N.S. Inst. Sci., Vol. 14, 201-356. WALKER, E. M. (1953). The Odonata of Canada and Alaska. Vol. 1. Univ. of Toronto Press. 33 AMPHIPOD AND ISOPOD CRUSTACEANS By E. L. BOUSFIELD During recent faunistic surveys of Sable Island, several lots of amphipod and isopod crustaceans were collected in terrestrial and shallow-water habitats by Dr. A. H. Clarke, Jr., National Museum of Natural Sciences, and by Mr. J. E. H. Martin, Entomology Research Institute, Ottawa. As these collections had been made coincidental to more thorough searches for molluscs and insects, respectively, only a small fraction of the total peracaridan crustacean fauna of the island, particularly of sand-burrowing and infaunal types, is probably represented here. Although all species recorded in this short list range more or less widely in coastal regions of eastern Canada, all records are apparently new to the island, and a few are of zoogeographical significance. The marine amphipod Gammarus annulatus of southern coastal affinities is authentically recorded for the first time from Canada. The presence of the freshwater amphipod Crangonyx richmondensis richmondensis in sand dune ponds tends to support previous views (e.g., Bousfield 1962, 1967) that some elements of the Sable Island fauna are remnants of populations widely distributed over an island archipelago that existed in offshore regions of Nova Scotia and southern Newfoundland during glacial and immediate postglacial (to Hypsithermal) times, but that have since been almost completely obliterated by changing sea levels and marine erosion. The terrestrial oniscoidean isopods were undoubtedly introduced by human agency from Europe (see Palmen 1951) but the semi-terrestrial talitrid amphipods are marine dispersed and are American endemics. LISTS OF SPECIES AMPHIPODA GAMMARIDEA Family GAMMARIDAE 1. Gammarus oceanicus Segerstrale Wallace Lake, 15 June 1967, JEHM coll. — 1 © (br. Il), 3 subadult 99. A common intertidal and shallow-water euryhaline and eurytopic species of the North Atlantic region, more common along rocky and fucoid shores than on sand. 2. Gammarus annulatus Smith Wallace Lake, 15 July 1967, 16 August 1967, JEHM coll. — about 60 specimens, mostly ovig. 9? but including subadult males, all with well-developed characters diagnostic of this species. Pond B, 14 July 1967, JEHM coll. — 1 6, 9 99 ovig. A pelagic and epibenthic species of sandy costal marine areas of New England, from Long Island Sound north to New Hampshire (Bousfield, in prep.). Previous records of this species from Canada are referable to the closely related inshore and estuarine species Gammarus lawrencianus Bousfield. 34 3. Crangonyx richmondensis richmondensis Ellis Sta. 845, sand dune pond % mi. S. of Meteorol. Stn. (hardness 45 ppm) 8 June 1966, AHC coll. — 1 ? ovig., 9 juv. Freshwater pond, 15 July 1967, JEHM coll. — 1 © (ovig.) 2 29 (br. I and II), 7 juv. The nominate subspecies is known from American Atlantic drainages from Georgia north to Nova Scotia and Newfoundland (Bousfield 19584). Previous records are from acidic or humic permanent ponds and outflows, mainly in winter-mild igneous-rock areas. The animals live close to shore, in shallow water, under debris and among roots of aquatic vegetation. This subspecies complex has a surprisingly wide distribution in coastal areas of North America but is virtually absent from the continental interior (Mississippi and Hudson Bay drainages). However, the very disjunct nature of this distribution and the relatively restricted ecological preference and physiological tolerance of the component subspecies suggest that natural dispersal is accomplished primarily by migration through headwater bogs of adjacent and (or) interconnecting watersheds. Its presence on Sable Island, therefore, is strongly suggestive of former land connections, or