The Faunal Connections betzueen Europe and North America
By Carl H. Lindroth

The Faunal Connections

between

Europe and North America

BY

CARL H. LINDROTH

JOHN WILEY & SONS, INC., NEW YORK
ALMQVIST&WIKSELL- STOCKHOLM

© 1957
Almqvist & Wiksell / Gebers Forlag AB, Stockholm

PRINTED IN SWEDEN BY

Almqvist & Wikselh

BOKTRYCKERI AKTIEBOLAG
UPPSALA 1957

To
My Wife

Foreword

,^o^« "♦<».. -^'.

1 first encountered Carl Lindroth twenty-six years ago, by letter, when he wrote
me in an effort to establish the identity of European and American species of
Micralymma. The next year (1931) he published his fine book on the insects of
Iceland, and since then I have followed his work with increasing interest and ad-
miration. He has pursued much the same line of research that I have done — study
of carabid beetles, not only their taxonomy but also special problems of their
ecology, evolution, and zoogeography— so that I have had continual opportunities
for direct and critical judgment. His studies of North European Carabidae cul-
minated in 1949 with publication of the third part of his great work on "Fenno-
skandischen Carabidae". In 1948 he began to turn his attention to America,
planning a series of collecting and study trips which have taken him to Newfound-
land for two summers, to the M. C. Z. (the Museum of Comparative Zoology
at Harvard, where I work) for several months during the winter of 1950-51, and
elsewhere.

The present book on European and American faunal connections is important
for two reasons. It is important because it is by Lindroth. He knows both sides
of the subject (the European and American sides) better, I think, than anyone else
has ever done; and he brings to the subject important new data derived from his
first-hand knowledge of insects, especially carabid beetles. I know— and this is
the reason I am writing this foreword— that much that has been written in the
past about relationships of European and American insect faunas is untrustworthy,
marred by misidentifications and other errors, while what Lindroth writes expeci-
ally about Carabidae can be trusted. His taxonomy stands the ultimate test: it
describes situations as they really are in nature.

The other reason why this book is important is the growing importance of the
north, which is drawing increasing numbers of biologists, other scientists, and
"practical" people. This book describes, compares, and traces the history of the
two best-known northern faunas, those of Europe and of eastern North America.
The book may have no "practical" applications, but it is an important contribution
to basic knowledge of the north.

P. J. Darlington, Jr.

Curator of Insects

Museum of Comparative Zoology

Harvard University.

Contents

Foreword by Dr. P. J. Darlington, Jr 7

Author's preface 12

Introduction 13

CHAPTER I

List of land and freshwater animal species common to Europe
and North America

Introduction 15

Mammals, p. 17 - Birds, p. 21 - Fishes, p. 38 - Coleoptera, p. 40 - Lepidoptera,
p. 66 - Diptera, p. 82 - Hymenoptera, p. 84 - Odonata, p. 91 - Orthoptera
Saltatoria, p. 92 - Araneae, p. 93 - Chilopoda, p. 108 - Diplopoda, p. no -
Isopoda Terrestria, p. 112 - Mollusca, p. \i^- Lumbricidae, p. 123 ... 17

Summary 125

Bibliography 126

CHAPTER II

The human transport of animals across the North Atlantic

The five criteria of an introduced species 135

Westward transport. From where did they come? 143

The fauna of Newfoundland as an illustration 145

The European species, p. 145 - The Newfoundland trade. Historical review,
p. 154 - The ballast traffic, p. 157.

Application to other parts of North America 168

Nova Scotia, p. 168 - Prince Edward Island, p. 169 - New Brunswick, p. 169
- New England, p. 169 - The Pacific Northwest, p. 170.

Investigations on ballast-places in Southwestern England 172

Account of collecting, p. 173 - List of animals collected, p. 186 - Carabid
beetles, p. 192 -Plants, p. 196 -The selection of animal species which man-
aged to cross the Atlantic, p. 198.

Eastward transport 21 1

The reasons for its poor effect, p. 214 - Another explanation, p. 216.

The economic importance of introduced animals 218

On the minimum size of a viable population 223

Bibliography 225

CHAPTER III

The true, pre-human relationship between the Palaearctic
and Nearctic faunas

The so-called Amphiatlantic species 230

Doubtful cases, p. 233 - "Westarctic" species, p. 237 - The American element
of the British flora, p. 240.

The chance of spontaneous transatlantic dispersal at the present time 242

The theories of earlier transatlantic land-connections 253

The Iceland-Greenland bridge, p. 253 -Affinities in the south, p. 280 -Wege-
ner, pro and contra, p. 284.

Circumpolar animals 286

The fossil evidence 306

Bibliography 313

Conclusions 322

Index 324

73810

FIG. I. River Oulankajoki in Kuusamo, NE Finland.
(Photo A. V. Lumiala)

FIG. 2. Doctors Brook opposite St. John Island, NW Newfoundland.
(Photo Ernst Palmen)

Author's preface

1 he incentive to this book was given by entomological fieldwork in Iceland
(1926 and 1929) and Newfoundland (1949 and 1951).

The former island has a veritable key position in the northern Atlantic, situated
almost three times as far from the European mainland (Norway) as from Greenland
but in spite of this inhabited by an almost purely European fauna. This fact leads
to the assumption that a considerable part of the Icelandic fauna immigrated from
Europe in an interglacial period when land had a greater extension in the North
Atlantic. Parts of this faunal element apparently even succeeded in colonizing
Greenland.

An attempt was made to investigate the insect fauna of Newfoundland from the
same point of view, i.e. to discover a "European" element in the fauna of this
island too. The results obtained were unexpected, in more ways than one, and it
soon became clear that a satisfactory understanding of the faunal history of the
islands in the North Atlantic called for a general faunistic survey of the continents
of Europe and North America, at least for selected, sufficiently investigated and
otherwise suitable groups of animals.

From this enlarged amount of data, the attempt to explain the trends of animal
distribution in the islands considered was transformed into a comparison between
the Palaearctic and the Nearctic terrestrial fauna as a whole. It was also necessary to
consider the results obtained by botanists who, in biogeography, are often ahead
of zoologists, to benefit by palaeozoological evidence (when available), and—
finally— in order to understand the importance of dispersal of animals by human
agencies, to get an idea of the North Atlantic trade during the last four centuries.

This was a diversified and fascinating study, but dangerous at the same time.
It is perhaps to be regretted that a specialist in Carabid beetles, like the cobbler,
is not always wise enough to stick to his last.

Introduction

A Scandinavian biologist travelling in Eastern Canada is at once struck by the
close affinity of the nature of this country with that of his native land. Not only are
the soft, levelled profiles of the landscape similar, as are the heavily ground rocks,
the gentle hills of moraine, the countless lakes and ponds, the wide areas of bogs and
fens— all vestiges of the simultaneous periods of glaciation— but the main stock
of native animals and plants seems identical too.

This far-reaching coincidence is indisputable— and yet, largely fictitious. The
specialist in zoological or botanical taxonomy, after a closer comparison, in most
cases will find that the European and the North American representatives of the
"species" after all are not quite identical. Whether he is inclined to regard them
as different subspecies or prefers a specific separation, is often a matter of judgment
only.

An excellent illustration of this is given by arboreous plants. There is no dif-
ficulty in using a common trivial name for almost any kind of tree occurring in
northern regions of the two continents: in both of them grow alder, ash, aspen,
birch, elm, hazel, maple, oak, pine, rowan, spruce, and several others; but no
indigenous kind of tree is identical on both sides of the Atlantic— provided
the common juniper {Juniperus communis) is not counted as a tree. With the
possible exception of one of the birches {Betula tortuosa) and perhaps one alder
{Alnus incana), botanists even regard all of them as different species.

The number of "Eur-American" species (the word used here regardless of a
possible occurrence in Asia) may be greater within any of the two following groups
of organisms:

(a) arctic or subarctic animals and plants which, on the whole, often tend to
a circumpolar distribution;

(b) animals and plants at a lower evolutionary stage, e.g. many insects, arachnids,
tardigrades, "worms", rotatorians, etc.; mosses, lichens, fungi, etc. These may
in part be "older" as species and thus have had more time at their disposal for
distribution. Due to their generally reduced size, often combined with the power

14 INTRODUCTION

of asexual reproduction, they may in part be more suited for a passive dispersal
across the sea.

The following attempt to estimate the faunal exchange between Europe and
North America had to be based on suffficient comparative knowledge of the
taxonomy of the forms occurring in the two continents. Unfortunately, such a
happy state of affairs has been reached in only few groups of animals. European
and American zoologists are too inclined to work independently and a taxonomic
study of a certain species group, to be useful in zoogeography, must be carried out
by a specialist fully acquainted with both of the faunas. An attempt to compile a
"complete" list of terrestrial (and limnic) animals common to Europe and North
America would therefore be a hopeless enterprise. It was necessary to restrict the
task to groups of animals which may be regarded as sufficiently worked through
and to add some cases of special interest from other groups.

Marine animals and animals of the tidal zone were excluded from the investiga-
tion.

The bias of this study is also accentuated by the fact that the discussion often
starts, taxonomically from Carabid beetles, geographically from Newfound-
land. As an apology it may be pointed out that the insect family mentioned pro-
vides an unusually suitable subject to a student in zoogeography: (i) it is numerous
in species, especially in northern regions; (2) these usually possess good specific
and subspecific characters, above all in the male genitalia; (3) many species show
a marked geographic variation; (4) the means of dispersal are unusually varying,
many species being constantly short-winged, bound to the soil and conservative
in distribution, others constantly long-winged, flying more or less regularly,
whereas, in a third category, both long- and short-winged individuals occur which
often results in a characteristic pattern of distribution of the two forms; (5) finally,
the taxonomy of northern Carabidae is tolerably well-known in both continents.
—As regards Newfoundland, this island has a key position in the understanding of
the human transatlantic transport of animals and plants.

Chapter I

LIST OF LAND AND FRESHWATER ANIMAL SPECIES
COMMON TO EUROPE AND NORTH AMERICA

Introduction

This list is pronouncedly selective. It was necessary to restrict its content to
animal groups sufficiently known with regard to taxonomy and distribution. In
spite of this limited scope the groups considered are by no means equally well
suited for the purpose. In some, as in birds, Carabid beetles, and certain Lepido-
ptera, specialists have made direct and careful comparisons between Old and New
World forms, in others a modern revision is still needed and compilations had to
be made from the often rather routine-stamped records of catalogues or from scat-
tered revisions of smaller taxonomic units. I am fully aware that, owing to the
extensiveness of the task, parts of the purely taxonomic literature have been
overlooked, which is much regretted. Also, introduced species which have appeared
more or less sporadically in either or both continents, for instance in hothouses,
have certainly not been listed in full.

It is also evident that the species and subspecies concepts are not used identically
in different groups of animals and direct comparisons of the zoogeographical signi-
ficance of, for instance, a bird and an insect subspecies should be avoided. The
criterion of a species seems to be especially obscure among mammals so that
"species" of that group apparently often correspond to what is called "subspecies"
in others.

Genera and species, under each higher taxonomical unit, are listed in alphabetical
order. Names in brackets are synonyms. Subgeneric names have not been used.

Names in English, according to both American and European practice, if
different, are added for vertebrates only.

The records of distribution are quite summary. Their main purpose is to show
whether the species, in Eurasia as well as in America, is transcontinental or not.
More detailed records, for instance of the occurrence in the British Isles, are particu-
larly given within groups containing species which are introduced into North
America. Subspecies, if any, are listed in geographical order: if numerous sub-
species are described from a limited are, only their approximate number is given.
Subspecies outside the actual area are omitted.

I 6 CHAPTER I

Only animals living (breeding) on the continent of both Europe and North
America were considered. A species occurring only on the Bermudas or on the
arctic islands of Canada as well as in Greenland, is not counted as "American";
similarly, a species confined to Iceland, the Faroes, Jan Mayen, and (or) Spitz-
bergen, not as "European". The British Isles, however, are counted as belonging
to the European continent.

Freshwater animals occurring also in the sea, or regularly descending into the
sea, are omitted.

Not included in the list are furthermore the following types of animals:

a. Domesticated animals. An enumeration is superfluous.

b. Animals introduced and released for the purpose of game, fur, or fishery.
Examples: American Mink (Mustela vison Schreb.), Muskrat {Ondatra [Fiber]
zibethica L.; fig. 5), and Rainbow Trout {Saltno irideus Gibb., s.l.), in Europe;
European Hare {Lepus europaeus Pall.), and European Saibling [Salvelinus al-
pinus L.) in North America.

c. Animals used in "biological control" of noxious insects, plants, and
other pests. Besides instances mentioned below (p. 121 a.f.), leaf-beetles of genus
Chrysomela (Chrysolina) were imported to California against the European weed
Hypericum perforatum (St. John's Wort).

d. "Pet animals", imported and released, for instance in order to still the
nostalgia of emigrants. Examples: Starling {Sturnus vulgaris L.; fig. 4) and
English or House Sparrow (Passer domesticus L.), in North America; the American
"Robin" (Turdus migratorius L.), in Great Britain.

e. Animals imported for study or fancy and unintentionally escaped. Example:
the Gypsy Moth (Lymantria dispar L.) 1868 or 1869 in North America, and prob-
ably several freshwater animals, released by aquarium fanciers.

f. Unintentionally imported animals which have not become established in
the new continent. Most of them have been found in ports or in wares immediately
after these have been unloaded, especially on the quarantine stations. Therefore
the majority of cases reported by Kraepelin (1901) from Hamburg have been
neglected whereas species found in hothouses (Kew Gardens, &c.) were considered.
Several similar cases, however, may have been overlooked.

Summarising, the list below tries to give, within the animal groups considered,
all species common to Europe and North America, and breeding (or supposed to
breed) in both continents, which are not purposely brought across by man in
either direction.

Species believed or stated to be unintentionally introduced into either or
both continents are indicated by a cross (f), in doubtful cases put in brackets.

List of land and freshwater animal species

17

The author is anxious to receive any addition to or correction of the list, which
can be sent under the address of: Zoological Institute, University, Lund, Sweden.

Mammalia — Mammals

The list was largely compiled from Anderson (1946), Ellerman & Morrison-Scott
(1951), and Burt & Grossenheider (1952).

There is great confusion among mammologists concerning the definition of
species and subspecies, the North American ones (e.g. Anderson, 1946) as a rule
being inclined to assign specific value to rather small taxonomic units. Unfortu-
nately no thorough comparison has been made between the Palaearctic and the
Nearctic mammal fauna and the relations indicated below are therefore in some
cases merely suggestive, especially in the Rodentia, where the opinions of spe-
cialists largely disagree.

The following Old World Rodentia, which according to Ellerman & Morrison-
Scott (195 1) "doubtless", "probably" or "perhaps" are represented also in North
America, have been omitted pending further evidence: — Clethrionomys glareolus
Schreb., Marmota marmota L.^, and Microtus agrestis L.

Insectivora

? SOREX ARCTicus Kerr^
(Arctic Shrew and Tundra Shrew)

sbsp. lapponicus

sbsp. europaeus sbsp. petschorae sbsp. middendorffi

Melander

Strog. Ognev

Ognev

N Sweden.

Kola Penins. NE Eur.

Russ. NW Sib.

sbsp. ultimus Allen

sbsp. tundrensis Merr.

/• typ.

NE Sib.

Alas., N. W. Terr.
Carnivora

ALOPEX LAGOPUS L.

(Arctic Fox)

Gr. Bear L.-Nova Scot.,
Labr.

(?) shsp.f uliginostis

sbsp. spitzbergensis f. typ.

sbsp. heringensis

Bechst.

B.-Ham.

Merr.

Icel.

Spitzb. Norw.-Kamch. Bering Isl.

sbsp. innuitus Merr.

sbsp. ungava Merr.

sbsp. groenlandicus Bechst

Alas.-Huds. B.-Baffins L. N Labr.

Ellesmere L., Greenl.

^ Accepted as American by Rausch (1953, p. 116 a. f.).

- This arrangement of northern Sorex is quite tentative. The association of petschorae,
middendorffi, and ultimus with tundrensis was made by Chaworth-Muster, according to

2 — 565597 Lindrolh

i8

CHAPTER I

/. typ.
Eurasian taiga.

CANIS LUPUS L.

(Gray Wolf)

sbsp. albus Kerr
Eurasian tundra.

sbsp. hattai Kish.
Sakh., ? Kuril.

Several more southern sbspp.

FELIS (lynx) lynx I>.

(Lynx)

/■ typ. sbsp. icrangeli sbsp. canadensis

Ognev Kerri

Eur.-Yenisei. Sib. E of Yenisei. Alas.-Labr., Nov,

Scot.
4 more southern sbspp.

Many sbspp.
Alas.-Labr., arct.
isl., N Greenl., in
the mts. S to Mex.

sbsp. subsolanus

Bangs!

Nfld.

GULO GULO L.

(Wolverine; Glutton)

/• typ.
Scand.-Ussuri, Sakhalin.

sbsp. luscus L. (incl. 3 forms)
Alas.-Labr., arct. isl., S to Color.

3 sbspp.
Brit. Isl.
sbsp. arctica Merr.

MUSTELA ERMINEA L.

(Ermine; Shorttail Weasel; Stoat)

/. typ. many sbspp. sbsp. orientalis

Ognev
Fennoscand. N Asia. NE Sib., Jap., Sak-

hal., Kuril,
sbsp. richardsoni Bonap. (and several
other forms)
Huds. B.-Labr., Nov. Scot., Nfld.

? Kamch.; Alas., N Can. W of Huds. B.,
arct. isl. E to Ellesmere Isl., N & NE
Greenl.

/• typ.
Main part of Eur.

MUSTELA NIVALIS L.

(Least Weasel)

sbsp. pygmaea sbsp. namiyei Kur.

Allen

E Sib., Mong., N Jap., Kuril.

Manch.

Several more southern sbspp.

sbsp. rixosa Bangs^
(incl. 3 forms)
Alas.-S Labr.

Ellerman & Morrison-Scott (195 1. P- 49), who themselves refer these forms, as well as
europaeiis and lapponicus, to caecutiens Laxm. Burt & Grossenheider (1952) think that
arcticus and tundrensis are of the same species. If so, the former name has priority.
^ Usually regarded as distinct species.

List of land and freshwater animal species

19

/• typ.

Scand -Stanovoi
(Sib.).

URSUS ARCTOS L.

(Brown Bear)

sbsp. heringianus sbsp. lasiotiis Gray sbsp. mtddendorfii
Midd. Merr.i (incl. many

forms) 2
E Sib., Manch., Mong., Manch., N Pacif. coast of Alas.

Kamch. Jap., Kuril. -N Brit. Col.

About 4 more southern sbspp.

/. typ.

Scand.

Many sbspp.
N Eur., N Asia

VULPES VULPES L.

(Red Fox)

sbsp. beringiana

Midd.

NE Sib., Kamch.

sbsp. fulva Desm.'
(incl. many forms)
Alas.-Labr., Nfld.,
S to the Mexic.
boundary.

/. typ.

Eur.-Baikal.

Ungulata

ALCES ALCES L. (fig. 54)

(Moose; Engl.: Elk)

sbsp. cameloides and pfizenmayeri Zuk.
M.-Edw.*

Sib. E of Yenisei, Mong., Manch.

sbsp. americanus
Clint.* (incl. 4
forms)

Alas.-Nov. Scot.
(Nfld., introduced),
S to Wyom.

CERVUS ELAPHUS L.

/• typ.

Many sbspp.

sbsp. xanthopygus
M.-Edw.

sbsp. canadensis
Erxl.3

(Red Deer)

(Elk or Wapiti)

Sweden.

C Asia, S

Mong., Manch.,

Originally trans-

Siber.

Amur, Ussuri.

american, now dis-

Several other

Europ

junct: Brit. Col.-N

sbspp.

Calif., E to Manit.

S to N. Mex.

^ Due to the many "species" described by Merriam, the taxonomy of N. American bears
is very confused (of. Simpson, 1945, p. 225, footnote).

^ According to Rausch (1953, p. 95 a. f.) even the grizzly {Urstis horribilis Ord), with
its forms, belongs to the species U. arctos L.

^ Usually regarded as distinct species.

* Flerow (1934, p. 141 a. f.) treats cameloides {pfizenmayeri Zuk.) as sbsp. oi americanus,
which he regards as distinct species but, in 1952 (p. 207), he joins them as above. Vide also
Peterson (1952).

20

CHAPTER I

/. typ.

N Eur.

RANGIFER TARANDUS L.' (fig. 53)

(Caribou; Reindeer)

sbsp. platyrhynchus sbsp. sibiricus Murr. sbsp. arcticus Rich.
Vrol. (and 4 other sbspp.) (incl. some minor

forms)

Spitzb.

sbsp. groenlandicus Bor.
W, N & NE Greenl.

Ural Mts.-E Sib.
Sakhal.

Arct. N Amer.: N &
C Alas., Brit. Col.-
N Labr. (caboti
All. 2) and Ellesmere
lsl.{pearyiA\l.^).

/. typ-

N Scand.-Baikal,
Mong.

Rodentia (in widest sense)

CLETHRIONOMYS RUTILUS Pall.

(Tundra or Northern Redbacked Vole)
Many sbspp.

N Asia.

sbsp. amurensis
Schrk.
Amur, Sakhal.

sbsp. dawsoni Merr.^
Alas.-Huds. B., S to Brit. Col.

sbsp. jochelsoni

Allen

Kolyma, Kamch.

/• typ.
N Russ.-N Sib.

DICROSTONYX TORQUATUS Pall

(Collared Lemming)
sbsp. lenae Kerr

Lena-Anadyr.

sbsp. unalascensis

Merr2.

Aleut. Isl.

sbsp. hudsonius Pall.^
N Labr.

sbsp. groenlandicus

Traill^

N. Amer. tundra:

Alas.-Hudson B.,

arct. isl., Greenl.

LEMMUS SIBIRICUS Kerr (obensis Brants)
(Brown Lemming)

/. typ. 3-4 sbspp.

White Sea-W Sib. NE Sib.

sbsp. kittlitzi

Midd.

Kamch.

sbsp. trimucronatus

Rich. 2

Alas. & Brit. Col., E

toNLabr.&Baff.L.

^ According to Flerow (1952, p. 222-247). Jacobi (iQSO bas a different arrangement of
Rangifer.

* Usually regarded as distinct species.

List of land and freshwater animal species

LEPUS TIMIDUS L.

sbsp. hiberniciis sbsp. scoticus Hilzh. /. typ.

Bell (Blue, Mountain, or Varying Hare)

Ireland. Scotland. Scand., N Russ.

21

Many sbspp.

One or two more south-
ern sbspp. in the Old
World.

sbsp. othus Merr.i
(Tundra Hare)
W Alas.

C Russ.-Kamch.,
Sakhal., N Japan,
Kuril.

sbsp. arcticus Ross^ (Arctic

Hare)

NW Terr.-Labr., Nfld.,

arctic isl., Greenl.

Several sbspp.
Alaskan islands.

MICROTUS OECONOMUS Pall.

(Tundra Vole; Root Vole)

sbsp. ratticeps f. typ. sbsp. yakutatensis

Keys. & Bias. (and several sbspp.) Merr.

Scand., Russ., Sib., E to Kamch., Alas.

N C. Eur. Kuril.

sbsp. tnacfarlani Merr.
NW Terr.

MUS MUSCULUS L.

