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National Museum of Natural Sciences

Publications in Zoology, No. 4

Fauna of Sable Island
and its Zoogeographic
Affinities

National Museums of Canada

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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

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.

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

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.

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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

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0
.02
0
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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

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.

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.

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.

34.

35?

36.

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.

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

59;

60.

OL:

62.

63.

64.

65.

66.

67.

68.

69.

70.

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.

13.
74.

15.
76.

78.

19.

80.

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),

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.

103.

104.

105.

106.

107.

108.

109.

110.

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.

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

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

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’

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

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.

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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.

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

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.

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.

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.

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.

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.

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.

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.

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.

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.

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. The peritoneum is dusky or
black.

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The specimens agree in most characters with Fundulus heteroclitus, but in one
character they tend toward Fundulus diaphanus. Differential characters are
reported by Hubbs ef al. (1943), Brown (1957), and Scott and Crossman (1964).
The specimens agree with heteroclitus in the low number of dorsal rays, the high
gill raker count, the dark instead of lightly specked peritoneum, and the short
snout. In the lateral line scale count, the dorsal fin index (of Scott and Crossman
(1964)), and the body depth, the specimens are intermediate, alchough tending
towards heteroclitus. The caudal peduncle depth is also intermediate, but tends
towards diaphanus. Thus four characters are typical of heteroclitus. Four other
characters are intermediate, three being closest to heteroclitus and one to
diaphanus. The differences from typical heteroclitus may have resulted from past
hybridization with diaphanus; a hybrid between the two species has been
reported by Hubbs ef al. (1943). Recent hybridization would appear unlikely,
however, since F. diaphanus is a freshwater species that only occasionally
penetrates brackish water. The differences may also be due to population
divergence. The latter would not be unexpected, as the Sable Island populations
are about 100 miles distant from the closest mainland populations in Nova
Scotia, and as Bigelow and Schroeder (1953: 163) report “So closely, indeed, do
they hug the shore that a line drawn 100 yards out from land would probably
inclose practically all the mummichugs in the Gulf of Maine.” Brown (1957), in
a review of the genus, declined to recognize a northern (continental) subspecies,
considering the diagnostic characters to be clinal, and added that a detailed study
of variation in this species was needed. Taxonomic recognition of Sable Island
populations would seem premature until such a variational study is undertaken.

Family GASTEROSTEIDAE

Pungitius pungitius (Linnaeus, 1758) ninespine stickleback
épinoche à neuf épines

Collections: NMC66-124(2); NMC66-216(2); NMC66-124-A(28)
NMC66-508(3). Standard lengths 9.4 to 41.5 mm.

Meristic characters: Dorsal spines VIII-XI, x=9.5; dorsal rays 9-12, x= ie
19-11, x=19.7; VI1,x 11.0; P 10, x=10.0; lateral plates 1-4, x=1.9; G.R. 1-3 +
9-11 = 11-13, x total number 11.5.

Proportions: Pectoral fin length / pelvic spine length 1.3-2.0, x=1.61.

McPhail (1963) described two forms of ninespine stickleback in North
America, one coastal in distribution, probably originating during glaciation in the
Bering Sea area, the other inland, probably originating in freshwaters of the
Mississippi basin. In the present specimens, two characters, plate number and
dorsal spine number, are intermediate but tend towards the freshwater form; the
other two characters, pectoral spine length and gill raker number, are definitely
characteristic of the coastal form. The weight of the characters, including the
least variable one, gill raker number, is thus towards the coastal Bering form.
This is indeed what would be expected if Sable Island was populated from the
Atlantic provinces and the New England states, which are inhabited by the
coastal Bering form.

Apeltes quadracus (Mitchill, 1815) fourspine stickleback
épinoche à quatre épines

Collections: NMC67-509(5). Standard lengths 19.6 to 45.0 mm.

Meristic characters: D I-III], 110-13, x=2.8, 111.6;A 19, x=9.0; V 12; P 11,
x=11.0; G.R. 7-9, x=7.5.