near connections, With mainland Nova Scotia, from which it has been increasingly isolated by postglacial submergence and erosion of surrounding archipelagos (see Bousfield 1962, 1967). Family HY ALELLIDAE 4. Hyalella azteca Saussure Sta. 845, June 1966, AHC coll. — about 400 specimens, in 2 lots. Sta. 848, Lily pond near West Light (hardness 10 ppm) 8 June 1966, AHC coll. — about 200 specimens (incl. dd and 99 ovig.). Freshwater pond, 7 July 1967, JEHM coll. — 35 specimens. Freshwater pond, 15 July 1967, JEHM coll. — 13 specimens, mostly 99 ovig. An extremely hardy and eurytopic freshwater species, widely distributed in North America south of the tree line. Its occurrence on all coastal islands having bodies of permanent fresh water is very probably attributable to wind dispersal. The present material has two dorsal pleosomal spines or mucronations. Family TALITRIDAE S Talorchestia longicornis Say Sta. 841, North Beach near West Light. 7 June 1966, AHC pan trap coll. — 7 dd, 9 29, 50 imm. Sta. 851A South Beach, opp. West Light landward side of drift line 150 ft. from water. 13 June 1966, AHC pan trap coll. — 3 dd, 1 9, 5 imm. Sta. 851B South Beach, landward side of drift line., 13 June 1966 AHC pan trap coll. — 10 dd, 3 99 subad., 9 imm. Sta. 851C South Beach — 22 66, 4 99, 18 imm. Shore, 1 July 1967, JEHM coll. — 3 dd, 3 subadult 99. Shore, 14 July 1967, JEHM coll. — 8 dd, 11 99, 3 subadult specimens. 35 This semi-terrestrial species was also collected at Lawrencetown Beach, mainland Nova Scotia, along with Orchestia platensis Kr. on 20 July 1967 by Martin. Talorchestia longicornis ranges from northern Florida to eastern Canada (Chaleur Bay, Long Peninsula, Burin Peninsula) (Bousfield 1958b, in prep.). It occurs more frequently than does T. megalophthalma on protected and estuarine sandy beaches, on steeper slopes, and farther from shore on the dunes. 6. Talorchestia megalophthalma Bate Sta. 841, North Beach, 7 June 1966, AHC pan trap coll. — 16 dd, 4 99. Sta. 851B, South Beach, 13 June 1966, AHC pan trap coll. — 1 d. Sta. 851C, South Beach drift line. 13 June 1966, AHC pan trap coll. —9 dd, 2 22, No specimens were collected in 1967. This species occurs more abundantly on open, flat surf-pounded beaches, closer to the water’s edge, and is more northerly in distribution, ranging from Georgia to the north shore of the Gulf of St. Lawrence (Bousfield 19585). Family CALLIOPIIDAE 7. Calliopius laeviusculus Kr. Wallace Lake, 15 July 1967, JEHM coll. —3 dd, 19 ovig. A very common pelagic and epibenthic coastal marine species; amphi-Atlantic — in North America south to the Middle Atlantic States. ISOPODA VALVIFERA Family IDOTEIDAE 1. Chiridotea caeca (Say) Wallace Lake, 15 July 1967, JEHM coll. — 20 specimens, mostly dd, a few 99. Ibid. 16 August 1967 — about 30 specimens, mostly 99. A species of sandy beaches, Florida north to Cape Breton Island. Common on open Atlantic beaches of mainland Nova Scotia, burrowing in the intertidal sands (Bousfield 1956). ISOPODA ONISCOIDEA Family ONISCIDAE 1. Oniscus asellus L. Sable Island, Gen. coll., terrestrial. AHC., June 1966 — 2 adults specimens. A European species, fairly widely introduced in North America, but previously recorded in eastern Canada only from ice-free, winter-mild coastal localities of Nova Scotia and Newfoundland (Palmen 1951; Bousfield 1962). 2. Porcellio scaber Latr. West Light, 13 June 1966, AHC pan trap coll. — 1 d, 1 imm. Sable Island, 17 July 1967, JEHM — about 100 specimens in 3 lots. 36 A hardy, very eurytopic European species, widely distributed in North America through human agency. Common in mainland Nova Scotia and Newfoundland, able to survive away from human habitation (Palmen 1951). 3. Porcellio sp. (dilatatus Brandt? ) Sable Island , June 1966, AHC gen. coll. — 1 9. 