(House Mouse)

Cosmopolitan, probably native of Mediterr., E Eur. & Asia.
Eur. (generally). f N & C Amer. (generally).

RATTUS NORVEGicus Bcrkcnh.
(Norway or Brown Rat)

Cosmopolitan, possibly native of Palaearctic Asia.

t Eur. (generally).

t N & C Amer. (generally).

RATTUS RATTUS L.

(Black or House Rat)

Almost cosmopolitan, native of Asia.
t Eur. (formerly generally). f N Amer. (both coasts).

Aves — Birds

The main source was Peters (I-VI, 1931-48), who gives an explicit, conservative
account with high demands on "the species". For the Passer if ormes, which Peters
did not manage to complete before his death, Hartert (1910-38) and the North

Usually regarded as distinct species.

22 CHAPTER I

American "Check-List" (1931), with subsequent Supplements in "The Auk",
were the foremost substitutes. The British "Handbook" (1945) also rendered
excellent service. Valuable information was obtained from Dr. Gustaf Rudebeck,
Lund.

Only breeding areas were considered.

Gaviiformes

GAVIA (COLYMBUS) ADAMSII Gray

(White-billed Northern Diver; Yellow-billed Loon)
N Finl. (?), N Russ., N. Zeml., Arct. Alas. -Mackenzie, Boothia Penins.

Sib. E to Bering Strait.

GAVIA (cOLYMBUS) ARCTICA L.

/. typ. sbsp. viridigidaris Dwight sbsp. pacifica Lawr.

(Black-throated Diver) (Green-throated Loon) (Pacific Loon)

Scand.- Yenisei, Altai. NE Sib., Alas. Alas.-Baffins L.

GAVIA (cOLYMBUS) STEI.LATA Pont.

(Red-throated Diver or Loon)
Iceland-Scand.-Kamch. Alas.-Nfld.-Greenl.

Podicipitiformes

PODICEPS (cOLYMBUS) AURITUS L.

(Slavonian or Horned Grebe)
/. typ. sbsp. corniitus Gmel.

Icel., Scotl., Scand.-Sakhalin. Brit. Columbia-Magdalen Isl.

PODICEPS (coLYMBUs) CASPicus Habl. (nigrtcollts Brehm)

/. typ. sbsp. californicus Heerm.

(Black-necked Grebe) (Eared Grebe)

Eur., Asia E to Amur. W & NW N. Amer. E to Manitoba.

PODICEPS (cOLYMBUS) GRISEIGENA Bodd.

f, typ. sbsp. holboelli Reinh.

(Red-necked Grebe) (Holboell's Grebe)

N. Eur., W. Sib. E. Asia, N. Amer., E to Ungava.

Procellariiformes

FULMARUS GLACIALIS L. (fig. 27)

/. typ. sbsp. rodgersii Cassin

(Fulmar Petrel; Atlantic Fulmar) (Pacific Fulmar)

NE arct. N. Amer., Greenl., NW Eur. NE Asia E to Bering Sea, Kuriles, islands

oflF Alas.

List of land and freshwater animal species 23

OCEANODROMA LEUCORHOA Vieill.

/. typ. 3 sbspp.

(Fork-tailed or Leach's Petrel) (Other American names)

N Pacific (inch Aleutian); N Atlantic: Pacific coast of N. Amer.
Maine-Brit. Isl.

PUFFINUS PUFFINUS Brunn.

/. typ. sbsp. opisthomelas Coues

(Manx Shearwater) (Black-vented Shearwater)

NW Eur. (incl. Iceland), Macaronesia; Islands oflf Lower Cahf. & Mex.
Bermudas.

Pelecaniformes

MORUS (sula) bassanus L. (fig. 28)
(Gannet)
Brit. Isl., Faeroes, Iceland. NE Canada, Nfld.

PHALACROCORAX CARBO L.,/. typ.

(European Cormorant)

Nova Scot., Labr., Greenl. Iceland, NW Eur. E to White Sea.

Other sbspp. farther south.

Ciconiiformes

CASMERODIUS (EGRETTA) ALBUS L.

/. typ. • sbsp. egretta Gmel.

(Great White Heron) (American Egret)

SE Eur., Asia. From Oregon and N. Carolina S to Pata-

gonia.
Other sbspp. farther south.

NYCTICORAX NYCTICORAX L.

/. typ. sbsp. hoactli Gmel.

(Night Heron) (Black-crowned Night Heron)

C Eur.-Japan, &c. N. Amer. (greater part)-Argent., Hawaii,

PLEGADIS FALCINELLUS L.,/. typ.

(Eastern Glossy Ibis)

S Eur., W & C Asia, Africa. S N. Amer., C Amer.

One tropical sbsp.

24

CHAPTER I

Iceland, Eur.-Kamch.

/• typ.
(European Teal)
Eur.-Japan.

Anseriformes

ANAS (dAFILA) acuta L.

(Pintail)

W & NW N. Amer., Greenl.

ANAS (nETTION) CRECCA L.

sbsp. nimia Friedm. sbsp. carolinensis Gmel.

(Green-winged Teal)
Aleutian Isl. W & NW N. Amer.

/• typ.

ANAS PLATYRHYNCHA L.

(Common Mallard)

sbsp. conboschas Brehm

Eur.-Japan; N. Amer. (doubtful E of Greenland.
Hudson Bay).

ANAS (CHAULELASMUS) STREPERUS L.

(Gadwall)
Iceland, Eur.-Kamch. NW N. Amer. W of Hudson Bay (not Alas.).

/■ typ.
Icel.(?), NE Eur.-Bering Str.; arct
N. Amer.

ANSER ALBIFRONS Scop.

(White-fronted Goose)

sbsp. flavirostris Dalg. & Sc.
W Greenland.

/• typ.
Icel., N Eur.-Kamch.

AYTHYA (nYROCA) MARILA L.

(Greater Scaup Duck)

sbsp. mariloides Vig.
Bering Island.

sbsp. nearctica Stejn.
Aleut.-Hudson Bay.

/• typ.
(Dark-breasted Brent)

N Eur. -Taimyr.

branta bernicla L.
sbsp. nigricans Lawr. sbsp. hrota O. F. Miill.

(Black Brent) (Pale-breasted or American

Brent)
Taimyr-NW arct. Amer. NE arct. Amer., Greenl.,

Spitzb.

/. typ.
Scand.-Kamch.

bucephala (glaucionetta) clangula L.
(Golden-eye)

sbsp. americana Bonap.
Alas.-Maine.

List of land and freshwater animal species

25

CLANGULA HYEMALIS L.

(Long-tailed Duck; Old-squaw)

Icel., Faeroes, Orkney, arct. Eur- Aleut., arct. N. Amer., Greenl.

Bering Str.

/• typ.
(Velvet Scoter)
Scand. -Yenisei.

MELANITTA FUSCA L.

sbsp. stejnegeri Ridgw. sbsp. deglandi Bonap.^

(White-winged Scoter)
Altai-Kamch. Alas.-E Canada.

/• typ.
(Goosander)
Icel.-Kamch., Kuril.

MERGUS MERGANSER L.

sbsp. americanus Cass.
(American Merganser)
Alas.-Nfld.

/• typ.
Icel.-Kamch.; Alas.-Labr.

/. typ.
(Common Scoter)
Icel.-Taimyr.

MERGUS SERRATOR L.

(Red-breasted Merganser)

sbsp. schioleri Salom.
Greenl.

OIDEMIA (MELANITTA) NIGRA L.

sbsp. americana Swain.
(American Scoter)
NE Asia, Alas.-Nfld.

/. typ.
Icel.-N. Zeml.

somateria mollissima L.
(Common Eider)
sbsp. v-nigrum Gray sbsp. sedentaria

Snyd.
NE Sib.; Alas.- Huds. Bay.

Victoria Isl.

sbsp. horealis Brehm
Queb. -Greenl.

somateria spectabilis L.
(King Eider)
Spitzb., N. Zeml.; Kola Penins.- Alas.-Labr. -Greenl

Bering Strait.

SPATULA CLYPEATA L.

(Shoveller)
W Eur.-Ussuri. N. Amer., W of Hudson Bay

sbsp. dresser i Sha rpe
Huds. Bay-Labr.

^ Often regarded as distinct species.

26

CHAPTER I

Falconifortnes

ACCIPITER (ASTUr) GENTILIS L.

(Goshawk)
/. typ. sbsp. buteoides sbsp. albidiis

Menzb. Menzb.

NW Eur.-N Russ. NE Russ., W Sib. NE Sib., Kamch.
(Several sbspp. in other parts of Eur.
and Asia).

sbsp. striatulus Ridgw.
Pacific region of N. Amer.

sbsp. atricapillus

Wils.

Alas.-Nfld.

/• typ.
W Eur.-W Sib.

AQUILA CHRYSAETOS L.

(Golden Eagle)

2-3 sbspp. in

E Asia.

Other sbspp. farther south.

sbsp. canadensis L.
Alas.-Ungava, S to Mex.

/• typ.
N Eur.

BUTEO LAGOPUS Pont.

(Rough-legged Hawk or Buzzard)

sbsp. s.-johannis Gmel.

sbsp. kamtchatkensis

Dement.

N Sib.; NW Alas.

Alas.-Labr., Nfld.

/• typ.
(Hen-Harrier)
Eur.-R. Lena.

CIRCUS CYANEUS L.

sbsp. taissiae But.
NE Sib.

sbsp. hudsonius L.
(Marsh Hawk)
Alas.-Nfld., S to Tex.

FALCO COLUMBARIUS L.

(Pigeon Hawk; Merlin)

sbsp. aesalon sbsp. regulus Pall. sbsp. insignis Clark sbsp. pacifictis

Tunst. Stegm.

Icel.(?), NW Eur. NE Eur., W Sib. N Sib. E Sib.

sbsp. bendirei Swann
Alas.-Sask.

/. typ.
Manit.-Nfld.

Two more southern sbspp.
in N. Amer.

/• typ.

FALCO PEREGRINUS Tunst.

sbsp. calidus Lath. sbsp. pealei Ridgw.

(Peregrine Falcon) (Peale's Falcon)

W Eur.-Ural. W Sib.-Kamch. NW N. Amer.

Many more southern sbspp.

sbsp. anatum
Bonap.
(Duck Hawk)
Alas.-Baff. L., S
to Tex.; Greenl.

List of land and freshwater animal species

27

sbsp. islandus
Briinn.

Iceland.

FALCO RUSTICOLUS L.

(Gyrfalcon)

/. typ. sbsp. uralensis

Gmel.

N Eur. W Sib.-Bering Str.

W Alaska.

sbsp. obsoletus Gmel.
(incl. candicans

Gmel.)

Alas.-Labr.,

Greenl.

/. typ.
Eur.-Kamch.

PANDION HALIAETUS L.

(Osprey)

sbsp. carolinensis Gmel.
Alas.-Labr., Nfld., S to Mex.
Other sbspp. farther south.

/. typ.

Scand.-Kamch.;
Alas.-Ungava.^

Galliformes

LAGOPUS LAGOPUS L.

(Willow Ptarmigan)

sbsp. alexandrae sbsp. leucopterus
Grinn. Tav.

Alas., Brit. Col. Arct. isl. of N.

Amer.

sbsp. alleni Stejn.
Nfld.

LAGOPUS MUTUS Mont.

sbsp. islandorum

Fab.

Iceland.

Several sbspp.
N Asia.

sbsp. millaisi Hart.
(Scottish Ptarmigan)
Scotland.

/. typ.

NW Eur.

sbsp. rupestris
Gmel. s. /.2
(Rock Ptarmigan)
Alas.-Ungava-Baff. Nfld
L., Greenl. (3 sbspp.).

sbsp. hyperboreus
Sundev.

Spitzb., Fr. Jos.
Land.

sbsp. zvelchi Brewst. 2 sbspp.

Greenland.

Gruiformes

GALLINULA CHLOROPUS L.

/. typ. sbsp. indica Blyth sbsp. cachinnans Bangs

(Moorhen) (Florida Gallinule)

W Eur.-Ussuri, N Afr. C, E, & S Asia. Calif.-Ont., S to Mex.

Many sbspp. in southern regions of the Old & New World.

^ The N. American form is referred to sbsp. albus Gmel. in Check List (193 1).
^ The N. American forms are often treated as species rupestris Gmel.

28

CHAPTER I

Charadriiformes, Charadrii

/• typ-

(European Turnstone)
Greenl., Icel., Eur.-Kamch.

ARENARIA INTERPRES L.

sbsp. morinella L.^
(Ruddy Turnstone)
W Alas. Arct. N. Amer.: Mackenzie-Baffins L.

sbsp. faeroensis Brehm
(Faeroe Snipe)

Icel., Faeroes.

/• typ.
Brit. Isl., Scand. -Bering
Str.

/• typ.
(Ringed Plover)
WEur.

CAPELLA GALLINAGO L.

/• typ.
(European or Common
Snipe)
Scand.-Kamch.-Japan.

sbsp. delicata Ord
(American or Wilson's
Snipe)
Alas.-Nfld.

CHARADRIUS ALEXANDRINUS L.

(Snowy or Kentish Plover)

sbsp. nivosus Cass. sbsp. tenuirostris Lawr.

Wash.-Lower Calif. C & SE U.S.A., West Ind.

Many sbspp. farther south.

CHARADRIUS HIATICULA L.

sbsp. tundrae Lowe sbsp. semipalmatus Bonap.^

(Semipalmated Plover)
Spitzb., N Scand.-Chuk- Bering Str.-Baff. L.-Nov.
chi Renins. Scot,

sbsp. psammodroma Salom.^
Baffins L., Greenl., Icel., Faeroes.

sbsp. arctica Schiol. sbsp. schinzii
Brehm

EROLIA (PELIDNA, CALIDRIS) ALPINA L.

/. typ.

N Greenl.

(Southern Dunlin)
E Greenl., Icel.,
Faeroes, W Eur.

(Northern Dunlin)
Scand.-Sib.: Yalmal;
Spitzb., N. Zeml.

sbsp. centralis But.

Yenisei-Yana.

sbsp. pacifica Coues

(Red-backed Sandpiper)

NE Asia; Alas.-Hudson Bay.

EROLIA (arquatella, calidris) maritima Brunn.*
(Purple Sandpiper)
Icel., Faeroes, arct. Eur. isl., Scand.- Arct. isl. of N. Amer.: Melville Isl.-
Taimyr. Ellesm. Isl., Greenl.

^ "A very unsatisfactory' race" (Peters, 1934, p. 271).
2 Often regarded as distinct species.
' Very doubtfully distinguished from forma typica.

* E. ptilocnemis Coues, with its races, from NE Sib. and Alaska, is sometimes regarded
as a sbsp. of maritima.

List of land and freshwater animal species

29

HAEMATOPUS OSTRALEGUS L.

(Oyster-catcher)

sbsp. malacophaga sbsp. occidentalis f. typ.

Salom. Neum.

Icel., Faeroes. Brit. Isl. N & C Eur.

3 sbspp.
Both coasts of N. America.

Other sbpp. farther south.

2 sbspp.
N Asia.

HIMANTOPUS HIMANTOPUS L.

(Stilt)

/. typ. sbsp. mexicanus P. L. S. Mull.

Mediterr., S Russ. -China. Oregon-Peru.

Several sbspp. farther south.

LIMOSA LAPPONICA L.

/. typ.
(Bar-tailed Godwit)
N Scand. -Taimyr.

sbsp. baueri Naum.
(Pacific Godwit)
NE Asia, Alaska.

LOBIPES (PHALAROPUS) LOBATUS L.

(Northern or Red-necked Phalarope)
Icel.-Hebr., Spitzb., Scand.-Sakhalin. Alas.-Labr., W Greenl.

sbsp. islandicus
Brehm

Icel., Faeroes.

NUMENIUS (PHAEOPUS) PHAEOPUS L.

/. typ. sbsp. variegatus

Scop.
(Whimbrel) '

Scand.-W Sib.

E Sib.

sbsp. hudsonicus
Lath.

(Hudsonian Cur-
lew)

Arct. N. Amer. W
of Hudson Bay.

Kanin Penins.-E Sib.

SQUATAROLA SQUATAROLA L.

(Black-bellied or Grey Plover)
Alas .-Baf fins L.

Charadriiformes, Lari

CATHARACTA (STERCORARIUS) SKUA Brunn., /. typ.

(Northern or Great Skua)

Arct. NE N. Amer. (perhaps not permanent), Greenl. (ditto), Icel.-Orkney.
Several other sbspp. in the southern hemisphere.

3°

CHAPTER I

/. typ.
Eur., W Sib.

CHLIDONIAS NIGRA L.

(Black Tern)

sbsp. surinamensis Gmel.
Alas.-New York.

/• typ.
Eur. (local)-S Mong.,
and farther south.

GELOCHELIDON NILOTICA Gmel.

(Gull-billed Tern)

sbsp. aranea Wils.
Atl. coast of N. Amer.
Virg.-Tex., W. Ind.
Other sbspp. farther south.

sbsp. vanrossemi Bancr.
Calif.

HYDROPROGNE CASPIA Pall, {tschcgrava auctt.)
(Caspian Tern)

Eur.-Ussuri, Afr.

N. Amer. (local): Calif., C Can., Atl. coast.

LARUS ARGENT ATUS Pont.

(Herring Gull)

/■ typ.

sbsp. omissus Pleske

5 sbspp.

sbsp. smithsonianus
Coues

NEur.

N Fennoscand.

Russ.-Sib.

Alas. -Maine.

sbsp. thayeri W. S. Brks.
Arct. isl. of N. Amer.

Several sbspp. farther south.

/• typ.

(Common Gull)
Eur.: Faeroes- Kanin
Penins.

LARUS CANUS L.

sbsp. kamtschatschensis
Bonap.

Siber.-Kamch.

sbsp. brachyrhynchus Rich.

(Short-billed Gull)
NW N. Amer.

LARUS HYPERBOREUS Gunn.

(Glaucous Gull)

Jan Mayen, Spitzb., Kola Penins.- Alas.-N Greenl.

Petchora, arct. Sib.

LARUS MARINUS L.

(Great Black-backed Gull)

Atl. coast of N. Amer. (Nova Scot.- Greenl.-Icel.-NW Eur., E to Petchora.

Labr.).

List of land and freshwater animal species 31

RISSA TRIDACTYLA L.

(Kittiwake)
/. typ. sbsp. polUcaris Ridgw.

Arct. isl. of NE N. Amer., Greenl., NE Sib., Bering Sea, Aleut.

Spitzb., Icel.-NW Eur., Sib. isl.

STERCORARIUS LONGICAUDUS Vieill.

(Long-tailed Jaeger or Skua)
Spitzb., Scand.-Sib. (incl. New Sib. Alas.-Labr.-Greenl.

Isl.).

STERCORARIUS PARASITICUS L.

(Parasitic Jaeger; Arctic Skua)
Icel., Spitzb., Scand.-Commander Isl. Alas.-Baffins L., Greenl.

STERCORARIUS POMARINUS Tcmm.

(Pomarine Jaeger; Pomatorhine Skua)
Kanin Penins., N. Zeml., N Sib. Alas.-Baffins L., W Greenl.

STERNA ALBIFRONS Pall, {minuta L.)

/. typ. sbsp. browni sbsp. athalassos sbsp. antillarum

Mearns Burl. & Low. Less.

(Little Tern) (Brown's Tern) . (Least Tern)

Brit. Isl.-N Russ. Calif.-Mex. Inland of U.S.A. Atl. coast of N.

-Tarbagatai. Amer.: Mass.-

Venez.

Other sbspp. farther south.

STERNA DOUGALLII Mont.,/. typ.

(Roseate Tern)

Atl. coasts of Eur. and N. Amer., N to Denm. and Nova Scot., S to Azor. and Venez.

Also farther south, in part other sbspp.

STERNA HIRUNDO L.

(Common Tern)
/. typ. sbsp. minussensis Sushk. sbsp. longipennis Nordm.

N. Amer. W to Gr. Slave C Asia. NE Asia.

L., S to Venez.; Eur. N
to N Scand., W Sib.

STERNA PARADiSAEA Pont. {tnacYuva Naum.)

(Arctic Tern)

Icel., N Eur. (S to Holl.), N Asia- Alas.-Baffins L., Greenl.

Commander Isl.

32

CHAPTER I

/• typ.

(Sandwich Tern)
W & S Eur.

THALASSEUS (sterna) sandvicensis Lath.

sbsp. acuflavidus Cab.

(Cabot's Tern)

Atl. coast of N. Amer.: Virg.-W. Ind.

Charadriiformes, Alcae

ALCA TORDA L.

(Razor-billed Auk; Razorbill)

/• typ-

Nova Scot.-Labr., Greenl., Icel., NW
Eur.-White Sea.

sbsp. britannica Ticeh.
Faeroes, Brit. Isl., W France.

sbsp. atlantis
Salom.

Maine-Labr.; Icel.,
W Scand.-Brit. Isl.

CEPPHUS (uRIA) GRYLLE L.^

(Black Guillemot)
/• typ.

Baltic Sea.

sbsp. niandti

Mandt

NE Greenl., Spitzb.,

N. Zeml., arct. Sib.

sbsp. arcticiis Brehm
Labr., Baffins L., Greenl.

sbsp. idtimiis

Salom.

Arct. isl. of N.

Amer., N Labr., N

Greenl.

/■ typ.
(Atlantic Puffin)
Ungava-Maine; S
Greenl., Icel., Bear Isl.,
N Norway.

FRATERCULA ARCTICA L.

sbsp. naumanni Nort.
(Large-billed Puffin)
N Greenl., Spitzb., N.
Zeml., Kola Penins.

sbsp. grabae Brehm
(Southern Puffin)
Faeroes, Brit. Isl., W
France, S Scand.

URIA AALGE Pont.

/• typ.

4 sbspp.

(Atlantic Murre;

Northern Guillemot)

Nova Scot.-Labr.; NW Eur.

Greenl., Icel.-

Scotl., Heligol.,

Scand., N. Zeml.

sbsp. inornata
Salom.

NE Asia; Alas.
Wash.

sbsp. calif ornica
Bryant
(Californian Murre)

Calif.

^ The sbspp. of Cepphus grylle seem to be little pronounced and have often been treated
differently.

List of land and freshwater animal species

33

URIA LOMVIA L.

/• typ.
(Briinnich's Murre or Guillemot)
NE Can. (Huds. Str.-Gulf St. Lawr.);
Greenl., Icel., Spitzb., Fr. Jos. L.,
Kola Penins., N. Zeml., Taimyr.

sbsp. arra Pall.

(Pallas's Murre)

E arct. Sib. (S to Japan); Alaska.

/• typ.

(Tengmalm's Owl)

Eur.-Ob (Sib.). C Sib.-Sakhalin.

Strigiformes

AEGOLIUS (CRYPTOGLAUX) FUNEREUS L.

sbsp. sibiricus But. sbspp. jakutorum sbsp. richardsoni

& magnus, But.
NE Sib.

Bonap.

(Richardson's Owl)
Alas.-Nova Scot.

Brit. Isl.-Sakhalin.

ASIO FLAMMEUS Pont.,/. typ.

(Short-eared Owl)

Aleut., Alas.-Ungava, New Jersey,
Greenl.
Several more southern sbspp. in the New World.

ASIO OTUS L.
(Long-eared Owl)

/. typ.
Brit. Isl.-Japan.

sbsp. wilsonianus Less.
Brit. Col.-Nfld.

Scand.-Bering Str.