Proportions: Length of the first dorsal spine comprised 9.1-15.4, x=11.4, and
pelvic spine length comprised 6.9-9.1, x=8.1, of standard length.

Krueger (1961) reports that freshwater populations of fourspine sticklebacks
have strong modes of 5 dorsal spines (all spines counted) whereas those of high
salinities show strong modes of 4 spines. Although the present sample is not
adequate, the few specimens present have counts of 3 or 4, which are typical of
the marine populations. This might be expected if Sable Island was populated
from coastal populations,

DISCUSSION

The question naturally arises as to how these fishes came to Sable Island,
which is 100 miles from the nearest populations on the mainland. As the three
species are euryhaline, salt water would not appear to constitute a barrier. Their
ability to cross narrow ocean straits is demonstrated by their presence on
Newfoundland (Scott and Crossman 1964), the Magdalens, and on Anticosti
Island (Cox 1921; Leim and Scott 1966; McPhail 1963; Schmitt 1904), although
Apeltes quadracus has not yet been reported from the latter island. These islands
are not, however, more than 50 miles from the closest populations of the species.
The shore-hugging distribution of Fundulus heteroclitus has already been men-
tioned. But Fundulus heteroclitus has managed to populate Bermuda, some 600
miles off eastern United States, where it forms an endemic subspecies F. h.
bermudae Ginther (Brown 1957). These facts show that despite their shore-
hugging habit they are still capable of reaching offshore islands. Bigelow and
Schroeder (1953: 308) note of Pungitius pungitius, “Since the range of the
nine-spined stickleback hardly touches the open waters of our Gulf...” and of
Apeltes quadracus (p. 312) “But it is so much more closely restricted to
estuarine situations than is its three-spined relative (p. 310) that we have never
taken it in our tow nets nor do we find a single record of it in the open sea.”

Yet despite these habits, since they have populated an island 50 miles away, it
is not difficult to conceive their reaching an island 100 miles away. Even if the
adults are shore-hugging in habit, it is within the realm of possibility that either
adults or young were carried offshore by unusual storms or currents. It is
possible that colonization might not have been from Nova Scotia, although this
provides the closest landmass, but from further south with the immigrants being
carried by the Gulf Stream.

Another explanation is possible. Clarke (1967) and Clarke et al. (1967)
suggest that Sable Island was much wider during the Hypsithermal period and
that portions of the continental shelf between the mainland and Sable Island
may also have been above sea level during that period and later. Larger size
would increase the probability of drifting specimens meeting the island, and

intermediate islands would provide stepping stones to Sable Island. There is
presently insufficient data to determine which hypothesis is most favourable.

The forms of Fundulus heteroclitus, Pungitius pungitius, and Apeltes qua-
dracus found on the island are morphologically the saltwater forms. This would
support invasion of the island by coastal forms from the mainland, perhaps by
intermediary islands. No direct connection with the continent is necessary to
explain the presence of these species on the island.

SUMMARY

Fundulus heteroclitus, Pungitius pungitius, and Apeltes quadracus are reported
for the first time from Sable Island, 100 miles off mainland Nova Scotia, in fresh
and slightly brackish ponds. The presence of these species on the island does not
call for a past land connection with the mainland, as they are capable of crossing
stretches of salt water to islands.

ACKNOWLEDGMENTS

The author is grateful to Dr. A. H. Clarke, Jr. Dr. H. F. Howden, and J.
E. H. Martin, who kindly collected fish specimens in the course of other studies.

REFERENCES

BIGELOW, HENRY B., and WILLIAM C. SCHROEDER
(1953). Fishes of the Gulf of Maine. Fishery Bulletin of the Fish and Wildlife Service, 53:
1-577.

BROWN, JERRAM L.

(1957). A key to the species and subspecies of the cyprinodontid genus Fundulus in the
United States and Canada east of the continental divide. Journal of the Washington Academy
of Sciences. 47(3): 69-77.

CLARKE, ARTHUR H.
(1967). Recent and post-Pleistocene mollusca from Sable Island, Nova Scotia. Annual
Reports for 1967 of the American Malacological Union, 11-12.