4. Trachaeoniscus (Trachelipus) rathkei (Brandt) Sta. 844, near West Light, in fields, under boards, 7 June 1966, AHC coll. — 299 ovig. Sable Island, June 1966, AHC gen. coll. (terr.) — 25 adult specimens. Sable Island, 17 July 1967, JEHM — about 50 specimens in 3 lots (with Porcellio). A very hardy European species with distribution in North America similar to that of P. scaber, Although common in mainland Nova Scotia, it is rare in New- foundland. It is expected to survive climatic conditions of the Newfoundland region and thus probably spread further (Palmen 1951). REFERENCES BOUSFIELD, E. L. (1956). Malacostracan Crustaceans from the Shores of Western Nova Scotia. Proc. N. S. Inst. Sci. 24 (1): 25-28, figs. (19582). Fresh-water Amphipod Crustaceans of Glaciated North America. Can. Field-Nat. 72(2): 55-113. (1958b). Distributional Ecology of the Terrestrial Talitridae (Crustacea; Amphipoda) of Canada. Proc. Tenth Int. Congr. Ent., vol. 1 (1956): 883-898. (1962). Studies on Littoral Marine Arthropods from the Bay of Fundy Region. Bull Nat. Mus. Canada No. 183:42-62. (1967). Postglacial Dispersal Patterns of Littoral Marine Mollusks and Crustaceans in Eastern Canada. Ann. Repts. for 1967, Amer. Malacol. Union: 42-44 (Abstract). in prep. The Shallow-water gammaridean Amphipoda of New England. Systematics-Ecology Program, Marine Biol. Lab., Woods Hole. PALMEN, ERNST (1951). A Survey of the Oniscoidea (Isopoda Terr.) of Newfoundland. Ann. Zool. Soc. Bot. Fenn. ‘““Vanamo”’ 14(1): 1-27. 37 THE FRESH AND BRACKISH WATER FISHES By D. E. McALLISTER INTRODUCTION This paper reports three species of fishes from Sable Island, a small, low, sandy isolated island 100 miles east of mainland Nova Scotia on the Atlantic coast of Canada. The author is unaware of any previous publications on fishes of Sable Island. The island is a crescent-shaped sand shoal, 21 miles long and up to 1 mile wide, lying near the edge of the continental shelf. The strong surf pounding the shores prevents collecting in the sea, although one egg case of the small skate Raja erinacea Mitchill 1825 was collected from the thousands found washed up about 0 to % mile east of the West Light on the North Beach, 7 June 1966. Collections were made, however, in some of the small and moderately sized ponds found on the island. The details on the collection sites are presented in Table I. METHODS Counts and measurements of paired structures were made on the left side. The last two rays of the dorsal and anal fins were counted as one in the Cyprinodontidae and separately in the Gasterosteidae. The specimens are cata- logued in the fish collection of the National Museum of Natural Sciences, National Museums of Canada, Ottawa. The catalogue numbers are given in the table and in the species accounts. Family CYPRINODONTIDAE Fundululus heteroclitus heteroclitus (Linnaeus, 1766) mummichug choquemort Collections: NMC66-219(3); NMC66-216(27); NMC67-509-A(15). Standard lengths 21.3 to 63.8 mm. Meristic characters: D 12-13, x=12.3; A 10-12, x=10.9; P 18-20, x=18.6; scales above lateral line 6-7, x=6.2; lateral line scales 36-40, x=38.1; G.R. 1-2 + 6-8=8-10, x total number=8.5. Proportions: Body depth comprised 3.8-5.1, x=4.68 and caudal peduncle depth 7.4-9.5,x=8.60, of standard length. Caudal peduncle depth comprised 2.1-4.0, x=2.96, in distance from dorsal insertion to midlateral base of caudal fin. The dorsal fin index was (3.6)4.2-21.8, being the predorsal length minus the postdorsal length (mean not given, since this factor varied so greatly with size). Except for two specimens, most (being females or young) have long thin vertical bars on the sides; the two exceptions have thicker bars, as though they were formed by the union of two ordinary bars. A black spot is found just in front of the dorsal fin. The fins are without spots. 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