NYCTEA SCANDIACA L. (tiyctca L.) (fig. 49)

(Snowy Owl)

Alas.-Ungava, Greenl.

/• typ.
Scand.-Kamch.

SURNIA ULULA L.

(Hawk Owl)

sbsp. tianschanica

Smallb.

C Asia.

sbsp. caparoch P. L. S.

Mull.

Alas.-Ungava.

STRIX (SCOTIAPTEX) NEBULOSA J. R. Forst.

(Great Gray Owl)

sbsp. lapponica Thunb.
N Scand.-Sakhalin.

/. typ.
Alas.-N Ontario.

3 ~ 565597 Lindroth

34

CHAPTER I

/• typ.
Brit. Isl., France, Medi-
terr.

TYTO ALBA Scop.

(Barn Owl)
sbsp. guttata Brehm
S Swed., Germ.-W Russ.,
Balcan.

sbsp. pratincola Bonap.
Transamerican in U.S.A.,
S to C Amer.

Numerous more southern sbspp.

/• typ.

Scand.-Amur
& Korea.

sbsp. fasciatus Baird
Alas.-Mont.

Piciformes

PICOIDES TRIDACTYLUS L. (fig. 50)

(Three-toed Woodpecker)
sbsp. crissoleucus sbsp. albidior Stejn.
Reich.
N Sib. Kamch.

sbsp. sakhalinensis

But.

SakhaHn.

3 more southern sbspp. in Asia.

sbsp. dorsalis Baird sbsp. bacatus Bangs

Rocky Mts. Mackenzie-Nfld.

Passeriformes

ACANTHIS (CARDUELIS) FLAMMEA L. {Utiaria L.)i

(Common or Mealy Redpoll)
/, typ. sbsp. holboellii sbsp. fiiscescens sbsp. rostrata Coues

Brehm Coues

Eur., Sib.; Alas. - N Sib. NE Labr., Baffins Greenl., ? Icel.

Ungava. L. ("islandica Htz.")

ACANTHIS (CARDUELIS) HORNEMANNII Holb.^

(Hornemann's Redpoll)

sbsp. exilipes Coues /. typ.

N Scand.-N Japan. Alas.-Ungava. Greenl., Icel., Jan Mayen,

Spitzb.

sbsp. kleinschmidti sbsp. littoralis

Hart. Brehm

(Scandinavian
Rock-Pipit)

Faeroes. Coasts of Scand

sbsp. japonicus Temm. & Schleg.

Lena R.-Kamch., Kuril.

ANTHUS SPINOLETTA L.

/. typ.

sbsp. blakistoni
Swinh.

(Water-Pipit)

C & S Eur. C Asia.

sbsp. rubescens Tunst.

(American Pipit)

Alas.-Nfld., S to New Mex., Greenl.

^ The specific distinction between flammea and hortiemanni, as well as the validity of
their "races", is still disputed.

List of land and freshzoater animal species

35

/• typ.
Scand.-E Sib.

BOMBYCILLA GARRULUS L.

(Waxwing)
sbsp. pallidiceps Reich. sbsp. cedrorum Vieill.

(doubtfully distinct)
Alas.-Manit. Brit. Col.-Nova Scot.

CALCARIUS LAPPONICUS L.

/. typ. sbsp. coloratus

Ridgw.
(Lapland Longspur)
N Scand., N Asia; NE Asia.
Mackenzie-N Queb.

sbsp. alascensis sbsp. siibcalcaratus

Ridgw. Brehm

(Alaska Longspur)

Alas. -Mackenzie. Greenl.

sbsp. britannica
Ridgw.
Brit. Isl.

CERTHIA FAMILIARIS L.

(Creeper or Tree-creeper)
/. typ. several sbspp.

N Eur.

C Asia.

7 sbspp.
N. & C. Amer.: Alas.-New York.

sbsp. orientalis

Dom.

E Sib., Sakh., N

Japan, Kuril.

sbsp. islandicus

Hntz.

Icel.

CORVUS CORAX L.

(Raven)
/. typ. several sbspp.

Eur.-W Sib. N Asia.

sbsp. behringianus

Dyb.

Bering Str.

sbsp. principalis Rigdw. sbsp. sinuatus Wagl.

Alas.-Ellesm. L., Greenl.; S to Georgia. Brit. Col.-C. Amer; E to Mississippi.

EREMOPHILA (OTOCORIS) ALPESTRIS L.

sbsp. flava Gmel. several sbspp. /. typ.

(Shore-Lark) (Horned Lark)

N Eur., N Asia E to Alas.-Huds. Bay, S to Huds. Bay-Nfld.,

Transbaik. Colombia. Greenl. (?)

Many sbspp. in southern mts. of Palaearctic region.

HIRUNDO (cHELIDON) RUSTICA L.

/. typ. sbsp. gutturalis sbsp. tytleri Jerd.

Scop.
(European Swallow)

Eur.-Yenisei, Transbaik.-Sakhal.; NE Sib., Kamch.

N Afr. Japan.

sbsp. erythrogaster

Bodd.

(Barn Swallow)

Alas.-Queb., S to

Mex.

36

CHAPTER I

/• typ.
(Great Grey Shrike)
Eur.-R. Ob.

Eur.-E Sib.

LANIUS EXCUBITOR L.

sbsp. mollis Ev. sbsp. invictus

Grinn.

(Northwestern

Shrike)
Yenisei-Kamch. Alas.-Sask.
Numerous southern Palaearctic sbspp.

LOXIA CURVIROSTRA L.^

(Crossbill)
shsp. japonica sbsp. pusilla Glog.

Ridgw.

Ussuri, Kuril., N Alas.-Queb.
Japan.
Many southern sbspp. in both continents.

sbsp. borealis

Vieill.

(Northern Shrike)

Huds. Bay-Queb.

sbsp. percna Bent
Nova Scot., Nfld.

sbsp. bifasciata Brehm
(Two-barred Crossbill)
N Eur.-Kamch.

LOXIA LEUCOPTERA Gmel.

/• typ.
(White-winged Crossbill)
Alas. -Nova Scot.

/. typ.
Scand.-Yenisei.

/• typ.

(Blue-headed

Wagtail)

Eur., N to C

Scand.

sbsp. alascensis Ridgw.

(Yellow Wagtail)

NE Sib.; W Alas.

LUSCINIA (cYANOSYLVIA) SUECICA L.

(Red-spotted Bluethroat)

sbsp. robusta But.
E Sib.; W Alas.
Several southern Palaearctic sbspp.

MOTACILLA FLAVA L.

sbsp. thunbergi Several sbspp.

Billb.

(Grey-headed W.)

sbsp. simillima
Hart.

Scand.-R. Ob. N Asia. Kamch.

Several southern Palaearctic sbspp.

sbsp. leucorrhoa Gmel.
NE N. Amer. S to N
Queb., W Greenl.

OENANTHE (sAXICOLA) OENANTHE L.

(Wheatear)
sbsp. schioleri Salom. /. typ.

E Greenl., Icel., Faeroes. Eur.-Bering Str.; Alas.

Several southern Palaearctic sbspp.

1 Dr. W. L. Brown, Harvard, informs me that the arrangement of N. American sbspp.
of Loxia curvirostra is in need of revision.

List of land and freshwater animal species

PARUS (PENTHESTES) ATRICAPILLUS L.

sbsp. coUetti Stejn. sbsp. borealis Sel. Several sbspp.

37

sbsp. kamtschat-
kensis Bonap.

(Northern Willow-
Tit)

W Norway. Scand.-N Russia. N & C Asia.

sbsp. turneri sbsp. septentrionalis sbsp. occidentalis

Ridgw. Harr. Baird

(Yukon Chickadee) (Long-tailed Ch.) (Oregon Ch.)

N Alas. Alas.-Manit., S to Brit. Col.-Calif.

N. Mex.

sbsp. bartletti Aldr. & Nutt. Many sbspp. farther south.

Nfld.

Kamch.

/• typ-

(Black-capped Ch.)
Ont.-N. Engl.

/• typ.
N Scand.-W Sib.

PARUS (PENTHESTES) CINCTUS Bodd.

sbsp. obtectus Cab. sbsp. alaskensis Prazak^

Yenisei-Kamch.

(Alaska Chickadee)
NE Sib.(?); Alas., Mac-
kenzie.

/• typ.

(Eversmann's War-
bler)
N Scand.-E Sib.

/• typ.

W Eur .-Ural.

PHYLLOSCOPUS (aCANTHOPNEUSTE) BOREALIS BlaS.

sbsp. kennicotti

Baird

(Willow Warbler)

sbsp. xanthodryas sbsp. examinandiis
Swinh. Stres.

N Japan, Kuril. Kamch., Kuril.

PICA PICA L.
(Magpie)
sbsp. hemileucoptera sbsp. leucoptera
Stegm. Gould

W Sib.-Irkutsk. E Sib.

W Alas.

sbsp. kamtschatica

Stejn.

Kamch.

sbsp. hiidsonia Sab.

Alas.-Manit. (casual farther E), S to

New Mex.

sbsp. nuttalli Aud.
Calif.

PINICOLA ENUCLEATOR L.

(Pine-Grosbeak)
/. typ. sbsp. pacata Bangs, sbsp. kamtschat-

(incl. sbsp. stschur Port.) kensis Dyb.

Scand.-W Sib. E Sib. Kamch., Sakhalin.

6-7 sbspp.

Transamerican :
Alas.-Nfld., S to
New Mex.

^ P. hudsonicus Forst., with its races, is sometimes referred to cinctus. /i \j^' ^n^

'^^oo*,*; c

V5.V "^-il^^

38

CHAPTER I

PLECTROPHENAX (PASSERINA) NIVALIS L.

(Snow Bunting)
sbsp. ihsulae f. typ. sbsp. townsendi

Salom. Ridgw.

Icel., Faeroes. N Eur., N Asia; Both sides of

Alas.-N Queb. Bering Str.

sbsp. subnivalis

Brehm

Greenl.

/• typ.

Eur.-Sib. (E limit?);
Alas.-Queb., S to Tex.

RIPARIA RIPARIA L.^

(Bank Swallow; Sand-Martin)

sbsp. ijimae Lonnb. sbsp. diluta Sharpe &

Wyatt
Kamch.-Japan; Alas.(?). NE Sib.-India.

/• typ.

Alas.-Nfld., S to
N. Mex.

sbsp. corea Og.-

Grant

Korea.

SITTA CANADENSIS L.

(Red-breasted Nuthatch)
3 sbspp.

China.

sbsp. tvhiteheadi Sharpe
Corsica.

sbsp. kruperi

Pelz.

Cauc.-As. Minor.

sbsp. islandicus

Hart.

Icel.

Several sbspp.
Aleut. Isl., &c.

TROGLODYTES (NANNUS) TROGLODYTES L.

(Wren)
sbsp. borealis Fisch. /. typ. Several sbspp.

(and related forms)
Faeroes. Eur.-Ural. N Asia, E to

Kamch., Japan,
Kuril,
sbsp. pacificus Baird sbsp. hienialis Vieill.

Alas. -Alberta, S to Color. Alberta-Nfld.

Reptiles and Amphibians

No species in common.

Pisces — Fishes

Main literature: Jordan, Evermann & Clark (1930), Berg (1933, 1948-49),
Wynne-Edwards (1952).

^ The differentiation of sbspp. in R. riparia is uncertain.

List of land and freshwater animal species 39

? COTTUS GOBIO L.^

(Bullhead)

/. typ. sbsp. sibiricus sbsp. cognatus Rich. sbsp. ricei Nels.

Kessl.
Eur., E to Petchora. Sib.: Ob-Lena. Alas.-Marit. Prov. SE Alas.-Ontario.

of Canada.

Esox LUCIUS L. (estor LeS.)
(Pike)

Irel. & Scand.-Anadyr (extreme NE Alas.-Labr., S to Ohio & New York.

Sib.), S to Italy, Asia Minor, W Tur-

cestan.

LOTA LOTA L.

(Burbot)

/. typ. sbsp. leptura Hbs. & sbsp. maculosa LeS.

Schtz.

Palaearct., E to Lena R. NE Sib. (Kolyma & Ana- Rest of N. Amer.

dyr); NW N. Amer. (Alas.
& Mackenzie R.).

Insecta — Insects

The insect fauna, when properly worked out in a remote future, will no doubt
provide the best foundation for a comparative analysis of the Europe — North-
America relations. At the present time only a small fraction of this, the largest class
in the animal kingdom, has been the subject of modern revisions and is thus suit-
able for our purpose. It was difficult to find, among other things, reliable data on
the distribution of most species in northern Asia, a matter of great significance for
classifying the geographical type and the history of the species.

The list of insects here given could possibly have been extended to include other
groups but the selection has been made in such a way that the main biological
types, above all the types of dispersal, should be represented, which was considered
more important than taxonomic completeness.

^ Berg (1933, p. 169; 1949, p. 1 148) regards sibiricus as distinct from gobio. Wynne-
Edwards (1952, p. 22) says that cognatus is "scarcely distinguishable from C. gobio" but regards
ricei as the North American representative of sibiricus. The taxonomic difficulties in this
group are also pointed out by Jordan, Evermann & Clark (1930).

40 CHAPTER I

Coleoptera — Beetles

The standard sources have been Junk & SchenkHng, Catalogus (1928-33), and
Leng, Catalogue (1920), with five supplements (1927-48). Special references are
given below. It is very much to be regretted that no catalogue of the Coleoptera
of Siberia has appeared after Hey den (1880-98).

Carabidae — Ground-Beetles

A review of the species common to Europe and North America has already been
published (Lindroth, 1954). Only three species are here added: Bradycellus harpa-
linus (Hatch, 1955), Har pal obr achy s leiroides (previously overlooked), and Somo-
trichus elevatus F. (Hatch, 1953, p. 153).

ACUPALPUS MERIDIANUS L.

Brit. IsL, Eur. -Iran, Cauc, S to f Wash., Oreg. (Hatch, 1953).

Mediterr.

AGONUM BOGEMANNI Gyll.

N Eur., mts. of C Eur., Corsica, Sib. Alas.-Labr., Nfld., S to Color.

AGONUM CONSIMILE Gyll.

Scand.-Ural; Kamch. Alas.-Nova Scot., Labr., Nfld.

AGONUM EXARATUM Mnh. {aldatitcum Popp.)
Kola and Kanin Penins., E Sib. Alas., Mackenzie Delta.

AGONUM MANNERHEIMI Dej.

/. typ. (Transitional) sbsp. stygicum Lee.

Scand., N Russ., W Sib. E Sib. Alas.-Labr., Nfld., S to

New York.

AGONUM MULLERI Hbst.

Brit. Isl., Eur., Cauc, W Sib. f New Engl., E Can., Nfld.; Oreg., Wash.,

Brit. Col.

AGONUM QUADRIPUNCTATUM DeG.

Brit. Is!., Eur.-Kamch. Alas.-Labr., Nfld.

India, Philipp.

List of land and freshwater animal species 41

AGONUM RUFICORNE Gze.

Brit. Isl., Eur., Mediterr., Macaron. f Maine, N. Brunsw., Nova Scot., Nfld.

AGONUM THOREYI Dej.

Brit. Isl., Eur.-E Sib.: Amur. Alas.-Nova Scot., Nfld., S to Calif.

AMARA AENEA DeG.

Brit. Isl., Eur., N Afr., Hither & C f NE U.S.A., E Can., Nfld.

Asia, W Sib.

AMARA ALPINA Payk. (fig. 40)

/. typ. (Transitional) sbsp. brunnipennis Dej.

Scotl., Scand.-W Sib. E Sib. Alas.-Labr., Nfld., Baffins

L., S to Color., mts. of New
Engl.

AMARA ANTHOBIA Villa

Brit. Isl., W, C & S Eur., Asia Minor, f Wash., Oreg. (Hatch, 1953).
Cauc.

AMARA APRICARIA Payk.

Brit. Isl., (i loc), Eur.- Sib.: Amur. (f) NE U.S.A., Can.: Nfld., Labr.-E

Brit. Col.

AMARA AULICA Panz.

Brit. Isl., Eur., Cauc, W Sib., W. f Nova Scot. (Cape Bret. Isl.), Nfld.

Turcestan.

AMAR.\ BIFRONS Gyll.

Brit. Isl., Eur., Cauc, W. Turcestan. f Nova Scot., Nfld.

AMARA ERRATICA Dft.

Eur. (boreo-alpine), Cauc, Sib. E to Alas.-Labr., Nfld., S to Color.

Kamch.

AMARA FAMILIARIS Dft.

Brit. Isl., Eur., Cauc, W Sib., N Afr. f NE U.S.A., E Can.-Nfld.; Oreg.-

Brit. Col.

AMARA FULVA DeG.

Brit. Isl., Eur., Asia Minor, Cauc, W f E Can., Nfld.
Sib.

42 CHAPTER I

AMARA HYPERBOREA Dej.

N Finl.-E Sib. Alas.-Labr., Nfld., S to Lake Sup.

AMARA INTERSTITIALIS Dej.

Scand.-Kamch., Cauc. Alas., Yukon Terr.

AMARA LUNICOLLIS Schio.

Brit. Isl., Eur.-Kamch., S to Iran. (f) NE U.S.A., E Can.-Nfld. (probably

introduced); C & N Can. (probably native)

AMARA QUENSELi Schnh. {remotestriata Dej.)

Brit. Isl., Eur. (boreo-alpine), Cauc, Alas.-Labr., Nfld., S to N. Mex.

Sib. E to Kamch.

AMARA TORRiDA 111. {cylittdrica Lee, &c.)
N Scand.-Kamch. Alas.-Labr., Nfld., S to Color.

ANISODACTYLUS BINOTATUS F.

Brit. Isl., Eur.-W Sib., N Afr. f Oreg., Wash., Brit. Col. (Hatch, 1953).

ASAPHIDION FLAVIPES L.

Brit. Isl., Eur., Cauc, W Sib., N Afr. f New York.

BEMBiDiON BRUXELLENSE Wesm. (rupcstre auctt.)
Brit. Isl., Eur.-Sib.: Lena R. f E Can., Nfld.

BEMBIDION CRENULATUM F. Sahib.

sbsp. /)ono;'e«5e J. Sahib. f- typ. shsp. farrarae Hatch

Kola Peninsula. E Sib. Oreg., Wash.

BEMBIDION DAURICUM Mtsch.

N Scand., Sib. E to Ochotsk. Alas., N. W. Terr., N Manit.

BEMBIDION GRAPEI Gyll.

Icel., N Eur.-Kamch. (figs. 45-47) Alas.-Labr., Nfld., S to Sierra Nevada;

Greenl.

BEMBIDION HASTi C. R. Sahib.

Scand.-N. Zeml., Sib. E at least to Brit. Col. & N. W. Terr.-N Labr.

Baikal.

List of land and freshwater animal species 43

BEMBIDION HYPERBORAEORUM Munst.

N Scand., Kanin Penins., Sib. (Yenisei Alas., N. W. Terr.
& Lena).

BEMBIDION LAMPROS Hbst.

Brit. Isl., Eur., Cauc, Sib.: E to R. f Brit. Col.; Nfld.

Lena.

BEMBIDION LAPPONICUM Zett. {bryantl Carr)

/. typ. sbsp. latiusculum Mtsch. sbsp.(?)

N Scand., N Russ., Sib. E Sib., Kamch. Alas., N. W. Terr.

E at least to Yenisei.

BEMBIDION MCKiNLEYi Fall (scandicum Lth.)
N Scand. Alas.: Mt. McKinley.

BEMBIDION OBSCURELLUM MtSCh. (fllSClCrUS MtSch.)

Denm.: Jutland; NE Eur., N & C Asia, NW N. Amer., S to N. Mex.
E to Kamch.

BEMBIDION PETROSUM Gebl.

sbsp. siebkei J. Miill. /. typ. sbsp.(?)

Scand., W. Sib. C Sib. Alas.-Labr., Nfld., S to

N. Mex.

BEMBIDION PROPERANS Steph.

Brit. Isl., Eur., Cauc, Sib. E to Amur, f Nova Scot.

BEMBIDION QUADRIMACULATUM L.

/. typ. sbsp. oppositum Say

Brit. Isl., Eur., Cauc, Sib. E to Brit. Col.-Nova Scot., Nfld., S to Tex.

Ussuri. & N. Mex.

BEMBIDION STEPHENSI Crotch

Brit. Isl., Eur., E to Poland & Slovakia, f Mass., E Can., Nfld.

BEMBIDION TETRACOLUM Say (ustulatum auctt.)

Brit. Isl., Eur., Cauc, Hither Asia, W f Oreg.-Brit. Col.; E U.S.A.. E Can.,
Sib., N Afr. Nfld. (fig. 12)

BEMBIDION TRANSPARENS Gebl.

N & E Eur., Sib. E at least to R. Lena. Alas.-Labr., Nfld., S to Indiana, (fig. 58)

44 CHAPTER I

BEMBiDiON YUKONUM Fall {grapeioides Munst.)
N Scand., Sib.: Jakutsk. Alas., Yukon Terr., N. W. Terr.

BLETHISA ESCHSCHOLTZI Fisch. (fig. 56)

SE Russ., SW Sib. Texas.

BLETHISA MULTIPUNCTATA L. (fig. 56)

/. typ. sbsp. aurata Fisch.

Brit. Isl., Eur.-W & C Sib. E Sib.; Brit. Col. & Yukon Terr.-Labr.,

Nfld.

BRADYCELLUS HARPALINUS Serv.

Brit. Isl., Eur. E to C Russ., Cauc, N f Vancouver, B. C. (Hatch, 1955).
Afr., Madeira.

CALATHUS FUSCIPES Gze.

Brit. Isl., Eur., W Asia E to Iran, f Vancouver, B. C.

Cauc, N Afr.

CALATHUS MICROPTERUS Dft.

/. typ. sbsp. ingratus Dej.

Brit. Isl., Eur., Cauc, W, C & (?) E Sib. Alas.-Labr., Nfld., S to Color.

CARABUS AURATUS L.

Brit. Isl. (not established), Europe f New^ Engl.: Mass., Vermont, Maine,

(excl. the N & E).

CARABUS CANCELLATUS 111.

Brit. Isl. (not established), Eur., Sib. f Wise, N. Carol, (doubtfully established).

E to Lena R. (Numerous sbspp.)

CARABUS GRANULATUS L. (fig. 8)

sbsp. hibernicus Lth. /. typ.

Brit. Isl. (espec Ireland). Brit. Isl., N, W & C Eur. E to Ural.

t Nova Scot., N. Brunsw. f New Engl., Wash., Brit. Col., occas. in

E Can. and N. W. Terr.
Other sbspp. in S Eur. & Asia, E to Amur, Sakhalin, & Japan.

CARABUS NEMORALIS Mull. (fig. 7)

Brit. Isl., Eur., E to C Russ. f Oreg.-Brit. Col.; NE U.S.A., E to Wise,

E Can., Nfld.

List of land and freshwater animal species 45

CARABUS TRUNCATICOLLIS Eschz.

sbsp. polaris Popp. /. typ.

NE Russ. (Petchora), N. Zeml., N Sib. Alas. W to Bering Str., E to Mackenzie

E to Anadyr. Delta.

CLIVINA COLLARIS Hbst.

Brit. Isl., Eur., Cauc, W Asia E to W f Wash.; Mass., E Can.
Turcestan.

CLIVINA FOSSOR L.