D. J. STANLEY, J. C. MEDCOF, and R. L. DRINNAN
(1967). Ancient oyster and bay scallop shells from Sable Island. Nature 215 (5106):
1146-1148.

COX, PHILIP
(1921). List of fishes collected in 1917 off the Cape Breton coast and the Magdalen Islands.
Contributions to Canada Biology 1918-1920, (11): 109-114.

HUBBS, CARL L., W BOYD W. WALKER, and RAYMOND E. JOHNSON

(1943). Hybridization in nature between species of. American cyprinodont fishes. Contribu-
tions from the Laboratory of Vertebrate Zoology, University of Michigan (23): 1-21.
KRUEGER, WILLIAM H.

(1961). Meristic variation in the fourspine stickleback, Apeltes quadracus. Copeia (4):
442-450.

LEIM, A. H., and W. B. SCOTT

(1966). Fishes of the Atlantic coast of Canada. Bulletin of the Fisheries Research Board of
Canada (155): 1-485.

McPHAIL, J. D.

(1963). Geographic variations in North American ninspine sticklebacks, Pungitius pungitius.
Journal of the Fisheries Research Board of Canada. 20(1): 27-44.

SCHMITT, J.
(1904). Monographie de L’isle d’Anticosti. Paris, Plon-Nourrit, 367.

SCOTT, W. B., and E. J. CROSSMAN
(1964). Fishes occurring in the fresh waters of insular Newfoundland. Department of

Fisheries, Queen’s Printer, Ottawa, 124.

APPENDIX I
SABLE ISLAND NOCTUOIDEA

E. W. ROCKBURNE and D. F. HARDWICK*

ARCTIIDAE

Diacrisia virginica Fabr.

NOCTUIDAE

Argrotis volubilis Harv.
Aletia oxygala Grt.
**Amathes c-nigrum Linn.
Amathes smithi Snell.
Amphipoea interoceanica Sm.
Apamea finitima Gn.
Apamea inordinata Morr.
Autographa precationis Gn.
Euxoa detersa Wik.

Euxoa messori Harr.

Faronta diffusa Wik.
Lacinipolia lorea Gn.
Nephelodes emmedonia Cram.
*Ochropleura plecta L.

Oligia bridghami G. & R.
Ommatostola lintneri Grt.

Paipema sp. probably nelita Stkr.

*Perodroma saucia Hbn.
*Pseudaletia unipuncta Haw.
Polia pulverulenta Sm.
Rhynchagrotis cupida Grt.
Spaelotis clandestina Harr.

No. of specimens

*Entomology Research Institute, Canada Department of Agriculture, Ottawa.
**Holarctic species, presumed native to North America; remainder endemic to North America.

APPENDIX II
THE FRESHWATER AND TERRESTRIAL ANNELIDA
By G.E. GATES and J. E. MOORE

During the Sable Island faunistic survey work of 1966, three species of leeches
(Hirudinea) and two species of earthworms (Oligochaeta) were obtained. The
former have been identified by Dr. J.E. Moore, Edmonton, Alberta, and the latter
by Dr. G. E. Gates, Bangor, Maine. All three species of leeches are widely
distributed and North American endemic, whereas the two oligochaete worms are
both of European origin and were probably introduced by human agency. All
records appear to be new to Sable Island.

SPECIES LIST

HIRUDINEA

The following material was obtained from Lily Pond, near West Light, Station
848, 10 June 1966, A.H. Clarke coll.

Haemopis marmorate (Say) — 1 adult specimen.

Helobdella stagnalis (L.) — 11 immature specimens (in 2 lots).
Erpobdella punctata (Leidy) — 1 adult specimen.
OLIGOCHAETA

The following material was obtained during general terrestrial collecting by A.H.
Clarke, June 1966.

Lumbricus rubellus Hoffmeister 1843 — several specimens (NMC 1159, 1160).
Dendrobaena sp. (poss. rubida Sevigny) — 2 small specimens (NMC 1161).

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