Brit. Isl., Eur.-Kamch. t Wash.; E Can., Nfld. (fig. 13)

DIACHILA ARCTICA Gyll.

/. typ. sbsp. amoena Paid.

N Eur.,? NW Asia. S & E Sib.; Alas., NW Can., Labr.

DIACHILA POLITA Paid.

NE Eur.-Kamch., S to Baikal region. Alas., N. W. Terr. (Mackenzie Delta).

DYSCHIRIUS HELLENI J. Miill.

Scand., Sib.: Yenisei. Alas., N Manit., Labr.

DYSCHIRIUS POLITUS Dej.

Brit. Isl., Eur., Iran, Sib. E to New York, New Hampsh., Nfld. (prob-

R. Lena. ably overlooked).

ELAPHRUS LAPPONICUS Gyll.

Brit. Isl., Scand., Sib. E to Kamch. Alas. -Labr.

ELAPHRUS RIPARIUS L.

Brit. Isl., Eur., N & C Asia E to Alas.-Labr., Nfld., S to Calif.

Kamch.

HARPALOBRACHYS LEIROIDES MtSch.

NE Russ., Sib. E to Ussuri. N. W. Terr. (Port Smith).

HARPALUS AFFiNis Schrk. (aenetis F.)

Brit. Isl., Eur., N Asia, E at least to R. f Wash., Brit. Col.; E U.S.A. (S. to Plor.,
Lena and Transbaik. W to Kansas), E Can.-Labr., Nfld.

HARPALUS FULIGINOSUS Dft.

Eur. (not Brit. Isl.), N Asia-Kamch., Alas.-Labr., Nfld., S to Mich.

Japan.

46

CHAPTER I

HARPALUS NIGRITARSIS C. R. Sahib.

/• typ-

N Fennoscand., Sib. (?).

sbsp. proximus Lee.

Alas.-Labr., Nfld., S to N. Hampsh.

HARPALUS RUFIPES DcG. {pubescens O. F. Miill.)

Brit. Isl., Eur., N Afr., N Asia E at f E Can., Nfld.

least to R. Lena, Japan.

LiciNUS PUNCTATULUS F. (sUphoides F.y

S Engl., W & S Eur. N to W Germ., f Mass., Maine (temporarily established).
N. Afr., Hither Asia.

MISCODERA ARCTICA Payk.

/. typ. sbsp. americana Mnh.

(little pronounced)
Brit. Isl., N & C Eur., Sib. E at least Alas.-Labr., Nfld., S to New Engl,
to R. Lena.

/• typ.

NEBRIA GYLLENHALi Schnh. (rufescens Strom)

sbsp. besseri Fisch. sbsp. castanipes Kby.

Greenl., Icel., Brit. Isl.,
Eur. (boreo-alpine), N
Asia (E limit uncertain).

(transitional)
E Sib.

NEBRIA NIVALIS Payk.
sbsp. femorata Mtsch.

/. typ.

(transitional)
Scotl., Scand., W Sib., E E Sib.
at least to R. Ob.

Alas.-Labr., Nfld., S to New
Engl.

sbsp. bifaria Mnh.

Kamch., Alas .-Baf fins L.,
Queb. (Gasp6), Maine
(Mt. Katahdin), Nfld.

NOMIUS PYGMAEUS Dej.

S Eur. N to Paris, N Afr., W Asia. Brit. Col.-Lake Sup., S to Georgia &

Calif.

NOTIOPHILUS AQUATICUS L.

Icel., Brit. Isl., Eur., N Asia E to Alas.-Labr., Nfld., S to New Mex.

Kamch.

^ L. sUphoides Rossi 1790 is a different species {vide Jeannel, 1942, p. 994-996) but the
American specimens belong to sUphoides Fabricius 1792, the right name of which is pimcta-
tulus Fabricius 1792.

List of land and freshwater animal species 47

NOTIOPHILUS BIGUTTATUS F.

Icel., Brit. Isl., Eur. E to Ural., Cauc. f Nfld. (fig. 14)

PATROBUS SEPTENTRIONIS Dej.

/. typ. sbsp. labradorinus Csy. (validity un-

certain)
Greenl., Icel., Brit. Isl., Eur. (boreo- Alas.-Labr. (in part transitional), Nfld.,

alpine). Sib. E to Amur & Kamch. S to Color.

In Eurasia two more southern sbspp.

PELOPHILA BOREALIS Payk.

/. typ. sbsp. eschscholtzi Mnh. sbsp. ulkei Horn

(transitional)
Brit. Isl., N Eur., Sib. E Kamch., Alas. NW Can.-Labr., Nfld.

at least to R Lena.

PERIGONA NIGRICEPS Dej.

Cosmopolitan,
t Eur. (rapidly dispersing). f U.S.A. (widely distributed).

PLOCHIONUS PALLENS F.

Cosmopolitan
t W & S Eur. t U.S.A. (both coasts).

PRISTONYCHUS (laemosthenes) complanatus Dej.

Almost cosmopolitan,
t Brit. Isl., W & S Eur. Native of N. Africa. fPacific coast (Calif.-Brit.

Col.)

PRISTONYCHUS TERRICOLA Hbst.

Icel., Brit. Isl., Eur., Cauc. t E Can., Nfld.

PTEROSTICHUS ADSTRICTUS Eschz.

Icel., Brit. Isl., N Eur.-Kamch. Alas.-Labr., Nfld., S to Ariz.

PTEROSTICHUS (CRYOBIUS) BREVICORNIS Kby.

{fastidiosus Mnh., mandibidaris auctt.)
Kola Penins., N Russ.-Kamch. Alas.-Labr., Nfld.; mts. of New Engl.i

PTEROSTICHUS MELANARius 111. {vulgaris auctt.)
Brit. Isl., Eur., Cauc, Sib. E to Amur, f Oreg.-Brit. Col.; E Can., Nfld.

^ The form from E of Hudson Bay is probably subspecifically distinct.

48 CHAPTER I

PTEROSTICHIUS STRENUUS Panz.

Brit. Isl., Eur., Cauc, Sib. E to Amur, f Nfld.

PTEROSTiCHUS VERMICULOSUS Men. (tnnuitorum Bwn.)

NE Russ. (Petchora), Sib. E at least N. W. Terr,

to R. Lena.

SOMOTRICHUS ELEVATUS F.

Cosmopolitan,
t W Eur. t Wash.

STOMis PUMicATus Panz.

Brit. Isl., Eur. E to Ural, Cauc, Asia f Queb.
Minor.

TACHYS PARVULUS Dej.

S Engl., C & S Eur. f Wash.

TACHYTA NANA Gyll.

/. typ. sbsp. inornata Say

Eur. {not Brit. Isl.), N Afr., Hither Wash.-New York, S to Flor.

Asia, Sib. E to Amur.

TRECHUS DISCUS F.

Brit. Isl., Eur., Asia E to Japan. f E Can.

TRECHUS OBTUSUS Er.

Icel., Brit. Isl., W, C & S Eur., N Afr. f Oreg., Wash.

TRECHUS RUBENS F.

Icel., Brit. Isl., N, W & C Eur., Sib. f E Can., Nfld.
E to Lena R.

TRICHOCELLUS COGNATUS Gyll.

Icel., Brit. Isl., N, W & C Eur., Sib. Alas.-Labr., Nfld., Greenl.

E at least to Lena R.

Elateridae — Click-Beetles

Dr. E. C. Becker, Department of Agriculture, Ottawa, kindly gave information
on this family as a supplement to the records in Leng's Catalogue. For Asia,
besides Heyden (1880-98), Jacobson (1905-15) has been used.

List of land and freshwater animal species 49

AGRIOTES LINEATUS L.

Brit. Isl., Eur., N Afr., W & NW f Brit. Col.; Nova Scot., Nfld.

Asia.

t N. Zeal. t C & S Amer.

AGRIOTES OBSCURUS L.

Brit. Isl., Eur., N Asia E to Baikal & f Brit. Col.; Nova Scot., Nfld.
Amur.

AGRIOTES SPUTATOR L.

Brit. Isl., Eur., N Afr., W, NW & C f N. Brunsw., Nova Scot.

Asia.

CORYMBITES (cTENICERA) CRUCIATUS L.

/. typ. sbsp. festivus Lee. sbsp. pulcher Leo.

Eur. {not Brit. Isl.), N Brit. Col.-Manit. Ontar.-Nova Scot.

Asia.

CORYMBITES (ctenicera) sjaelandicus O. F. Miill.

Brit. Isl., N & C Eur., N Asia S to (f) NE U.S.A., W to Mich., N Ontar.-

Amur, Korea. Gaspe.

EANUS (corymb iTEs) cosTALis Payk.

/. typ. sbsp. decoratus Mnh.^

N Eur., N Asia E to Amur. N.W.Terr.-Labr.,N Nfld., New Hampsh.

ELATER (aMPEDUS) NIGRINUS Hbst.

Brit. Isl., Eur., N Asia E to Amur. Alas.-Labr., S to NE U.S.A.

harminius (athous) undulatus DeG.

/. typ. sbsp.?

Brit. Isl., N & C Eur., N Asia E to Queb., Ont. (certainly wider distr.)

Amur & Kamch.

HYPNOiDus (hypolithus) hyperboreus Gyll.

Eur. (boreo-alpine), N Asia E to Alas., NW Can.

Kamch.

HYPNOIDUS (hypolithus) rivularius Gyll.

Eur. (boreo-alpine), Altai, Sib. E at Alas.-Labr., Nfld.; N. Hampsh.

least to Lena R.

1 Fall (1934, p. 35) was unable to make a definite separation between decoratus and
the European costalis; Brown (1936, p. 247) regards them as different species.

4 ~ 565597 Lindroih

50 CHAPTER I

MELANOTUS CASTANIPES Payk.

Brit. Isl., Eur., Cauc, W & N Asia (f) NE U.S.A. W to Mich. & Indiana;

E to Amur. E Canada, Nfld.

SERicus BRUNNEUS L. (incoTtgruus Lee.)
Brit. Isl., Eur., Sib. E to Amur. B. C.-E Can., S Labr., Nfld.i

Coccinellidae — Ladybirds

For the Palaearctic fauna, Dobrzhanskij (1926) and Mader (1926-37) have been
consulted. Valuable supplementary informations have been received from Dr. E.
A. Chapin, West Medway, Mass.

ADALIA BIPUNCTATA L.

Brit. Isl., Eur., N Afr., Asia E to f Wash., Idaho (Hatch, 1953); NE

Baikal and N China. U.S.A., E Can., Nfld.

ADALIA FRIGIDA Schn.

N Eur., Sib. E to Amur & Kameh. Brit. Col., Wash.-Maine, Nfld., S to

Ariz.

ANATIS OCELLATA L.

Brit. Isl., Eur., N Asia E to Amur, f Wash.; NE U.S.A. W to Lake Sup.,

Kamch. & Japan. Nfld.

ANISOCALVIA 14-GUTTATA L.

/. typ. sbsp. i2-?naculata Gebl.^

Brit. Isl., Eur.-Sakhalin, Kuril.; Brit. E Sib.; N. Amer.: Wash.-Nfld.
Col.-Maine, Nfld.

ANISOSTICTA STRIGATA Thunb.

/. typ. sbsp. bitriangularis Say^

N & C Eur., Sib. E at least to Baikal. Wash, Oreg.-Nova Scot., S Labr.

BULAEA LICHATSCHOVI Humm.

Hung., S Russ., N Afr. Hither Asia, f New Jersey (probably accidental).

S & W Sib., Tien-shan.

* The Sericiis of N. America may be subspecifically distinct.
^ By Chapin {in lift.) regarded as a colour aberration only.

* Often regarded as distinct species.

List of land and freshwater animal species 51

COCCINELLA HIEROGLYPHICA L.

/. typ. sbsp. mannerheimi sbsp. humboldtiensis sbsp. tricuspis Kby.

Muls. Nun.

Brit. Isl., Eur., W E Sib.-Amur, Sak- Wash., N Calif. C Can.-Mich.

Sib. halin.

COCCINELLA TRIFASCIATA L.

Eur. (boreo-alpine), N & C Asia E to Wash., Oreg.-Nova Scot., Nfld.
Pacific coast.

COCCINELLA 1 1 -PUNCTATA L.

/. typ. sbsp. ainu Lew. (?) sbsp. menetriesi Muls.

Icel., Brit. Isl., Eur.-C Japan. Alas.-N Calif.

Asia, N Afr.
t NE U.S.A., E Can., Nfld.

HIPPODAMIA 13-PUNCTATA L.

/. typ. sbsp. tibialis Say.

Brit. Isl., Eur., Asia (inch Sib.) E to Alas.-Maine, S Labr., Nfld., S to Calif.

Amur, Kamch. & China.

NEOMYSIA OBLONGOGUTTATA L.

/. typ. 4 sbspp.

Brit. Isl., Eur., Asia E to Amur; (sbsp. pullata Say, &c.)

(?) Oreg. Oreg.-Labr.

STETHORUS (SCYMNUS) PUNCTILLUM Wsc.

Brit. Isl., Eur., Asia E to Japan. f Mass., Ontar.

Scarabaeidae — Dung-Beetles

The main source was Schmidt (1922). Valuable information was also obtained from
Mr. B.-O. Landin of the Zool. Inst., Lund.

AEGIALIA ARENARIA F.

Brit. Isl, Baltic coasts, W & C Eur. f Massach. (Darlington, 1927).

AEGIALIA RUFA F.

Brit. Isl., Baltic coast, W & C Eur. \ Miquelon (S of Nfld.).

52 CHAPTER I

APHODIUS BOREALIS Gyll.

Brit. Isl., N & C Eur., Sib. {i.a. Nfld. (probably indigenous in N. Amer.

Baikal)'. and overlooked).

APHODIUS DEPRESSUS Kug.

Brit. Isl., N & C Eur., N & C Asia E f New York,
to Amur.

APHODIUS DisTiNCTUS O. F. Miill. {inquinatus Hbst.)
Brit. Isl., Eur., N Asia (at least W Sib.), f Wash.; Atl. States of U.S.A.

APHODIUS ERRATICUS L.

Brit. Isl., Eur., N Afr., N & C Asia E f NE U.S.A., E Can., Nov. Scot,
at least to Baikal.

APHODIUS FASCiATUS Ol. (putridus Hhst., foetidus F. nee Hbst., tenellus Say)

Brit. Isl., Eur. (widely), Cauc, Sib. f Wash.; E U.S.A. S to New Mex. &

E to Kamch. Color., N to S Labr.

APHODIUS FIMETARIUS L.

Brit. Isl., Eur., N Afr., N & C Asia to f Brit. Col., N.W. Terr., Wash.; Tex.-
E Sib. Nov. Scot., Nfld.

APHODIUS FOSSOR L.

Brit. Isl., Eur., N Afr., Asia to E Sib. f Brit. Col.; N. Carol.-Nov. Scot. & S

Labr., W to Mich. & Ont., Nfld.

APHODIUS GRANARIUS L.

Almost cosmopolitan.

Brit. Isl., Eur. (generally), N & C Asia f N. Amer. (widely distr.), N to Wash,
to E Sib. & Nov. Scot.

APHODIUS HAEMORRHOIDALIS L.

Brit. Isl., Eur., Sib., N & C Asia. f NE U.S.A., E Can.-Nov. Scot., Nfld.

APHODIUS LIVIDUS Ol.

Almost cosmopolitan.

Brit. Isl., Eur. (except the North), N & f S U.S.A., N to Indiana & Calif.
C Asia. W. Ind.

APHODIUS PRODROMUS Brahm
Brit. Isl., Eur., N & C Asia. f E U.S.A., S to Kent., E Can.

List of land and freshwater animal species 53

APHODIUS RECTUS MtSch.

(f ) Greece. Native of E Asia. f Wash.

APHODIUS RUFIPES L.

Almost cosmopolitan.

Brit. Isl., Eur. (generally), Asia E to f E U.S.A.

Amur, China.

APHODIUS SCROFA F.

Brit. Isl., Eur., W Asia. t Maine, Queb., N. Brunsw.

APHODIUS SUBTERRANEUS L.

Brit. Isl., Eur., W & C Asia E at least f NE U.S.A., E Can.-Nov. Scot,
to SW Mongol.

GEOTRUPES STERCORARIUS L.

Brit. Isl., Eur., N Asia to E Sib. & f N. Brunsw., Pr. Edw. Isl.

Japan.

GEOTRUPES STERCOROSUS Scriba {sylvaticus Panz.)
Brit. Isl., Eur., W Asia. t Rhode Isl. (i ex.).

HYBOSORUS ILLIGERI Rche.

S Eur., Afr., W Asia. t S U.S.A.

ONTHOPHAGUS NUCHICORNIS L.

Brit. Isl., Eur., W Asia. t Brit. Col.; NE U.S.A., E Can.-Nov.

Scot., Nfld.

OXYOMUS SILVESTRIS Scop.

Brit. Isl., Eur., W Asia. t Wash.; NE U.S.A., Nov. Scot. (Hatch,

1946).

PLEUROPHORUS CAESUS CrtZ.

Almost cosmopolitan.

Brit. Isl., Eur., N Afr., Asia Minor, f Wash., Oreg.; NE U.S.A., W to

W. Turcest. Indiana.

RHYSSEMUS GERMANUS L. {puncticolUs Bwn.)

Brit. Isl., Eur., Afr., N & C Asia E to f Ontario.
N China.

54 CHAPTER I

TROX SCABER L.

Cosmopolitan.
Brit. Isl., Eur. (generally). f Flor., Indiana.

Cerambycidae — Longhorn Beetles
Records of the occurrence in Asia were taken from Plavilstshikov (1936, 1940).

ACMAEOPS PRATENSIS Laich

N & C Eur. {not Brit. Isl.), Sib. E to Alas.-Labr., Nfld., S to New Mex. &
Amur., Kamch., Korea, & Sakhal. NE U.S.A.

CALLIDIUM VIOLACEUM L.

Brit. Isl., Eur., Sib. E to Amur, Sakhal. f NE U.S.A., E Can., Nfld.
& Japan.

GRACILIA MINUTA F.

Brit. Isl., Eur., N Afr., Hither Asia. f Wash.; E U.S.A.

t Japan.

HYLOTRUPES BAJULUS L.

Brit. Isl., Eur., N Afr., Hither Asia. f E U.S.A.

"Siberia" (probably W).

t China.

JUDOLIA SEXMACULATA L.

Brit. Isl., N & C Eur., Sib. E to Amur, Wash., Huds. Bay Terr.-S Labr. & New
Sakhal. & Japan. Engl., S to Calif.

LEPTIDEA BREVIPENNIS Muls.

Brit. Isl., S & C Eur., E to S Russ., f Calif.

N. Afr., Cauc, Hither Asia.

LEPTURA RUBRA L. (planata Sw. & Hopp.)
Brit. Isl., Eur., N. Afr., Sib. E to Baikal, f Iowa.

NEOCLYTUS ACUMiNATUS F. {erythrocephaliis F.)

t Trieste, NW Jugosl., Austria, S Can.; Wash.-New Engl., S to Flor.

Germ.; Gr. Brit, (accidental). (G. Mul-
ler, 1949-53, P- 148-149)-

List of land and freshwater animal species 55

PACHYTA LAMED L.

N & C Eur., Sib. S to Altai, E to Alas.-Queb. & Penns., S to Calif.

Sakhal.

PHYMATODES LIVIDUS Rossi

Brit. Isl., C & S Eur., Eur. Russ., f NE U.S.A., E Can.

Cauc, N. Afr.

PHYMATODES TESTACEUS L.

Brit. Isl., Eur., N. Afr., Hither Asia. f Calif.; E U.S.A., S to Georgia, W to

Kansas.

RHAGIUM INQUISITOR L.

/. typ. sbsp. stshukini Sem. sbsp. rugipenne Rtt. sbsp. lineatiim Ol.

Brit. Isl., Eur., N. Cauc. E Sib., Manch., Alas.-Queb. &

Afr., Sib. -Amur, Korea, Kamch., New Engl.

Kamch., Japan. Sakhal.

SAPERDA POPULNEA L.

/. typ. sbsp. moesta Lee.

Brit. Isl., Eur.-E Sib. Wash.-Calif. E Can.-Wyom., Penns.

TRAGOSOMA DEPSARIUM L.

N Eur., C Eur. mts., Sib. E to Amur. Brit. Col.-Nfld.

Chrysomelidae — Leaf Beetles

Apart from current literature, valuable information has been obtained from Mr.
W. J. Brown, Ottawa, and Dr. Ernst Palmen, Helsingfors. Records of European
species of Phyllodecta (Phratora) and of Melasoma lapponica L. from N. America
are false, according to Brown (1951 and in litt.).

ADOXUS OBSCURUS L.

/. parthenogen.^ f. bisex.

Eur. (not Brit. Isl.), Sib. E to Kamch., Alas.-Nfld., S to New Mex.

China, Japan.

AGELASTICA ALNI L.

Brit. Isl., Eur., W Sib., Japan. t Conn., New Jers., New York.

^ Single males of Adoxus obscurus have been observed in S Eur. but it seems that the
species is constantly parthenogenetic in the whole of Europe.

56 CHAPTER I

CASSIDA FLAVEOLA Thunb.

Brit. Isl., N & C Eur. t E U.S.A. W to Minnes., Can. W to

Alberta, N to Churchill, Man., & Nor-
man Wells, N. W. Terr.

CASSIDA RUBIGINOSA O. F. Miill.

Brit. Isl., Eur., Asia E to Japan. f E Can., New Brunsw. to Lake Huron;

Maine, New York.

CHRYSOMELA (cHRYSOLINA) STAPHYLEA L.

Icel., Brit. Isl., N & C Eur., Sib. E to f Nov. Scot., Nfld.
Kamch. (sbsp. daurica Gebl.)

CRIOCERIS ASPARAGI L.

Brit. Isl., Eur. (except the north), N. f Wash.-Calif.; E U.S.A.
Afr., N Asia.

CRIOCERIS I2-PUNCTATA L.

Eur. {not Brit. Isl.), Sib. E to Amur. f Wash.; E U.S.A.

GALERUCELLA LUTEOLA O. F. Miill. (xatithomelaena Schrk.)

Eur. {not Brit. Isl.), Hither Asia, C f Wash.-Calif.; NE U.S.A., E Can.

Asia.

GALERUCELLA NYMPHAEAE L.

Brit. Isl., Eur., Sib. E to Amur, Kamch. Alas.-S Labr. & Nfld., S to Texas.
& Japan.

GASTROIDEA POLYGONI L.

Brit. Isl., Eur., N Asia to E Sib. f Canada, Nov. Scot, to Sask.; U.S.A.

W to Mississ.

LEPTINOTARSA DECEMLINEATA Say (fig. 3)

t W & C Eur., N to S Denm., E to Native of Rocky Mts. (Color., New

Polon. & Rouman. Mex.), now generally distributed in U.S.A.

& S Can.

LILIOCERIS LILII Scop.

Brit. Isl., Eur. (except the north). Sib. f Montreal (Brown, 1946, p. 47).
E at least to Baical.

LONGITARSUS succiNEUS Foudr. {ovalis Gent.)
Brit. Isl., Eur., Sib. E to Vladiv. f Nfld.

List of land and freshwater animal species 57

LONGITARSUS WATERHOUSEI Ktsch.

Brit. Isl., W, C & S Eur., Asia Minor. f Oreg.; Mich., Indiana.

MANTURA CHRYSANTHEMI Koch

Brit. Isl., Eur., N. Afr. f Massach,

PHAEDON ARMORACIAE L.^

Brit. Isl., Eur., Sib. E to Kamch. & (f) Brit. Col.-Maine, S to Color.

Japan.

PHAEDON COCHLEARIAE F.^

Brit. Isl., Eur., Asia Minor. (t) NE U.S.A., E Can., W to Utah &

Alberta.

PHYLLOTRETA AEREA AUard

Brit. Isl., C & S Eur., Asia Minor. f NE U.S.A.

PHYLLOTRETA ARMORACIAE Koch

Eur. (except the north & Brit. Isl.), f NE U.S.A., E Can., W to Nebr.

Sib.

PHYLLOTRETA UNDULATA KtSch.

Brit. Isl., Eur., N. Afr., Asia E to f Maryland.

Amur.

PHYLLOTRETA viTTATA F. (striolota F., sinuata Redtb., nee auctt. americ.)

Brit. Isl., Eur., Asia E to Amur, Alas.-Nov. Scot., Nfld., S to Calif. &

Corea, Japan & Sakhal. the Gulf of Mex.

PHYLLOTRETA ziMMERMANNi Crotch {sinuata auctt. americ.)

NE Eur. (Finl.); E Sib. (Ussuri). New Engl .-Georgia, W to Manit.

(probably wider distr.).

PHYTODECTA AFFINIS Gyll. {arctlca Mnh.)2

Eur. (boreoalpine), Sib., Altai. Alas.-Huds. Bay., Color. (Holdhaus &

Lindroth, 1939).

PLAGIODERA VERSICOLORA Laich

Brit. Isl., Eur., N. Afr., Sib. E to f NE U.S.A., E Can.

Amur, China & Japan.

^ Fall's identification of Phaedon is possibly in need of confirmation.
^ Brown (1942, p. 100) questions this synonymy.

58 CHAPTER I

PRASOCURIS PHELLANDRII L.

Brit. Isl., Eur., W Sib. Brit. Col.-Queb., S to Color.

PSYLLIODES CHRYSOCEPHALA L.

Brit. Isl., Eur., Hither Asia, Sib. f Nfld.

PSYLLIODES CUCULLATA 111.

Eur. (not Brit. Isl.), N Asia E to Pacif. f E Canada.: New Brunsw. & Gaspe.
coast.

ZEUGOPHORA SCUTELLARIS SuflFr.^

N & C Eur. (not Brit. Isl.), W Sib. f Wash., Oreg.; Mich., Indiana, New

Mex.

Rhynchophora {Anthribidae & Curculionidae) — ^Weevils

The number of known species common to Europe and North America is larger in
this group of beetles than in any other, but a close and modern comparison of the
species in question has actually been made only for those of European origin in-
troduced into the Maritime Provinces of Canada, including Newfoundland (Brown,
1940, 1950; Palmen, in litt.). Therefore several of the earlier records must be re-
garded with some doubt and, pending further evidence, it seems safe to omit the
following species still listed by Leng (1920): — Anthonomus pomorum L., Brachy-
deres incanus L., Cionus scrophulariae L., Grypus hrunnirostris F., Lepyrus capu-
cinus Schall., Phyllobius glaucus Scop., Phytobius velutus Beck [velatus auctt.),
Sitona lineellus Bonsd. and 5. tibialis Hbst. — Blatchley & Leng (19 16) has been
very useful. Invaluable information has been given by Mr. W. J. Brown and Dr.
E. Palmen,

ACALYPTUS CARPINI Hbst.

Brit. Isl., Eur., Asia E to Amur & f NE & C U.S.A., E Can., Nfld.

Kamch.

AMALUS HAEMORRHOUS Hbst.

Brit. Isl., Eur., Asia E to Amur. (f) Wash.; E Can. & NE U.S.A., W to

Manit. & Iowa.

APION LONGIROSTRE Ol,

S & C Eur., N. Afr., W Asia. f C U.S.A., S Ontario.

^ Brown {in litt.) suggests that this identification is in need of confirmation.

List of land and freshwater animal species 59

APION SIMILE Kby.

Brit. Isl., Eur., N. Afr., Hither Asia, (f) Nfld.

W Sib.

ARAECERUS FASCICULATUS DeG.

Cosmopolitan.

t Brit. Isl., Eur. (widely). t Oreg.; E U.S.A.

Probably native of India.

AULEUTES EPILOBII Payk.

N & C Eur. {not Brit. Isl.), Sib. E at Alberta-Nov. Scot., Nfld., S to Texas,
least to Baical.

BARYNOTUS MOERENS F.

Brit. Isl., N, C & SW Eur. t Nova Scotia.

BARYNOTUS OBSCURUS F.

Brit. Isl., N & C Eur. f New Brunsw., Nov. Scot., Nfld.

BARYNOTUS SQUAMOSUS Germ, (schonherri Zett.)

Icel., Brit. Isl., NW & W Eur. f NE U.S.A., New Brunsw., Nov. Scot.,

Nfld.

BARYPITHES PELLUCIDUS Boh.

Brit. Isl., NW & W Eur. t Brit. Col.-Calif.; NE U.S.A., Nfld.

BR.\CHYSOMUS ECHINATUS Bonsd.

Brit. Isl., N & C Eur., Cauc. t Queb., Nfld.

CEUTHORRHYNCHUS ASSIMILIS Payk.

Brit. Isl., Eur., N. Afr., Cauc. f Brit. Col.-Calif.

CEUTHORRHYNCHUS ERYSIMI F.

Brit. Isl., Eur., N. Afr., W & N Asia f New York, Ontario.
E to Amur.

CEUTHORRHYNCHUS FLORALIS Payk.

Brit. Isl., Eur. only. t Nov. Scot., Nfld.

6o CHAPTER I

CEUTHORRHYNCHUS PUNCTIGER Gyll. {marginatus auctt. americ.)^

Brit. Jsl., Eur. only. t Wash.; E U.S.A., W to Missouri, E

Can.

CEUTHORRHYNCHUS QUADRIDENS Panz.

Brit. Isl., Eur., N. Afr., Cauc. f NE U.S.A.

CEUTHORRHYNCHUS RAPAE Gyll.

Brit. Isl., N & C Eur., W Asia. t Wash.-Calif.; New Engl., E Can., Nfld.

CEUTHORRHYNCHUS SULCICOLLIS Payk.

Brit. Isl., N & C Eur., N. Afr., Cauc, f Maryland.
Sib.

CLEONUS PIGER Scop.

Brit. Isl., Eur., N. Afr., Asia E to f New York, Queb., Ont.

Amur.

CRYPTORRHYNCHIDIUS LAPATHI L.

Brit. Isl., Eur., Sib. E to Amur & (f) Wash.; E U.S.A., W to Dakota.

Japan.

ELLESCHUS BIPUNCTATUS L.

Brit. Isl., N & C Eur., Asia E to f NE U.S.A., W to Mich., E Can., S

Kamch. Labr., Nfld.

ELLESCHUS scANicus Payk.
Eur. {not Brit. Isl.), ? W Sib. f Penns., New Jers.

GRYPUS (gRYPIDIUS) EQUISETI F.

Brit. Isl., Eur., Sib. E to Amur. Alas.-Labr., S to Color.

GYMNETRON ANTHIRRINI Payk.

Brit. Isl., Eur., N. Afr., Syria. f E U.S.A. W to Ohio, E Can.

GYMNETRON NETUM Germ.

C & S Eur. {not Brit. Isl.), N. Afr., W f NE U.S.A. W to Iowa.
Asia.

^ Hatch (1953, p. 27) repeats records of C. marginatus Payk. and cyanipennis Germ, from
N. America though these species were cancelled already in Leng (IV. suppl., 1939).

List of land and freshwater animal species 6i

GYMNETRON PASCUORUM Gyll.

Brit. Isl., Eur., N. Afr., Syria. f E. U.S.A.: New Jers.-Virg.

GYMNETRON TETRUM F. {teter auctt.)

Eur. {not Brit. Isl.), N. Afr., Asia E to f Brit. Col.-Oreg.; E U.S.A. W to
Baical. Iowa & Color., E Can.

LIOPHLOEUS TESSULATUS O. F. Miill.

Brit. Isl., Eur., Sib. f New York (i ex.).

MAGDALIS BARBICORNIS Latr.

Brit. Isl., Eur., Madeira. t NE U.S.A.

MECINUS PYRASTER Hbst.

Brit. Isl., Eur., N. Afr., W Asia. f Maryland.

MIARUS HISPIDULUS Lec.

(t) S Spain? {Vide Franz, 1947, p. 241- Ont., Massach.-Florida.
243).

MIARUS MERiDiONALis Bris. {consuctus Csy.)
S Eur., N. Afr. f Kansas.

MIARUS MICROS Germ.
Brit. Isl., Eur., N. Afr. . f Manitoba.

NOTARIS AETHIOPS F.

Brit. Isl., Eur., Sib. E to Amur. Alas.-Labr., Nfld., S to Color.

NOTARIS BiMACULATUS F. {wyomittgensis Chitt.)

Brit. Isl., Eur., Asia E to Amur & Oreg.-Color., S Labr., Nfld.

Kamch.

OTIORRHYNCHUS CRIBRICOLLIS Gyll.

S Eur., N. Afr. t California.

f Australia.

OTIORRHYNCHUS DESERTUS Rosenh.

Brit. Isl., NW, W & C Eur. t Nfld.

62 CHAPTER I

OTIORRHYNCHUS LIGNEUS Ol.

Brit. Isl., W Eur.: Scand. -Italy. f Maine, New Brunsw., Nov. Scot.,

Nfld.

OTIORRHYNCHUS LIGUSTICI L.

Brit. Isl., Eur. (except the south), Cauc. f N New York.

OTIORRHYNCHUS MERIDIONALIS Gyll.

France, Spain, Sizily; N. Afr. f California.

OTIORRHYNCHUS OVATUS L.

Brit. Isl., Eur., Asia E to Baical. f Brit. Col.-Calif., Utah; E Can.-New

Mex., W to Wyom.; Nfld.

OTIORRHYNCHUS PORCATUS Hbst.

Brit. Isl., W & C Eur., N to Norway. f Montreal.

OTIORRHYNCHUS RAUCUS F.

Brit. Isl., Eur. (generally), W Sib. f Ontario.

OTIORRHYNCHUS RUGIFRONS Gyll.

Icel., Brit. Isl., N, W & C Eur. f Nov. Scot., Nfld.

OTIORRHYNCHUS RUGOSOSTRIATUS Gze.

Faeroes, Brit. Isl., W & S Eur., N. Afr., f Brit. Col.-Calif., Utah, Color.; E & C
Cauc. U.S.A., Queb.

OTIORRHYNCHUS SCABER L.

Brit. Isl., N & C Eur. t Nov. Scot.

OTIORRHYNCHUS SINGULARIS L.

Brit. Isl., Eur. (widely). t Brit. Col., Wash.; New Engl., E Can.,

Nfld.

OTIORRHYNCHUS SULCATUS F.

Brit. Isl., N & C Eur. f Brit. Col.-Calif.; NE U.S.A., E Can.,

Nfld.
t Australia.

PANTOMORUS GODMANI CrOtch

t SW Eur. t S. Afr., Austral. f U.S.A. (widely), E Can.

Native of S. Amer. or Hawaii.

List of land and freshwater animal species 63

PENTARTHRUM HUTTONI Woll.

Brit. Isl., W Eur. t Montreal.

PHILOPEDON PLAGIATUS Boh.

Brit. Isl., Eur., N. Afr. f New Brunsw., Pr. Edw. Isl., Magdal.

Isl., Nfld.

PHYLLOBIUS OBLONGUS L.

Brit. Isl., Eur. (except the north), W f New York, Ontario, New Brunsw.

Asia.

PHYTONOMUS (hypera) arator L. {polygoui L.)
Brit. Isl., Eur., ?W Asia. f Illinois.

PHYTONOMUS (hYPERA) MELES F.

Brit. Isl., Eur., N. Afr., Asia Minor. f NE U.S.A., E Can., Nfld.

PHYTONOMUS (hYPERA) NIGRIROSTRIS F.

Brit. Isl., Eur., N. Afr., Asia Minor. f Brit. Col.-Oreg.; E & C U.S.A., E

Can., Nfld.

PHYTONOMUS (hypera) POSTICUS Gyll. {variabilis Hbst., murinus auctt. americ.)i

Brit. Isl., Eur., N. Afr., Sib. E to f C U.S.A., Calif, (fig. 6); since 1952

Yenisei, C Asia. also in the East.

t India.

PHYTONOMUS (hYPERA) PUNCTATUS F.

Brit. Isl., Eur., C Asia. t Brit. Col.-Calif.; NE & C U.S.A., E

Can., Nfld.

PHYTONOMUS (HYPERA) RUMICIS L.

Brit. Isl., Eur., N. Afr., Cauc. t Iowa, N. Dak., Alberta. *

POLYDROSUS IMPRESSIFRONS Gyll.

S & C Eur. {not Brit. Isl.). t NE U.S.A.

POLYDROSUS SERICEUS Schall.

Brit. Isl., Eur., N. Afr., ?Sib. t Indiana; Queb.

^ There seems to be no reason for the opinion of Lehmann & Klinkowski (1942, p. 28)
that posticus Gyll. and the true murinus F. should have been confused by North American
authors.

64 CHAPTER I

PSELACTUS (phloeophagia) spadix Hbst.

Brit. Isl., Eur., N. Afr. t NE U.S.A.

t Australia.

RHINONCUS CASTOR F.

Brit. Isl., Eur., Asia E to Japan. f Brit. Col., Oreg.; NE U.S.A. W to

Illin., E Can., Nfld.

RHINONCUS PERICARPIUS F.

Icel., Brit. Isl., Eur., N. Afr., Sib., f New York, Nfld.

Japan.

RHYNCHAENUS (ORCHESTES) SALICIS L.

Brit. Isl., Eur., Sib. E to Kamch., Wash., Calif.; E U.S.A. W to Wise,

Japan. Nfld. (Probably transamerican.)

SCIAPHILUS ASPERATUS Bonsd.

Icel., Brit. Isl., Eur., Casp. region f Brit. Col.; NE U.S.A., E Can., Nfld.

SITONA CRINITUS Hbst.i

Brit. Isl., Eur. E to Ural. t Wash.-Calif.; Nfld. (Certainly wider

distr.)

SITONA CYLINDRICOLLIS Fahr.

Brit. Isl., C & S Eur., W & C Asia. f NE U.S.A. & E Can., W to Alberta.

SITONA DiscoiDEUS Gyll. (humeralts Steph.)*
Brit. Isl., Eur., Mediterr., W & C Asia, f Rhode Isl.

SITONA HISPIDULUS F.

Brit. Isl., Eur., Cauc, Syria. f Wash.-Calif.; NE U.S.A. & E Can.,

W to Nebr., Nfld.

SITONA LEPIDUS Gyll. {flavescens Mrsh.)

Brit. Isl., Eur., Mediterr., Hither f Wash.; E U.S.A. & E Can., W to

Asia, W Sib. Kans., Nfld.

SITONA LINEATUS L.

Brit. Isl., Eur. (generally). f Brit. Col.

^ Also other European species of the crinitits group have been reported from N. America
but the identifications require confirmation.

* According to Brown {in lift.) this record requires confirmation.

List of land and freshwater animal species 65

SITOPHILUS (cALANDRA) GRANARIUS L.

Cosmopolitan,
(t) Brit. Isl., Eur.; Asia. t U.S.A., Canada.

^ SITOPHILUS (cALANDRA) ORYZAE L,

Cosmopolitan,
t Brit. Isl., Eur.; Asia. t U.S.A.

STENOPELMUS RUFINASUS Gyll.

tBrit. Isl., Holl., Belg., Germ., France. S U.S.A. N to Indiana.

STOMODES GYROSICOLLIS Boh.

C & S Eur. t Maine.

STROPHOSOMUS MELANOGRAMMUS Forst.

Icel., Brit. Isl., C & W Eur., Madeira. f Brit. Col.; NE U.S.A., E Can., Nfld.

TANYSPHYRUS LEMNAE F.

Brit. Isl., Eur., ?N Asia. Manit.-Ontar., S to Flor. (probably

Japan. . wider distr.).

TRACHODES HISPIDUS L.

Brit. Isl., Eur. (except the north). f St. Pierre (S of Nfld.; probably acci-

dental).

TRACHYPHLOEUS BIFOVEOLATUS Beck

Brit. Isl., Eur., Cauc, Macaron. isl. f Brit. Col.; New York, E Can., Nfld.

TROPiDERES (gonotropis) dorsalis Thunb. {gibbosus Lee.)
N & C Eur., Sib. E to Amur. Color., Huds. Bay, Maine, Nfld.

TROPIPHORUS CARINATUS O. F. Miill.

Brit. Isl., N & C Eur., E to C Russ. f Nfld.

TROPIPHORUS OBTUSUS BoHsd.

Icel., Brit. Isl., N & C Eur. E to the f Nov. Scot.

White Sea.

TROPIPHORUS TOMENTOSUS Mrsh.

Brit. Isl., W & C Eur. t Queb., Pr. Edw. Isl., Nov. Scot.

5 ~ 565597 Lindroth

66 CHAPTER I

TYCHius (miccotrogus) PICIROSTRIS F.

Brit. Isl., Eur. only. t Brit. Col. & Wash. -Nov. Scot, (prob-

ably transamerican).

TYCHIUS STEPHENSI Schnh. {tomentosus Hbst.)
Brit. Isl., Eur. only. t E U.S.A., E Can.

Macro-Lepidoptera — Butterflies and Moths

Lepidoptera, especially the "Macros", are the most collected of all insects and the
literature dealing with them is excessively rich. In spite of this, the attempts to
get a tolerably complete picture of Eur- American species met with greater difficul-
ties than in any other group of animals here treated. Lepidopterists generally
seem too interested in local faunas and description of new aberrations (often termed
"subspecies"). Furthermore, the perpetual change of nomenclature, in names of
all categories, and the unfortunate splitting-up of the old genera, often on small
or obscure characters, have made the modern literature on Macrolepidoptera
almost unsurveyable to the non-specialist.

Seitz' "Gross-Schmetterlinge "(1909-24) has been very useful but, of course, is
not up to date. Djakonov (1931), Valle (1933), Kozhantshikov (1937, 1950), have
given supplements for the Palaearctis. For North America, McDunnough's Check
List (1938) provided excellent service though, unfortunately, it does not give the
distribution. Many Palaearctic species are by him marked as doubtful in North
America; these I have omitted, unless the occurrence was later confirmed. For
Butterflies, Klots (1951) has been used, for some of the other groups Forbes
(1923, 1948, 1954). Krogerus (1954) and Ferguson (1955) published excellent
local faunas, of Newfoundland and Nova Scotia respectively. For additional
papers the reader is referred to the bibliography of Chapter I (p. 126 a. f.).

The difficulties mentioned above induced me to send the preliminary list of
Eur- American Macrolepidoptera to Mr. D. C. Ferguson, Halifax, and Dr. E. G.
Munroe, Ottawa. The latter co-operated, on my behalf, with Dr. T. N. Freeman
concerning the Diurna and with Dr. D. F. Hardwick concerning the Noctuids.
Mr. Ferguson also consulted Dr. J. G. Franclemont. I am highly indebted to these
gentlemen for the additions and corrections made, and also to Dr. H. Krogerus,
Helsingfors, for valuable suggestions.

Among possible Eur-American species, the following have been omitted in
await of further evidence: Chrysophanes {Heodes, Lycaend) phlaeas L. {americana

List of land and freshwater animal species 67

Harris, hypophlaeas Bdv.), Plebeius (Lycaena, Lycaenopsis) argiolus L. (pseudargi-
olus Bdv. & Lee), P. (Lycaena) glandon Prun. (aquilo Bdv., and races), Plusia
(Caloplusia) hochemvarthi Hoch., and Schrankia turfosalis Wocke.

Diurna^

ADOPAEA (tHYMELICUS) LINEOLA Ochs.

Brit. Isl., Eur. (generally), N. Afr., N f Ontario (since about 1910); Mich.,
Asia E to Amur. Ohio.

ARGYNNIS2 CHARICLEA Schn.

/. typ. sbsp.? Several sbspp.

N Scand., Kola Penins., Sib. E to Kamch. Alas.-N Queb., Nfld.,

N. Zeml. Greenl. (N to ']%°\'j')-^

ARGYNNis EUNOMIA Esp. {aphiraphe Hbn.)

/. typ. Several sbspp.

N & C Eur., Asia E to Amur. Alas.-Labr., S to Maine and, in Rocky

Mts., to Color.

ARGYNNIS FREIJA Thunb.

N Eur., Sib. E to Amur, N Japan. Alas.-Labr., Nfld., S to New Mex.

ARGYNNIS FRIGGA Thunb.

/. typ. Several sbspp.

N Eur., Sib. E to Amur. Alas.-Labr., Baffin Isl., S to Color.

ARGYNNIS IMPROBA Butl.

N Scand., N. Zeml., NW & NE Sib. Alas. & Brit. Col.-Baffin Isl.

ARGYNNIS PALES Schiff.*

Eur. (boreoalpine), C & N Asia E to Alas., Brit. Col., N. W. Terr. E to

Kamch. Baker Lake.

ARGYNNIS POLARIS Bsd.

N Eur., W & NE Sib. Alas.-Labr., S to Manit.; Ellesm. Isl.,

Greenl. (N to 81 °5o')-

^ The famous Monarch {Danaus plexippus L.), native of America, is not listed here. It
has been observed repeatedly in western Europe, above all in Britain, but the occurrence is
accidental since the food-plant of the larva, Milkweeds (Asclepias, and related genera), is
not indigenous in Europe. Vide Ford (1946, p. 152 a. f.) and Klots (1951, p. 78).

^ Included in Argynnis are here: Boloria, Brenthis, Clossiana, &c.

^ The Greenland form transgredes into/, typ. (Petersen, 1954, p. 245).

* The division into sbspp. is very confused {vide Petersen, 1954, p. 237).

68 CHAPTER I

ARGYNNIS SELENE Schiff. {ttiyrina Cram.)

/. typ. Several sbspp.

Brit. 1^1., Eur., Asia E to Kamch., Alas.-Labr., Nov. Scot., Nfld., S to N.

Kuril., Japan. Carol.

ARGYNNIS TITANIA Esp. {amathusia Esp.)

sbsp. rossica Hemm. Several sbspp. 2 sbspp.

S Finl., Balticum, Eur. C & S Eur. mts. Sib. E to Amur.

Russ.

Several sbspp.
Wash .-Nov. Scot., Labr., S along the mts. to New Mex., isolated in White Mts.
(New Hampsh.).

CARTEROCEPHALUS (pAMPHILA) PALAEMON Pall.

Brit. Isl., Eur., N Asia E to Amur, Alas.-Labr., Nfld., S to NE U.S.A.

Kamch., & Kuril.

COENONYMPHA TULLIA O. F. Miill. (tiphott Rott.) (fig. 55)

Many sbspp.

Brit. Isl., Eur. (widely), Asia E to Alas., NW Can., S to Ariz. & New Mex.,

Kamch. E to Manit., as sbsp. inornata Edw. to

Labr. & Nfld.

COLIAS HECLA Lef.

N Scand., W & NE Sib. Alas.-N Labr., Baffin & Ellesm. Isl., S to

Churchill, Man., in Rocky Mts. to 54°;
Greenl. (N to 8i°3o').

COLIAS NASTES Bdv.

N Scand., N Zeml., E Sib. Alas.-Labr., N to Victoria & Baffin Isl.,

S to S Brit. Col.

COLIAS PALAENO L.

/. typ. sbsp. chippezva Kby. (pelidneides Stgr.)

N & C Eur., Sib. E to Amur & Kamch. Alas.-Labr.

EREBIA DISA Thunb.

/. typ. sbsp. mancina Dbl. & steckeri Holl.

N Scand., N Russ., Sib. (in the East Alas.-Labr.
transgreding into the N. American
sbspp.).

List of land and freshwater animal species 69

EREBIA DISCOIDALIS Kby.

Petchora tundra (Kusnezov, 1925); Alas.-Huds. Bay, S to Minnes.

Yenisei-E Sib., S to Altai.

EREBIA FASCIATA Butl.

Petchora tundra (Kusnezov, 1925); N Arct. N. Amer.: N. W. Terr, to Huds.

Ural., Yenisei-Anadyr. Bay, Victoria Isl.

HESPERIA (ERYNNIS, AUGIADES) COMMA L.

/. typ. Several sbspp.i

Brit. Isl., Eur. (generally), Asia E to Alas.-Labr., Nfld., S to Color.

Amur & China.

oeneis jutta Hbn.

/. typ. Several sbspp.

N Eur. S to E. Pruss., Sib. E to Amur Alas.-Nov. Scot., Nfld., S to New Engl.
& Kamch.

PAPILIO MACHAON L.

Several sbspp. sbsp. aliaska Scudd. (joannisi Ver.) &

hudsoniamis Clark^.
Brit. Isl., Eur. (generally), N. Afr., Alas.-Huds. Bay & Lake Sup.

Asia E to Kamch., China & Japan.

pieris napi L.

Several sbspp. Several sbspp.

Brit. Isl., Eur. (generally), N. Afr., Asia Alas.-Labr., Nfld., S to New York &

E to Amur, China, Kamch. & Kuril. Calif.

PIERIS RAPAE L.

Brit. Isl., Eur. (generally), Asia E to f Transamerican (except N Can.), Nfld.,

Amur, Japan, Kuril. S to Mexico.

t Hawaii, Austral.

PLEBEius (lycaena, lycaeides) IDAS L. {argytognomon auctt., nee Bergstr.)

Several forms 3 sbspp.

Eur. (except Brit. Isl.), N & C Asia E Alas.-Nov. Scot., Labr., Nfld., S along

to China & Kamch. the mts. to Color.

^ Often, in part at least, kept as separate species.

2 Forms from NE N. America {brevicauda Saund., &c.) are regarded as specifically
distinct.

yo CHAPTER I

PLEBEius (lycaena) optilete Knoch

/. typ. (incl. several forms) sbsp. yukona HoU.

N & C Eur., N Asia E to Amur, Kamch. NW N. Amer. E to Alberta.
& Kuril.

PYRGUS (hesperia, syrichtus) centaureae Ramb.^

Scand., N Russ., W & S Sib. (prob- Brit. Col.-Labr., Nfld., S to Color,

ably farther east).

VANESSA (nYMPHALIS) ANTIOPA L.

Brit. Isl. (immigr.), Eur. (generally), N. Alas.-Nfld., S to Venez.
Afr., N & C Asia E to Amur & Japan.

VANESSA (pYRAMEIS) CARDUI L.

Cosmopolitan.

Brit. Isl., Eur. (generally, but immigr. Entire N. Amer. (except the Arctic,
in the north & west), Afr., Asia E to immigr. in the north).

C Amer.
Japan & Kuril., Austral.

VANESSA (pYRAMEIS) ATALANTA L.

Brit. Isl., Eur. (generally, but immigr. Transamerican in U.S.A. & S Can.
in the north & west), N. Afr., W & C (immigr. in the north).
Asia E at least to Altai. C Amer.

Sphinges

celerio galii Rott. {gallii auctt.)

/, typ. sbsp. intermedia Kby.

Eur. (generally), N & C Asia E to Brit. Col.-Labr., Nfld., S to Calif. &

Japan, Kuril. Georgia. Mexico.

CELERIO LINEATA F.

sbsp. livornica Esp. /. typ.

Eur. (immigr. only), Afr., S & E Asia. S Can.-U.S.A.-C & S Amer. (as an

immigr. N to Brit. Col., Montr. & Nov.

Scot.).

SPHINX (hyloicus) pinastri L.

Brit. Isl., Eur. (generally). Sib. E to f NE U.S.A., Alberta (doubtfully estab-

Japan. lished).

^ In the East of N. Amer., Manit.-Labr., is freija Warr., usually regarded as distinct
species and doubtfully recorded also from the Scandinavian mountains.

List of land and freshwater animal species 71

Bombycimorpha
ARCTiA (hyphoraia, platyprepia) alpina Quens.

N Fennoscand., Sib. S to Altai, E at N Alaska, NW N. W. Terr,

least to Baikal.

ARCTIA caja L.
/• typ.
(incl. several forms) sbsp. phaeosoma Btlr. 5-6 sbspp.

Eur. (generally), Asia E E Asia, to Amur, Japan, Alas. & Brit. Col.-Labr, &
to Pamir, S to N India. Kuril. Nov. Scot., S to Color. &

Utah.
Transgreding forms in E Asia.

ARCTIA (hyphoraia, parasemia) lapponica Thunb. (festiva Bkh.)

/. typ. sbsp. gibsoni B. H. & hyper borea Curt.

N Eur., Sib. E at least to Lena R. N. W. Terr., NE U.S.A.

ARCTIA (ORODEMNIAS, APANTESIS) QUENSELI Payk.

/. typ. sbsp. norvegica 2-3 sbspp. sbsp. gelida

Strd. Moeschl.

C Eur. mts. N Fennoscand. N Mong., Lena R. Alas., Yukon Terr.

-Amur, N Japan. & Brit. Col.-Labr.,

New Hampsh.;
Greenl.

EUPROCTis (porthesia) chrysorrhoea L. {similis Fuessl.)

Brit. Isl., Eur. (except the north), N & f U.S.A. (not established), Ont. (green-

C Asia E to Amur, Japan, Sakhal. & house).

Kuril.

EUPROCTIS (nygmia) phaeorrhoea Don. {chrysorrhoea auctt., nee L.)

Brit. Isl., Eur. E to Ural, N. Afr., f E Can., Nov. Scot., NE U.S.A.

Cauc, Hither Asia.

GLUPHISIA CRENATA Esp.

/. typ. sbsp. septentrionis Wlk.

N & C Eur., Sib. E to Amur. Pacif. coast-Nov. Scot., Nfld., S to

Georgia.

HYPHANTRIA CUNEA Drury

t Hungar., Yugosl., Austria. S Can., U.S.A. (generally), S to Texas.

t Japan.

72 CHAPTER I

ORGYIA (nOTOLOPHUS) ANTIQUA L.

Icel., Eur. (generally), N. Afr., Cauc, Brit. Col.-Nov. Scot., Nfld., S to New
N Asia E to Amur, Corea & Japan. York.

t Sumatra, S S. Amer.

PARASEMIA (nEMEOPHILA) PLANTAGINIS L.

Many sbspp. Many forms (? sbspp.)

Brit. Isl., Eur. (widely), Asia E to Yukon Terr. & Brit. Col.-Ont., S to

Kamch., Japan & Kuril. Color.

PHRAGMATOBIA FULIGINOSA L.

/. typ. sbsp. borealis Stgr. Several sbspp. sbsp. rubricosa

Harr.
W & C Eur. N Fennoscand. Asia E to Amur & Brit. Col.-Nov.

Kamch. Scot. & Nfld., S

to S. Carol.

STILPNOTIA (lEUCOMA) SALICIS L.

Brit. Isl., Eur. (generally), N. Afr., f Brit. Col.-Oreg.; E Can., Nov. Scot.,

Asia E to Lena R. & N China. NE U.S.A.

UTETHEISA BELLA L.

(t) Wales (Skokholm Isl., i ex.; E Can., Nov. Scot., E U.S.A. (widely).

Williams, 1950).

Noctuae (Phalaenae)

ACRONYCTA (apATELA) LEPORINA L.

4

/. typ. 4 sbspp.

Eur. (generally), N & C Asia E to Brit. Col.-Nov. Scot., N U.S.A. S to

Amur, Sakhal. & N Japan. Color.

AMPHIPYRA (pYROPHILA) TRAGOPOGONIS L.

Brit. Isl., Eur. (generally), N. Afr., Asia (f) S Can., Nov. Scot., Nfld., N U.S.A.
E to W Sib. & N India. W at least to Mississ.; Brit. Col. Pos-

sibly transamerican.

ANARTA cordigera Thunb.

N & C Eur., Sib. (at least Amur). Alas.- Labr., Nov. Scot., S to Mass. &

Color.

List of land and freshwater animal species 73

ANARTA (sYMPiSTis) FUNEBRis Hbn. (funesta Payk.)

N Fennoscand., Alps; NE Sib., N Alas.-Labr. & Nov. Scot., S to New

Japan. Hampsh. & Wyom.

ANARTA (SYMPISTIS) LAPPONICA Thunb. (fig. 30)

N Fennoscandia, N. Zeml. N. W. Terr., Labr., Baff. Isl., Greenl.

ANARTA (lasiestra) leucocycla Strg. (incl. staudlngerl Aur.)

N Fennoscandia. Brit. Col., N. W. Terr. & Victoria Isl.-

Labr., Nfld., Baff. & Ellesm. Isl., S to
Color., isolated in White Mts. (New
Hampsh.); Greenl.

ANARTA (SYMPISTIS) MELALEUCA Thunb.

Fennoscandia; NE Sib. Mackenzie Delta-Labr., S to Gaspe.

ANARTA MELANOPA Thunb.

Shetl., Brit. Isl., Eur. (boreoalpine), E Brit. Col.-Nfld., New Hampsh., S along
to Kola Penins. the mts. to New Mex.

ANARTA RICHARDSONI Curt.

sbsp. dovrensis sbsp.? sbsp. asiatica Stgr. /. typ. & 3 sbspp.

Stgr.

N Fennoscand., W Sib. E Sib. Alas.-Labr., Grin-

N. Zeml. nell L., Greenl. (N

to 8i°3o').

ANARTA (SYMPISTIS) ZETTERSTEDTI Stgr.

N. Fennoscandia; Mongol, (sbsp.?). N. W. Terr. & Brit. Col.-Labr., Baff.

Isl., Greenl.

ANOMOGYNA (GRAPHIPHORA) LAETABILIS Zett.

N Fennoscandia; NE Sib., Sakhal. Brit. Col. & Yukon-Labr.

ANOMOGYNA RHAETICA Stgr.

/. typ. shsp. fennica Brdt. sbsp. homogena McD. &

sbsp. conditoides Benj.
Alps. N Fennoscandia. Brit. Col.-Labr., Nfld.,

New Hampsh.

ANOMOGYNA SINCERA H.-S.

Fennoscandia, C Eur. mts.; Sib.: Amur. Alberta-Labr., Nfld., NE U.S.A.

74 CHAPTER I

ANOMOGYNA (aplectoides) SPECIOSA Hbn.

/. typ. sbsp. mixta Walk.

Eur. (boreoalpine), N & C Asia E to N. W. Terr.-Labr., Nfld., NE U.S.A.

Kamch.

CHLOANTHA (lithomoia, calocampa) solidaginis Hbn.

/. typ. 3 sbspp.

Brit. Isl., N & C Eur., Sib. E to Amur. Alas. & Brit. Col.-Queb.

CRYMODES (HADENA, APAMEA) EXULIS Lef. (fig. 38)

Icel., Faeroes, Shetl., Scotl., Spitzb. N. W. Terr.-Labr., Nfld., Ellesm.

Isl., Greenl.

CUCULLIA LUCIFUGA Schiff.

/. typ. sbsp. intermedia Speyer

N & C Eur., N & C Asia to E Sib. N U.S.A., S Can. W at least to Alberta,

Nfld., S to Virg.

heliothis (melicleptria, canthylidia) scutosa Schiff".

C & S Eur., N. Afr., W & C Asia E W U.S.A., W Can., E at least to Alberta

to China & Corea. & Kansas, S to Color.

HiLLiA (oRTHOSiA, crasia) IRIS Zett. (crasis H.-S.)

N Fennoscandia, N Russia; E Sib.: Brit. Col.-Labr. & Nov. Scot., S to

Amur. Color. & NE U.S.A.

HYDROECIA MICACEA Esp.

Brit. Isl., N & C Eur., N Asia E to f NE U.S.A., E Can., Nov. Scot., Nfld.

Amur & Japan.

LAPHYGMA (caradrina) exigua Hbn. {flavimaculata Harv.)

8 Eur., Afr., W & C Asia E to Man- f Florida-Calif.; straggling E to Wise. &
chur. & Japan; S Asia. Kans.

t C Amer., Hawaii.

LEUCANIA (SIDERIDIS, HELIOPHILA) PALLENS L.

/. typ. sbsp. luteopallens Sm.

Eur. (generally), N. & C Asia E to Alas.-Nfld., S to the Gulf of Mexico.

Amur & Kamch.

List of land and freshwater animal species 75

LEUCANIA (SIDERIDIS, HELIOPHILA, PSEUDALETIA, CIRPHIS) UNIPUNCTA Haw.

Almost cosmopolitan.

(t) Brit. Isl., C & S Eur., C Asia, SE U.S.A. & S Can., Brit. Col.-Nov. Scot.,

Sib., Japan. Nfld.

(t) S Asia, Austral., Pacif. Isl. 3 sbspp. in C & S Amer.

MAMESTRA (sCOTOGRAMMA) TRIPOLI I Rott.

Eur. (generally), N. Afr., N & C Asia, Transamerican in U.S.A. & Can., Nov.
Japan. Scot., Nfld., S to Mexico. S. Amer.

PARASTiCHTis (hadena, septis) basilinea F. (fimtima Gn.)

Eur. (generally), N & C Asia E to Brit. Col.-Nfld., S to Color. & NE

Amur & Japan. U.S.A.

PARASTICHTIS (hadena, agroperina, septis) lateritia Hfn.

Eur. (generally), N & C Asia E to Yukon Terr. & Brit. Col.-Nfld., N

Amur, Japan & Kuril. U.S.A. (generally).

PLUSIA (SYNGRAPHA, AUTOGRAPHA) DIASEMA Bsd.

Fennoscandia; NE Asia: Amur & Yukon, N. W. Terr. & Alberta-Labr.,

Kamch. Greenl.

PLUSIA (pHYTOMETRA, AUTOGRAPHA, SYNGRAPHA) INTERROGATIONIS L.

Icel., N & C Eur., Asia E to Kamch. Alas. & Yukon-Labr. , Greenl.
& Kuril. NE U.S.A.

PLUSIA (SYNGRAPHA, AUTOGRAPHA) MICROGAMMA Hbn.

N & C Eur.; Kamch. Brit. Col.-Nov. Scot.

PLUSIA (chrysoptera, polychrysia) moneta F.

Brit. Isl., Eur. (except the north), N & (f) Alberta, Sask., Mont.
C Asia E to N China, Vladiv. & Kuril.

PLUSIA (AUTOGRAPHA, SYNGRAPHA) PARILIS Hbn. (fig. 31)

N Fennoscandia. N. W. Terr & Alberta-Labr., BaflF. &

Ellesm. Isl., Greenl.

PYRRHIA UMBRA Hfn.

N & C Eur., Asia E to SE Sib., Japan N U.S.A. (widely) S to N. Carol., Can.
& Kuril. W at least to Alberta, Nov. Scot., Nfld.,

S to Color.

76

CHAPTER I

RAPHIA PRATER Grote

(t) Engl. (Harpenden, i ex.; Williams, Can. & U.S.A. (widely), Nov. Scot., S
ig^o). to Color. & Texas.

RHYACIA (GRAPHIPHORA, NOCTUA, DIARSIA) C-NIGRUM L.

Eur. (generally), Cauc, N & C Asia E Transamerican in U.S.A. & Can., Nfld.
to Amur, Japan, Sakhal., Kuril.; S Asia. C Amer.

RHYACIA (actebia, ochropleura) fennica Tausch.

N Eur., N & C Asia E to Amur, Alas.-Nov. Scot., Nfld., N U.S.A.

Corea, Kamch. & Sakhal.

RHYACIA (PERIDROMA, lycophotia) margaritosa Haw. (saucia Hbn.)

Icel., Brit. Isl., Eur. (except the north), U.S.A. (generally), S Can., Nov. Scot.,

N Afr., Cauc, C Asia E at least to W Nfld.

Turcest. S Amer. (original patria?)

RHYACIA (diarsia) mendica F. (festiva Schiff., dislocata Sm.)

Icel., N & C Eur., Cauc, N & C Asia Brit. Col.-Labr., Nfld.
E to Amur, Kamch. & Sakhal.

RHYACIA (eUROIS, PERIDROMA) OCCULTA L.

Icel., N & C Eur., N & C Asia E to Brit. Col.-Nov. Scot., Nfld., S to Color.,

Kamch., Corea & Japan. Greenl.

RHYACIA (ochropleura. DIARSIA, PERIDROMA) PLECTA L.

Eur. (generally), N & C Asia E to Brit. Col.-Nov. Scot., Nfld., U.S.A.

Amur, Japan, Sakhal. & Kuril. (widely) S to Texas.

S Asia. t S. Afr.

RHYACIA (anaplectoides, EUROIS, aplecta) prasina SchiflF.

Eur. (generally), Cauc, N & C Asia E Brit. Col.-Nfld., N U.S.A. S to Color,
to Amur, Japan & Sakhal.

RHYACIA (pachnobia) tecta Hbn.

/. typ. sbsp. roosta Sm.

N Eur., Eur. Russia; Kamch. Alas. & Yukon-Labr.

RHYACIA (eUXOA, AGROTIS) YPSILON Rott.

Cosmopolitan.
Eur. (generally), Cauc, W, N & C Asia Brit. Col.-Labr., U.S.A. (generally),
E to Vladiv., Corea & Sakhal. Nfld.

S Asia, Austral., Hawaii, C & S Amer.

List of land and freshwater animal species 77

SCHOYENIA (anarta, archanarta, graphiphora, anomogyna) quieta Hbn.

N Fennoscandia; ace. to Valle (1933, p. N. W. Terr. & Baffin Isl.
100) in N Russ. & NE Asia.

scoliopteryx libatrix L.

Eur. (generally), Asia E to Kamch. Brit. Col.-Nov. Scot., Nfld., U.S.A.

(generally).

sPAELOTis (triphaena, amphitrota) clandestina Harr. {unicolor Wlk.)

sbsp. suecica Aur. /. typ.

Fennoscandia, N. Russia. Amur & Kamch.; Brit. Col.-Nov.

Scot., Nfld., S to Color. & Missouri;

Greenl.

xanthia (cosmia) lutea Strom (flavago F.)

Eur. (generally), N & C Asia Brit. Col.-Nov. Scot., S to Penns.

E to Kamch.

Geometrae

BOARMiA (ectropis) crepuscularia Hbn.
Eur. (widely), Asia E to Amur. Brit. Col.-Nov. Scot., S to W Virg.

CALOCALPE (hYDRIA) UNDULATA L.

N & C Eur., N Asia E to Kuril. Alas.-Nov. Scot., Nfld., S to N. Carol.

& Ariz.

CARSIA PALUDATA Thunb. {sororluta Hbn.)

N & C Eur., N & C Asia E to Amur Brit. Col.-Nov. Scot., Labr., Nfld., mts.
& Kamch. of NE U.S.A.

CHEIMATOBIA (oPEROPHTERA) BRUMATA L.

Icel., Brit. Isl., N & C Eur., Cauc; E f Nov. Scot, (f) Greenl.
Sib.

cidaria (xanthoroe) abrasaria H.-S.

/. typ. sbsp. congregata Wlk.

N Eur., W & NE Sib. Pacif. coast & Yukon Terr.-Nov. Scot.,

Nfld., S to New York.

cidaria (perizoma) alchemillata L.
Eur. (widely), Asia E to Altai. t Nfld. (Ferguson).

78 CHAPTER I

ciDARiA (epirrhoe) ALTERNATA O. F. MuIl.

Icel., Eur. (generally), Cauc, N Asia Alas.-Nov. Scot., Labr., Nfld., S to
E to kamch. & Kuril. Distr. of Col.

CIDARIA (dysstroma) citrata L. (tmmanata Haw.)

Icel., N & C Eur., N & C Asia E to Alas. & Brit. Col.-Nov. Scot., Labr.,

Kamch. & Kuril. Nfld., S to New York.

CIDARIA (xANTHOROE) FERRUGATA CI.

Eur. (generally), N & C Asia E to Brit. Col.-Nov. Scot, Nfld., S to Penns.

Kamch.

CIDARIA (hydriomena) furcata Thunb. (sordidata F.)

Icel., N & C Eur., N & C Asia E to Pacif. coast-Nov. Scot., Labr., N U.S.A.

China, Japan & Kamch.

CIDARIA (eULYPE, RHEUMAPTERA) HASTATA L.

Icel., N & C Eur., N & C Asia E to Alas.-Nov. Scot., Labr., Nfld., S to

China & Kamch. Penns.

CIDARIA (tHERA) JUNIPERATA L.

Brit. Isl., Eur. (widely). f Maine, Ontar., Nov. Scot.

CIDARIA (SPARGANIA, XANTHOROE) LUCTUATA Schiff.

/. typ. sbsp. obductata Moeschl.

N & C Eur., N & C Asia E to Kamch. Pacif. coast-Labr., Nfld., S to N. Carol.

CIDARIA (xanthoroe) munitata Hbn.

Icel., Faeroes, Eur. (boreoalpine), Sib. Pacif. coast-Nov. Scot., Nfld.
E to Amur & Kamch.

CIDARIA (pERCNOPTILOTA) OBSTIPATA F.

Almost cosmopolitan.

Eur. (except the north), Asia (widely). Pacif. coast-Nov. Scot., Nfld., S to the
Afr. Gulf of Mexico. S. Amer.

CIDARIA (dASYURIS) POLATA Dup.

N Eur., W & NE Sib. to Kamch. Alas.-Labr., Baffin & Ellesm. Isl., also

in Color.; Greenl.

List of land and freshwater animal species 79

ciDARiA (hydriomena) ruberata Frr.

Brit. Isl., N & C Eur. (Not known Brit. Col.-Nov. Scot., Nfld., S to Color,
from Asia but possibly confused with
coerulata F.)

CIDARIA (psychophora) sabini Kby. (incl. frigidaria Guen.)

N Fennoscandia, N. Zeml.; NE Asia Alas.-NE arct. N. Amer., Greenl.

to Kamch.

CIDARIA (diactinia) silaceata SchifF.

N & C Eur., Asia E to Kamch., Japan. Brit. Col. & Yukon -Nov. Scot., NE

U.S.A.

CIDARIA (eulype) subhastata Nolck.

/. typ. 2 sbspp.

N & C Eur., Sib. E to Kamch. Brit. Col.-Nov. Scot., Labr.

CIDARIA (dysstroma) truncata Hufn.

N & C Eur., Asia E to China. Alas. & Brit. Col.-Labr., Nfld., S to

Calif. & New York.

CIDARIA (colostigia) turbata Hbn.
N & C Eur., C & NE Asia E to Kamch. Alas. -Alberta.

CIDARIA (euphyia) unangulata Haw.

/. typ. sbsp. intermediata On.

N & C Eur., Sib. E to Kamch., Japan. Pacif. coast-Nov. Scot., Nfld.

ematurga atomaria L.

/. typ. sbsp. amitaria Guen.i

Eur. (generally), W, N & C Asia E to E Can., Nov. Scot., S to Penns., W at
Amur & Kamch. least to Alberta.

eupithecia albipunctata Haw. (tripunctaria H.-S.)

N & C Eur., Cauc, Sib. E to Amur. Brit. Col. & Calif.-Nov. Scot. & Labr.,

Nfld., S to Penns.

eupithecia castigata Hbn.

Eur. (widely), Asia E to Amur. Brit. Col.-Nov. Scot., Nfld., S to N.

Carol.

^ Often regarded as distinct species.

So CHAPTER I

EUPITHECIA GELIDATA Moeschl. {hyperboreata Stgr.)

N & northern C Eur.; ?Asia. Alas.-Nov. Scot., Labr., S to New Mex.,

Greenl.

EUPITHECIA INTRICATA Zett. Qielveticaria Bsd.)^

/. typ. sbsp. gibsonata Tayl.i

N & C Eur.; ?E Sib. Alberta-Ontar., New Brunsw., NE

U.S.A.

EUPITHECIA LARICIATA Frr.

/. typ. sbsp. bifasciata Dyar sbsp. luteata Pack.

N & C Eur., Cauc, Sib. Brit. Col. E Can.-Nov. Scot., Nfld.,

E to Transbaic. E U.S.A.

EUPITHECIA PALUSTRARIA Dbl. {pygttiaeata Wan., obiimbrata Tayl.)
N & C Eur., Lena R., Amur. Brit. Col.-Gaspe & Labr., S to Color.

EUPITHECIA SATYRATA Hbn.

/. typ. 3 sbspp.

Icel., N & C Eur., C Asia, ?Amur. Alas.-Nov. Scot., Labr., S to N. Carol.

EUPITHECIA SOBRINATA Hbn.

Icel., N & C Eur.; Kamch. ? Brit. Col.; E Can., Nov. Scot., E

U.S.A. S to Texas.

EUPITHECIA UNDATA Frr. {scriptaHa H.-S.)

N Fennoscandia, mts. of C Eur. & Brit. Col.-Labr., S to Color.

Hither Asia, C & NE Asia.

HIMERA (COLOTOIS) PENNARIA L.

Brit. Isl., N & C Eur., Cauc, Asia f E Canada (established?).

Minor.

ITAME FULVARIA Vill.

N & C Eur., Sib. E to Kamch., Japan. Pacif. coast-Nov. Scot., Nfld., S to

Massach.

ITAME (dYSMIGIA) LORICARIA Ev.

/. typ. sbsp. Julia Hulst

N Eur., Sib. E to Kamch. Yukon Terr.-Nov. Scot., S to Wyom. &

New York.

^ E. arceuthata Frr., likewise reported from both continents, is doubtfully distinct from
intricata Zett.

List of land and freshwater animal species 8i

LYGRIS TESTATA L.

N & C Eur., N Asia to E Sib. Brit. Col.-Nov. Scot., S to New Jers,

OPORiNiA (epirrita) autumnata Gn.

N & C Eur., Sib. E to Kamch. & Brit. Col.-Nov. Scot., Nfld., S to Penns.

Kuril.

scopuLA (acidalia) frigidaria Moeschl.

sbsp. schoeyeni Sp.-Schn. /. typ.

N Fennoscandia, N Eur. Russ. Kamch.; Yukon Terr.-Labr., Nfld.

VENUSIA CAMBRICA Curt.

N & C Eur., N & C Asia E to Kamch., Alas.-Nov. Scot., Nfld., S to N Calif. &
Japan & Kuril. New York.

Aegeriidae

AEGERIA (TROCHILIUM) APIFORMIS CI.

Brit. Isl., Eur. (generally), Cauc, W f NE U.S.A.

Asia E to Altai.

SYNANTHEDON (CONOPIA, THAMNOSPHECIA) CULICIFORMIS L.

Brit. Isl., Eur. (generally), Cauc, Sib., Alaska.
N China.

SYNANTHEDON (CONOPIA, RAMOSIA) TIPULIFORMIS CI.

Brit. Isl., Eur. (generally), Cauc, Sib. f Can. & U.S.A., W to Alberta & Wash.

f Australia.

Cossidae

ZEUZERA PYRINA L.

Brit. Isl., Eur. (except the north); f NE U.S.A.

N. Afr., Asia (except the north),
E to Corea & Japan. India.

t S. Africa.

Hepialidae

HEPIALUS (hepiolus) ganna Hbn.
/. typ. sbsp. hyperboreus Moeschl. ^

(inch several forms)
N & C Eur., Asia to E Sib. Alas.-Ontar.,? Labr., NE U.S.A.

^ Often regarded as distinct species.
6 — 365597 Lindroth

§2 CHAPTER I

Diptera — Two-winged flies

Only 'two groups of blood-sucking flies, the mosquitoes {Culicidae, CuUcinae) and
the horse flies & clegs {Tabanidae) were considered sufficiently known for the
present purpose. Information about the former was provided, for the Old World
by Stackelberg (1937), Marshall (1938), Natvig (1948), and Montshadsky (1951),
for North America by Dyar (1928), Matheson (1944) and Carpenter & La Casse
(1955). The Tabanids were mainly treated according to Olsoufiev (1937) and
Philip (1947). Mr. H. Kauri, Lund, made valuable suggestions on the synonymy
of this family.

Culicidae, Culicinae — Mosquitoes

AEDES (stegomyia) aegypti l. {fasciatus F.)
Cosmopolitan in tropic & subtropic regions.

t Brit. Isl. (accidentally). t U.S.A., S of about 40° N.

f S peninsulae of Eur.

Native of Africa.

aedes cataphylla Dyar
N & C Eur., Cauc, W & E Sib. Alas.-Color, E to Manit. & Wyom.

aedes cinereus Meig.

/. typ. sbsp. fusciis O. S.

Brit. Isl., Eur. (widely), Cauc, Sib. Alas.-Labr., Nov. Scot., S to Color.

E to Amur.

AEDES COMMUNIS DeG.

Brit. Isl., Eur. (widely), Sib. E to Alas.-Labr., Nov. Scot., S to Color.

Kamch.

AEDES DI ANTAEUS H., D. & K.

N & C Eur., Sib. E to Ussuri. Alas.-Labr., Nov. Scot., NE U.S.A., S

to New York & Wyom.

AEDES DORSALIS Meig.

Brit. Isl., N & C Eur., Cauc, N Asia Brit. Col.-Queb., S to Color. & Delaw.
E to Amur & N China.

AEDES EXCRUCIANS Wlk.

N, C & SE Eur., W & N Asia E to Alas.-Nov. Scot., Nfld., S to Color.

Ussuri.

List of land and freshwater animal species 83

AEDES FLAVESCENS O. F. Miill.

Brit. Isl., Eur. (widely), W & N Asia Alas.-Labr., S to Color.
E to Kamch.

AEDES IMPIGER Wlk. (nec Dyar) {nearcticus Dyar)

N & C Fennoscand.; Sib.: Taimyr, Alas.-Hudson Bay, S to Montana;

Kolyma & New Sib. Isl. EUesmere L.

AEDES INTRUDENS Dyar

N & C Eur., E to Ural, E Sib., Kamch. Brit. Col.-E Can. & NE U.S.A., S to

Color.

AEDES NIGRIPES Zett.

Spitzb., N Fennoscand., N Russ., N Alas.-N Queb.; Greenl.

Sib. E to Kamch.

AEDES PULLATUS Coq.

Eur.(boreomontane) E to Volga; Alas.-Queb., S to Color.

Kamch. (Stackelberg, not in Mont-

shadsky).

AEDES FUNCTOR Kby.

Brit. Isl., N & C Eur., Sib. E to Alas.-Labr., Nov Scot. S to Color.

Kamch.

AEDES RiFARius Dyar & Kn.
N & C Eur. E to Ural; E Sib. " Alas.-Ontar. & New York, S to Color.

AEDES STicTicus Meig. (hirsuteron Theob.)
Brit. Isl., N & C Eur., W & E Sib. Alas.-Queb. & New Engl., S to Texas.

AEDES V^XANS Meig.

Brit. Isl., Eur. (widely), W & N Asia Yukon & Brit. Col.-Nov. Scot., S to
E to Ussuri. Color. & Texas.

E & S Asia, Austral.

CULEX MOLESTUS Forsk.i

Brit. Isl., Eur. (widely), Russia (without Calif .-New York (limits not determined),
further details)., N. Afr.

^ Often regarded as a form of pipiens L.

84 CHAPTER I

CULEX PIPIENS L.
Brit. Isl., Eur. (generally), N. Afr., W Brit. Col.-Nov. Scot.; U.S.A. (generally).
& N Asia E to Amur & China.

S. Amer., E & S Afr.

CULEX TERRITANS Wlk. (apicalis auctt., nee Adams)

Brit. Isl., N, C & SE Eur., N. Afr., Alas.-New Engl., S to Flor. & Texas.

Cauc, W. Turcest.

CULISETA (tHEOBALDIA) ALASKAENSIS Ludl.

Brit. Isl.,^N & C Eur., Cauc, Sib. E Alas.-Queb. & Labr., S to Color,

to Kamch.

CULISETA (theobaldia) morsitans Theob.

Brit. Isl., N, C & SE Eur., Cauc., Brit. Col.-Labr., Nov. Scot. & NE

W Sib. U.S.A., S to Color.

Tabanidae — Horse flies & clegs

CHRYSOPS NIGRIPES Zett.

N Fennoscand., Eur. Russ. S to Lenin- Alas.-Maine.
grad, Sib. E to Kamch. & Sakhal.

TABANUS (hybomitra, tylostypia) aequetinctus Becker (flavipes Wied.)

NE Russ., Sib. E to Kamch. & Wise, Labr.

Anadyr.

TABANUS (hybomitra, tylostypia) lapponicus Wahlb. {astutus O. S.i, hyphus

Whitn.)
N & C Eur., Sib. E to Kamch. Brit. Col.-Labr., NE U.S.A.

TABANUS (hybomitra, tylostypia) sexfasciatus Hine {anderi Kauri^, borealis auctt.,

in part.)
N Fennoscand. (distr. imperfectly Alas., Labr.

known).

Hymenoptera — Wasps, &c.

Only a few groups of the Aculeata were considered sufficiently known for the
present purpose. In all of them, the catalogue of Muesebeck, Krombein & Townes
(195 1) provided an excellent source of knowledge.

^ This synonymy was established by Mr. H. Kauri.

List of land and freshwater animal species 85

Apidae, Bombinae — Bumblebees

Besides the catalogue of 1951, Lutz & Cockerell (1920) has been used for the
Nearctic, Skorikov (1937), Henriksen (1937), and Pittioni (1942-43) for the
Circumpolar fauna.

BOMBUS ARCTICUS Kby.^
N Eur., N. Zeml., Sib. E to Kamch. Alas.-Labr., Greenl. (N to 8i°5o').

BOMBUS BALTEATUS Dahlb.

/. typ. (and related forms). sbsp. kirbiellus Curt, (and related forms)

N Eur., N. Zeml., Sib. E to Kamch. Alas.-Labr., in Rocky Mts. S to Ariz.

& Bering Strait.

BOMBUS HYPERBOREUS Schnh.

N Eur., N. Zeml., Sib. E at least to Alas.-Baffin L., Greenl. (N to 81 "30').

Lena R.

BOMBUS LAPPONICUS F.

/. typ. (and related forms) sbsp. sylvicola Kby.^ (and related forms)

Brit. Isl., Eur. (boreoalpine), N Asia Aleut. Isl., Alas.-Labr., Baffin L., in

E to Kolyma & Kamch., S to Pamir. Rocky Mts. S to Ariz.

Crabronidae

Here all necessary information was gathered from Leclercq (1954, p. 126, &c.).

CROSSOCERUS (COELOCRABRO) AMBIGUUS Dahlb.

Brit. Isl., Eur. E to Ural, N China, f E Can., NE U.S.A., S to N. Carol.,

Hokkaido. W to Iowa.

ECTEMNIUS (hYPOCRABRO) CONTINUUS F.

Brit. Isl., Eur. (generally), N. Afr.,Asia Alas.-E Can., U.S.A. (generally).
E to Amur & Kamch., Japan. Mexico.

ECTEMNIUS DIVES Lep. & Brull.

Brit. Isl., Eur. (widely). Sib. E to Brit. Col.-E Can., U.S.A. to Color.

Kamch., Japan.

^ In the catalogue of 195 1 (p. 1253) and, doubtfully, by Lutz & Cockerell (1920, p. 507)
arcticus Kby. is regarded as a synonym oi hyperboreus Schnh. Cf. Pittioni, 1942 (p. 182, a.f.).
^ Often regarded as distinct species.

g6 CHAPTER I

ECTEMNius (cLYTOCHRYSUs) LAPiUARius Panz. (comptus Lep. & Brull.)

Brit. Isl., Eur. (generally), Cauc, W Alas.-Labr., Nfld., U.S.A. (generally).

Sib., Amur, Sakhal., Japan.

ECTEMNIUS (CLYTOCHRYSUS) NIGRIFRONS CreSS.

Brit. Isl., Eur. (generally), W Sib., Wash.-E Can., Nfld., U.S.A. (generally).

Kamch., Sakhal., Japan.

ECTEMNIUS (mETACRABRO) QUADRICINCTUS F.

Brit. Isl., Eur. (except the north). t E Can., NE U.S.A. S to Maryl., W to

Illin.

RHOPALUM (CRABRO, EUPLILIS) CLAVIPES L.

Brit. Isl., Eur. E to Ural. (t) Wash.-Calif.; NE U.S.A.

RHOPALUM (CRABRO) COARCTATUM Scop.

Brit. Isl., Eur. (widely), Sib. E to Brit. Col.-E Can., N U.S.A.

Baical.

Vespidae, Vespinae — True Wasps

The records for North America were obtained from the 195 1 catalogue, those for
the Old World mainly from Berland (1928) and Weyrauch (1937).

VESPA (vespula) adulterina Buyss.

/. typ. sbsp. arctica Rohw.

Mts. of C Eur., Cauc, Mongol., Sakhal. Alas.-N. Carol.
Brit. Col.-Color.

VESPA (vespula) austriaca Panz.
N & C Eur., Asia E to Kamch. Alas.-New Jers., S to Color.

VESPA CRABRO L. (incl. Several forms)

Brit. Isl., Eur. (widely), N. Afr., W & f Atl. States of U.S.A. (sbsp. germana
NAsia. Christ., of C Eur.).

VESPA (vespula) norvegica F.

/. typ. sbsp. albida Slad. sbsp. norvegicoides Slad.

Eur., Sib. E to Kamch. Alas.-Labr. Alas.-N. Carol.

& Sakhal.

List of land and freshwater animal species 87

VESPA (vESPULA) RUFA L.

/. typ. sbsp. schrenki Rad. 6 sbspp.

{sibirica And.)

Eur., W & C Asia E to E Sib., Mong., China. Alas.-Nov. Scot., S to

Kamch. Ariz.

\^SPA (vespula) vulgaris L.
Eur., Palaearctic Asia. Alas.-N. Carol., S to New Mex.

Formicidae — Ants

There is a good deal of difference of opinion among North American myrmecolo-
gists concerning the taxonomic relationship of Nearctic and Palaearctic forms
within such genera as Formica, Leptothorax, Lasius, Myrmica, &c. Marion R,
Smith, in the Catalogue of 1951, represents a more liberal view in this respect.
On the recommendation of Mr. I. H. H. Yarrow, British Museum, I have here
followed the more restrictive position taken by Creighton (1940). Genus Lasius
has been treated according to Wilson's monograph (1955) though with re-estab-
lished subspecific names, consistently abandoned by this author.— The distri-
bution in Eurasia was mainly compiled from Donisthorpe (1927) and Stitz (1939).

ANERGATES ATRATULUS Schenck {friedlandi Creighton)

Brit. Isl. (i loc), C Eur. N to S (t)i Connect.-Virginia.

Sweden, W & E Sib.

CAMPONOTUS HERCULEANUS L.

N & C Eur., Cauc, N & C Asia E to Alas.-Nfld., N U.S.A., S (in the mts.)
Amur, China & Sakhal. to New Mex. & Penns.

t Brit. Isl.

CAMPONOTUS NEARCTicus Emery (pavidiis Wheel.)
t Brit. Isl. Brit. Col.-Calif.; NE U.S.A., S to Tex.

CAMPONOTUS PENNSYLVANICUS DeG.^

t Brit. Isl. E Can., E half of U.S.A., S to the Mex.

Gulf.

^ Vide p. 234.

^ Regarded as sbsp. of hercideanus L. by Smith (195 1).

88 CHAPTER I

CREMATOGASTER LINEOLATA Say

t Brit. Isl. E Can., E U.S.A., S to Color. & Flor.

(often indoors).

FORMICA ciNEREA Mayr
/. typ. sbsp. lepida Wheel. ^

Eur. (except Brit. Isl. & the north), W Can.-Calif. & Utah, E to N. & S.

Cauc, Asia Minor. Dakota.

FORMICA FUSCA L.

Brit. Isl., N & C Eur., N & C Asia E Alas.-Labr., Nfld., N U.S.A. S to Ariz,
to Amur, China, Kamch. & Sakhal.

FORMICA RUFIBARBIS F.

/. typ. sbsp. occidua^ Wheel. sbsp. gnava Buckl.i

Brit. Isl., Eur. (widely), Wash.-Calif. Utah-Tex., Mexico.

Cauc, N. & C Asia E to
Lena R. & Manchur.

FORMICA SANGUINEA Latr.

/. typ. sbsp. siibnuda Emery^

Brit. Isl., Eur. (generally), Cauc, N & Alas.-Labr., Nfld., NE U.S.A., in Rocky
C Asia E to Amur, Japan. Mts. S to Ariz.

IRIDIOMYRMEX HUMiLis Mayr
Cosmopolitan.

t Brit. Isl., W & C Eur. (indoors). f S U.S.A., N to S. Carol.

Indigenous in S. Amer.

LASius ALiENUS Foerst.
/. typ. sbsp. americanus Emery^

(pallitarsus Prov.)
Brit. Isl., Eur. (widely), N. Afr., Cauc, Brit. Col. -Nov. Scot, (with gaps), S to
N & C Asia, China, Japan. Ariz. & Flor.; Mexico.

LASIUS FLAVUS F.

/. typ. sbsp. hrevicornis Emery^

Brit. Isl., Eur. (except the Wash.-Sask., S to E Can. & U.S.A. E of

north), Cauc, Asia Minor, Color. Rocky Mts., S to N. Carol.

Sib. E to Amur, Japan. & Tenness.

^ Possibly distinct species, according to Yarrow {in lift.).

2 Smith (1951) recognizes 5 North American sbspp. oi sanguinea Latr. Wilson & Brown
(1955) regard sanguinea Latr. as a purely Eurasian species.

' Wilson (1955, p. 62 a.f.) in principle does not use subspecific names in Lasius but the
Palae- and Nearctic forms are (in part) identical in flavus F. only.

List of land and freshwater animal species 89

LASIUS NIGER L.

/. typ. sbsp.i

Brit. Isl., Eur. (generally), N. Afr., Wash.-New Mex. (in the mts.).

Cauc, Sib. E to Amur & Kamch.,
China, Japan.

LASIUS UMBRATUS Nyl. (incl. mixtus Nyl.)

/. typ. sbsp.i

Brit. Isl., Eur. (widely), Cauc, Hither E Can., Nov. Scot., E U.S.A., W to
Asia. (Distr. in N & C Asia uncertain.) Idaho, S to Flor. & Ariz.

MONOMORIUM DESTRUCTOR Jerd.

Cosmopolitan.

t Brit. Isl. (hothouses). t SE U.S.A.

Indigenous in S Asia.

MONOMORIUM PHARAONIS L.

Cosmopolitan.

t Brit. Isl., Eur. (widely, indoors). f Can. & U.S.A. (widely, indoors;

fieldadaption in Flor.).
Probably indigenous in S Asia.

MYRMICA LAEVINODIS Nyl.

Brit. Isl., N & C Eur., N Asia to E f Queb., Mass.

Sib., Manchur., Japan.

MYRMICA LOBICORNIS Nyl.

/. typ. (incl. several forms) sbsp. lobifrons Perg. sbsp. fracticornis Emery
Brit. Isl., N & C Eur., C Alas.-Ariz. W Can.-Nfld., S to New

Asia. Mex.

ODONTOMACHUS HAEMATODA L.

F. typ. almost cosmopolitan in the tropics.
/. typ. 4 sbspp.

t W Eur. (hothouses). Not introduced Southernmost U.S.A., W Indies, Mexico,
from N. Amer.!

PARATRECHINA (PRENOLEPIS) LONGICORNIS Latr.

Cosmopolitan.

t Brit. Isl., W Eur. (indoors). t E U.S.A., N to New York (outdoors

in the Gulf Coast area only).
Probably indigenous in S Asia.

^ Wilson (1955, P- 62 a. f.) in principle does not use subspecific names in Lasius but the
Palae- and Nearctic forms are (in part) identical in flavus F. only.

go CHAPTER I

PARATRECHINA (NYLANDERIA) VIVIDULA Nyl.

Cosmopolitan.

t Brit. Isl., W Eur. (hothouses). t SE U.S.A. (in hothouses farther north).

Indigenous in S. America.

PHEiDOLE ANASTASii Emery

t Brit. Isl., W & C Eur. (hothouses). t E Can., E U.S.A. (fieldadaption in S

Flor.).
From C America.

POLYERGUS RUFESCENS Latr.

/. typ. 3-4 sbspp.

Eur. (except Brit. Isl.), N to S Sweden, Brit. Col.-Calif. & Ariz., E to Ontar. &
C Asia. Mich.

PONERA COARCTATA Latr. (fig. 35)

/. typ. sbsp. pennsylvanica Buckl.

Brit. Isl., S & C Eur., Cauc, N. Afr. Queb.-Gulf of Mexico, W to S. Dak. &

Kans.
Two further sbspp. are recorded from Africa & Austral.

PONERA ERGATANDRIA For.

sbsp. bondroiti For. /. typ.

t Belgium (hothouses). Texas, Flor.; W. Indies.

Not introduced from N. Amer. !
Probably indigenous in S. Amer.

PONERA OPACiCEPS Mayr
t Brit. Isl. (hothouses). S & SW U.S.A., C. & S. America.

TAPINOMA (TAPINOSOMA) MELANOCEPHALUM F.

Cosmopolitan in the tropics.

t Brit. Isl, (hothouses). t Flor., Georgia.

Probably indigenous in S. America.

tetramorium caespitum L.

Brit. Isl., Eur. (generally), N. Afr., (t)^ Calif.; NE U.S.A., S to Tenness.,

Cauc, W, N & C Asia E to Lena R., W to Nebr.
China, Japan.

2-3 sbspp. in tropical Africa.

^ Vide p. 234.

List of land and freshwater animal species 91

TETRAMORIUM GUINEENSE F.

Cosmopolitan in the tropics.

t Brit. Isl., C Eur. (hothouses). f S Flor., Texas, in hothouses farther

north.
Possibly indigenous in W Africa.

TETRAMORIUM SIMILLIMUM F. Smith

Cosmopolitan in the tropics.

t Brit. Isl. (hothouses). S Flor., in hothouses farther north.

(Intr. from N. Amer.?)

Native country unknown.

TRIGLYPHOTRIX STRIATIDENS Emery

Almost cosmopolitan.

t Brit. Isl., W Eur. (hothouses). f SE U.S.A. N to S. Carol.

Indigenous in S Asia.

WASMANNIA AUROPUNCTATA Rog.

t Brit. Isl. (hothouses). f S Flor., Calif.

Indigenous in C & S America.

Odonata — Dragonflies

For North America the new works of Walker (1953) and Needham & Westfall
(1955) have been consulted. The more scattered Old World literature is repre-
sented by Valle (1932, 1952) and Ander (1950). The almost cosmopolitan Pantala
flavescens F. seems not to have been found with certainty in Europe as yet and
was therefore omitted.

AESCHNA COERULEA Stroem

/. typ. sbsp. septentrionalis Burm.

Eur. (boreo-alpine), W Sib.-Lena R. Alas.-Nova Scot., Nfld., S to N. Hampsh.

AESCHNA JUNCEA L.

/. typ. sbsp. orientalis Bart. sbsp. ?

Eur., Asia E to Amur, NE Sib., Sakhal., Japan. Alas.-Labr., Nfld., S to

? Kamch. Color.

gZ CHAPTER I

AESCHNA SUBARCTICA Walk.

sbsp. interlineata Ander sbsp. elisabethae Djak. /. typ.

W & C Eur., N to S N & E Eur., E to Ar- Brit. Col.-Nova Scot., S

Sweden. chang. (Possibly confused to Michig.

with juncea in N Asia.)

ENALLAGMA CYATHIGERUM Charp.

Eur., entire N Asia, E to Kamch. Alas.-Nfld., S to Calif. & Utah.

LESTES DRYAS Kby.

Eur., Sib. E to Ussuri, Japan. Alas.-Nova Scot.

LIBELLULA QUADRIMACULATA L. (fig. 51)

Eur., Sib. E to Kamch., Sakhal., Alas.-Nova Scot., Nfld., S to Ariz.

Japan, S to Kashmir.

SOMATOCHLORA SAHLBERGI Tryb.

N Fennoscand. (Petsamo, Kola), N Alas., N. W. Terr.

Sib. (Yenisei, Lena R.).

SYMPETRUM DANAE Sulz.

Eur., Sib. E to Kamch., Ussuri, Japan. Alas.-Nfld., S to Calif. & Color.

Orthoptera Saltatoria — Grasshoppers & Crickets

The North American records were mainly compiled from Blatchley (1920), those
from the Old World from Ander (1951), Holdhaus (1954), &c.

AREOPEDELLUS (dASYHIPPUS) VARIEGATUS Fisch.

Eur. (boreoalpine), in the north from Alaska.
N Finl. through Sib. E to Kamch.

gryllotalpa gryllotalpa L. {vulgaris Latr.)

Brit. Isl., Eur. (ecxept the north), N. f NE U.S.A.

Afr., W & C Asia.

GRYLLUS (gRYLLULUS) DOMESTICUS L.

t Brit. Isl., Eur. (generally; indoors). f E U.S.A., E Can., W to Rocky Mts.

N. Afr., W & S Asia.

List of land and freshwater animal species 93

MELANOPLUS (pODISMA) FRIGIDUS Boh.

Eur. (boreoalpine), N Asia E to Man- Alas., N. W. Terr,
chur., Kamch. & Sakhal.

ROESELIANA ROESELII Hag.

Brit. IsL, C & S Eur. E to Ural, f Montreal (probably intr. with aircraft;

W Sib. Urquhart & Beaudry, 1953, p. 78).

TACHYCiNES (diestrammena) asynamorus Adcl. (mamioratus Deli., japanicus Blatch.)

Cosmopolitan.

t Icel., Brit. Isl., Eur. (widely; in hot- f U.S.A., E Can. (widely; in hothouses),
houses). t Greenl.

Probably indigenous in E Asia.

TETRix (tettix, acrydium) subulatus L. (granulatus Kby.)

Eur. (generally), W, N & C Asia, E to Brit. Col.-Nova Scot., S to New Jersey
Lena R. & Amur. & Kansas.

Araneae — Spiders

The literature dealing with the taxonomy and distribution of spiders is extensive
but very scattered and the list of Eur- American species here repesented is prob-
ably incomplete. Roewer's catalogue (1942, 1954) is most useful but the records
of distribution are quite summary. They were supplemented, for Europe by
Simon (1914-37), Reimoser (1919), Holm (1950), for northern Asia by Cha-
ritonov (1932, 1936), for North America by Chamberlin & Ivie (1947), Comstock
& Gertsch (1948), Gertsch (1949), Hackman (1954), and several other papers
listed in the bibliography (p. 126 a.f.).

Earlier, less complete lists of Eur-American species were published by Berland
(1932, p. 379) and Gertsch (1949, p. 259). My preliminary list, compiled from the
literature, was sent to Dr. W. J. Gertsch, New York, who kindly corrected and
enlarged it to a great extent. I also received many valuable suggestions from
Mr. H. Kauri, Lund, and Dr. A. Holm, Uppsala. The final list here given deviates
from the opinion of Dr. Gertsch only in so far that a few more species are
regarded as introduced; these are marked with a cross in brackets.

Erigone arctica White and E. atra Bl. have been excluded from the list on the
advice of Dr. Holm.

Q^ CHAPTER I

Dysderidae

DYSDERA CROCATA C. L. Koch

Cosmopolitan.

(t) Brit. Isl., S Eur. N to N France & f U.S.A. (widely), S Can.; Bermudas.
S Russ., Cauc, N. Afr.

Scytodidae

SCYTODES THORACICA Latr.

t Brit. Isl., Eur. (widely, indoors), f U.S.A. E of Mississippi, E Can.

E to S Russ.

S & E Asia.

Pholcidae

PHOLCUS PHALANGIOIDES Fuessl.

Cosmopolitan.

Brit. Isl., Eur. (widely, indoors), f Entire U.S.A., S Can. (usually in-

E to Russ., Cauc. doors).

Afr., Asia. t S. Amer.

PHOLCUS OPiLiONOiDES Schrank (maniieh Gertsch)
C & S Eur., S Russ., Turkest., China. f NE U.S.A.

psiLOCORUS (PSHYSOCYCLUS) siMONi Berl. {simplicior Ch. & Iv.)
t France, Engl, (indoors). Calif., Oreg.

Theridiidae

CRUSTULiNA STICTA Cambr.
Brit. Isl., NW, W & S Eur. N U.S.A., S Can.

CTENiuM (robertus) lividum Blackw.

Faeroes, Brit. Isl., Eur. (widely), N. Alaska.

Zeml., W & N Asia E to Kamch.

DIPOENA PRONA Menge {hamata Tullgr.)
Brit. Isl., N & C Eur., Hung., Corfu. (f) U.S.A. (widely).

STEATODA BIPUNCTATA L.

Icel., Brit. Isl. (mostly indoors), Eur. f Mass.; E Can., Nfld.

(widely), W & N Asia E to Kamch.

List of land and freshwater animal species 95

TEUTANA CASTANEA CI.

N, C & S Eur. {not Brit. Isl.), Turkest. f Ontario (hothouses).

TEUTANA GROSSA C. L. Koch

Almost cosmopolitan,
(t) Brit. Isl., Eur. (widely), W Sib. f Brit. Col.-S Calif.; N. Carol.-Tex.

TEUTANA TRIANGULOSA Walck.

C & S Eur. {not Brit. Isl.), N. Afr., f U.S.A. (widely), S Can.

Cauc, W. Turkest.

THERIDIUM BERKELEYI Em.

Sweden. Calif.-New Jers., Ontario.

THERIDIUM (nEOTTIURA) BIMACULATUM L.

Brit. Isl., Eur. (widely), W Sib. f Brit. Col., Wash.

THERIDIUM IMPRESSUM L. Koch

Brit. Isl., Eur., Cauc, W & N Asia Alaska-Alberta & N. W. Terr.

E to Kamch.

THERIDIUM MELANURUM Hahn {denticulatiim Walck.)

Brit. Isl., Eur. (generally), N. Afr., W Brit. Col.-Calif. & Utah.
& C Asia.

THERIDIUM OHLERTI Thor. {iimbratlcum L. Koch, liindbecki Sor., simulatum Em.)

N & C Eur. (boreoalpine). Sib. Alas., Can., Calif. & New Mex. (moun-

(Yenisei). tains); Greenl.

THERIDIUM ORNATUM Hahn {pictum Walck., zelotypum Em.)

Brit. Isl., Eur. (generally), N. Afr., Sib. N. W. Terr., Manit., Utah.
E to Kamch.

THERIDIUM (enoplognatha) ovatum CI. {redimitum CI., lineatum L.)

Brit. Isl., Eur. (widely), N. Afr., Cauc, f Brit. Col.-Calif.; Nov. Scot.-New
W. Turkest.; Japan. York.

THERIDIUM PETRAEUM L. Koch

C & S Eur., N. Afr. (f) Wash.-Calif.; Ontario, Maine.

96

CHAPTER I

THERIDIUM SIMILE C. L. Koch

Brit. Isl., Eur. (widely), E to S Russ., t Brit. Col.-Wash.
Mediterr.

THERIDIUM (achaearanea) tepidariorum C. L. Koch
Cosmopolitan.

t Icel., Brit. Isl., Eur. (widely, hot- f U.S.A. (widely) & S Can. (indoors in

houses); W Sib., China. the north).

Originally tropical.

THERIDIUM (enoplognatha) thoracicum Hahn
Brit. Isl., Eur. (widely), N. Afr. f Oregon,

THERIDIUM TINCTUM Walck.

Brit. Isl., Eur. (generally), Cauc; t Wash., Oregon.

Japan.

THERIDIUM VARIANS Hahn

Icel., Brit. Isl., Eur. (generally), N. f Brit. Col.-Wash.

Afr., W & N Asia E at least to Yenisei.

theridula gonygaster SIM. {opulenta auctt. eur., nee Walck.)
Almost cosmopolitan in the tropics.

•f France, Corsica (probably not in- Florida,

troduced from America).

Afr., Madag.; Japan. W. Ind., C. & S. Amer.

Nesticidae

NESTicus CELLULANUS CI. {terrestris Em.)
Brit. Isl., Eur. E to C Russ. f NE U.S.A.

NESTICUS (eidmanella) pallidus Em.

t England. Brit. Col., entire U.S.A., Nov. Scot.,

Nfld.
t Hawaii. C & S Amer.

Linyphiidae

agyneta cauta Cambr.
Brit. Isl., Eur. (boreoalpine). NE U.S.A., E Can., Nfld.

List of land and freshwater animal species 97

BATHYPHANTES (STYLOPHORA) CONCOLOR Wid.

Brit. Isl., Eur. (generally), E to S Russ., (f) NE U.S.A., W at least to Wise, E
Madeira, W Sib. Can., Nfld.

BATHYPHANTES GRACILIS Blackw.

Brit. Isl., Eur. (except the south). Sib. (f) Nfld.
E to Kamch. & Sakhal.

BATHYPHANTES (STYLOPHORA) NIGRINUS Westr.

Brit. Isl., Eur. (widely), E to S Russ., N U.S.A., W to Oreg., E Can.
W Sib.

BATHYPHANTES PULLATUS Cambr. (kiiratai Ch. & Iv.)

Brit. Isl., N & W Eur., Eur. Russ.; Alas., Wyom., Queb., Nfld.

Kamch.

BATHYPHANTES SETIGER F. Cambr. {hyperborciis Holm, josephus Ch. & Iv.)
Brit. Isl., N & C Eur. Alaska.

CENTROMERUS (CENTROMERITA, CENTROMERINUS) BICOLOR Blackw.

Faeroes, Brit. Isl., N & C Eur. E to (f) Nfld.

Moscow.

CENTROMERUS SYLVATICUS Blackw.

Brit. Isl., N & C Eur., Sib. E to (f) NE U.S.A., E Can., Nfld.

Kamch.

ESTRANDiA (linyphia) grandaeva Keys, {tridens Schenk., nearctica Banks)

N Fennoscand.; Kamch. Alas.-Nova Scot., Labr., Nfld., S to N.

Carol.

helophora (linyphia) insignis Blackw.

Brit. Isl., N & C Eur., Sib. E to Alas.-Nova Scot., Nfld., S to Penns. &

Kamch. Wise.

lepthyphantes complicatus Em. {umbraticola Keys., audax Sor.)
Icel., Scotl., Eur. (boreoalpine), Spitzb. Alas., Labr., Nfld.; Greenl.

lepthyphantes leprosus Ohl.

Icel., Faeroes, Brit. Isl., Eur. (generally), (f) N U.S.A., Nova Scot., Nfld.
Hither Asia; Kamch.

7 ~ 565597 Litidroth

gS CHAPTER I

LEPTHYPHANTES MINUTUS Blackw.

Ikit. Isl., N & C Eur. E to Eur. Russ. E U.S.A. W to Mich.

LEPTHYPHANTES NEBULOSUS Sund.

Brit. Isl., Eur. (widely), Sib. E to (f) U.S.A. (widely), W at least to Mon-

Kamch. tana & Utah; Nov. Scot.

LINYPHIA MARGINATA C. L. Koch {triangularis Walck.)

Brit. Isl., Eur. (widely). Hither Asia, Alas.; Nov. Scot., Nfld., S to Texas.

W Sib.; China, Japan.

MACRARGUS (microneta) multesimus Cambr.
N Scand., Sib. NE U.S.A.

microlinyphia (linyphia, pusillia) impigra Cambr. {cayuga Em.)
Brit. Isl., C Eur., W Sib. Alas.; Ont., New York.

microneta (micronetaria) viaria Blackw.

Brit. Isl., Eur. (generally) E to Volga, Alas.-E Can., S at least to Illin. & New
N. Afr., Hither Asia. York.

OREONETIDES vaginatus Thor. (adipatus L. Koch)

Brit. Isl., Eur. (boreoalpine), N. Zeml., Alas., Labr., NE U.S.A., Nfld.; Greenl.
Sib. E to Kamch.

Erigonidae (Micryphantidae)

AULACOCYBA SUBITANEA Cambr.

Icel., Brit. Isl., W Eur. N to Sweden, (f) Mass.
N. Afr.

COLLINSIA (coryphaeolana) holmgreni Thor. {mendica L. Koch)

Spitzb., Jan Mayen, Icel., Faeroes, Alas.; Labr., Baffin L., Greenl.

Scotl., Eur. (boreoalpine), N. Zeml.,
Sib. E to Kamch.

COLLINSIA (microerigone, typhochraestus) spetsbergensis Thor. {oxycephala

L. Koch)

Spitzb., Icel., N Scand., N. Zeml., Sib. Alaska; Greenl.
E to Lena R.

List of land and freshwater animal species 99

CORNICULARIA cuspiDATA Blackw. {breviculu Cby. & Bish.)

Icel., Faeroes, Brit. Isl., N & C Eur.; (f) New York, Nfld.
Kamch.

CORNICULARIA KARPINSKII Cambr.

Spitzb., Icel., Brit. Isl., Eur. (boreo- Labr., Nfld., Greenl.

alpine). Sib. E to Kamch.

CORNICULARIA UNICORNIS Cambr.
Brit. Isl., N & C Eur. E to Moscow. (f) Nfld.

DiPLOCENTRiA (scotoussa) bidentata Em. {rivalis Cambr.)

Icel., Brit. Isl., Eur. (boreoalpine), Alas.-Labr., Nfld., S to Color.

E to Czechosl.

diplocephalus cristatus Blackw.

Icel., Faeroes, Brit. Isl., Eur. (generally), (f) NE U.S.A. W to Wise, E Can.,
N. Afr., Sib. E to Kamch. Nfld.

t New Zeal.

DISMODICUS BIFRONS Blackw.

/. typ. sbsp. decemoculatus Em.

Icel., Brit. Isl., N & C Eur., Sib. E to Alas.-Nfld., N U.S.A.
Transbaic.

entelecara (mythoplastoides) media Kulcz. (sombra Ch. & Iv.)
Eur. (boreoalpine). Sib. E to Kamch. Alaska.

eperigone maculata Banks
t Switzerl. (hothouse). E U.S.A. W to Kansas, Nfld.

erigone psychrophila Thor.

Spitzb., Fr. Jos. L., Icel., Faeroes, N Alas., Grant L., EUesm. L., Greenl.

Scand., N. Zeml., Sib. E to Kamch.

erigone tirolensis L. Koch

Spitzb., Jan Mayen, Icel., Scotl., Eur. Alas.; Labr., Baffin L., Ellesm. L.,

(boreoalpine), N. Zeml., Sib. E to Greenl.

Kamch.

GONATIUM RUBENS Blackw.

Icel., Faeroes, Brit. Isl., Eur. (widely), (f) Ontar., E U.S.A., S to Flor., W at
W Sib. least to Wise.

lOO CHAPTER I

HILAIRA FRIGIDA Thor.

Jan Mayen, Icel., Faeroes, Brit. Isl., N Labr., Baffin L., Greenl.
Eur.;' Kamch.

HILAIRA (utopiellum) herniosa Thor. {mirabiUs L. Koch, consimilis Cambr.)
Eur. (boreoalpine), W Sib. Arct. Can., Nfld.

HYPSELISTES FLORENS Cambr.
Brit. Isl., W Sib. Alas., S Can., N U.S.A.

ISLANDIANA (ADUVA) ALATA Em.

N Scand. Alas.-NE U.S.A. (mts.), S to Wyom.;

Nfld.

MASO (caseola) sundevalli Westr. {herbicola Em.)

Icel., Brit. Isl., Eur. (widely). Sib. Alas., NE U.S.A., E Can., Nfld.

E to Kamch.

MONOCEPHALUS (thyreosthenius, hormathion) PARASITICUS Westr. {becki Cambr.,

limnatus Cby. & Bish.)

Icel., Brit. Isl., N & C Eur. NE U.S.A., W to Wise, Nfld.

MINYRIOLOIDES TRIFRONS Cambr.

/. typ. sbsp. aquatilis Cby. & Bish. (affinis

Schenk.)
Brit. Isl., N & W Eur.; Kamch. New York.

POCADICNEMIS PUMILA Blackw.

Brit. Isl., Eur. (generally), E to Moscow, Alas.-E Can., N U.S.A.

RHAEBOTHORAX (typhochraestus) borealis Jacks. (incl. thori Jacks.)
Spitzb., Icel., N Scand. Labr., Greenl.

RHAEBOTHORAX PAETULUS Cambr.

Eur. (boreoalpine). Alas.; Greenl.

SISICUS APERTUS Holm {longitarsi Ch. & Iv.)
N Scand., Switzerl. Alaska.

TIBIOPLUS NEARCTICUS Ch. & Iv.

N Sweden. Alaska.

List of land and freshwater animal species k

TRACHYNELLA (trACHYNOTUS) NUDIPALPIS WcStr.

Icel., Faeroes, Brit. Isl., Eur. E to (f) Nfld.

Moscow.

TRiCHOPTERNA (pELECOPSis, lophocarenum) mengei Sim. {excovata Em.)
Brit. Isl., N & C Eur. (not in Russ.). (f) NE U.S.A., E Can., Nfld.

WALCKENAERA (spiROPALPUS, cornicularia) vigilax Blackw. (spiralis Em.)

Brit. Isl., Eur. (widely), E to Volga and E U.S.A., W to Wyom. & Missouri, S
Crimea. to Louis. & Georg.

zornella (linyphia) cultrigera L. Koch

Fennoscand., W Sib. Alas. & Brit. Col.-NE U.S.A., S to

Utah; Nfld.

Araneidae (Argiopidae)

ARANEUS CORNUTUS CI. (foHatus Fourc.)

Icel., Brit. Isl., Eur. (generally), N & C Alas.-E Can., N U.S.A., Nfld.; Greenl.
Asia E to Kamch., China.

ARANEUS DiADEMATUS CI. (diadema L.)

Icel., Brit. Isl., Eur. (generally), N & C f Brit. Col. & Wash.-New Engl., Nov.
Asia E to Kamch. Scot., Nfld.; Greenl.

ARANEUS DISPLICATUS HentZ

sbsp. zvestringi Thor. ' /. typ.

Brit. Isl., Eur. (widely but local). Sib. Alas.-Nfld., S to Georgia.

E to Lena R.

ARANEUS MARMOREUS CI. {raji Scop.)

Brit. Isl., Eur. (generally), Cauc, Sib. Alas., N U.S.A., E Can.
E to Amur & Kamch.

ARANEUS NORDMANNI Thor.

N, C & E Eur. (not Brit. Isl.); Trans- Alas. & Oreg.-E Can., NE U.S.A.
baic.

ARANEUS PATAGIATUS CI. (ocellatus CI., dumetorum Vill.)

Icel., Brit. Isl., Eur. (generally), N. Alas.-Labr., Nfld., N & NE U.S.A.;

Afr., W & N Asia E to Kamch. Greenl.

I02 CHAPTER I

ARANEUS QUADRATUS CI.

Brit. Isl., Eur. (generally), Sib. E to Alaska, Yukon; Greenl.

Kamth.

ARANEUS sCLOPETARius CI. (sertcatus CI., oviger Panz.)

Icel., Brit. Isl., Eur. (widely) E to E U.S.A. (generally) W at least to Wise,

Volga & S Russ., C Asia, China, Japan. E Can., Nfld.

CERCIDIA PROMINENS WeStr.

Brit. Isl., Eur. (widely) E to Volga; E Can., NE U.S.A. W at least to Wise.

Kamch.

CYCLOSA CONICA Pall.

Brit. Isl., Eur. (generally), N. Afr., Alas.-Nfld., N U.S.A.

W & N Asia E to Transbaic.

META MENARDI Latr.

/. typ. sbsp. ovalis Gertsch

Brit. Isl., Eur. (except the east), N. NE U.S.A. W to Wise., E Can., Nfld.

Afr., W & C Asia,
t Madagascar.

ZYGiELLA (zilla) calophylla Walck. {atrica C. L. Koch)
Brit. Isl., Eur. E to S Russ.; Sakhal. f E U.S.A., E Can.

ZYGIELLA (zilla) MONTANA C. L. Koch (nec auctt. americ.)

Mts. of C Eur., Cauc, W. Turcest.; Brit. Col.-Calif.

Kamch,

ZYGIELLA (zilla) x-notata CI. (Htterata Ol.)

Brit. Isl., Eur. (generally) E to S Russ., f Alaska-Calif.; E Can.-New Engl.
C Asia, Sakhal. f S. Amer.

Tetragnathidae

PACHYGNATHA CLERCKi Sundev. {sewardi Ch. & I v.)

Brit. Isl., Eur. (generally). Sib. E to Alaska.

Kamch., Japan

TETRAGNATHA EXTENSA L. (groenlatidtca Thor., manitoba Ch. & Iv.)

Icel., Brit. Isl., Eur. (generally), N. Alas.-Nov. Scot., Labr., Nfld., N U.S.A.;

Afr., W & N Asia E to Kamch., China, Greenl.

Japan.

List of land and freshwater animal species 103

Agelenidae

TEGENARIA AGRESTis Walck. {magnacava Exl.)
Brit. Isl., Eur. E to C Russ, t Wash.

TEGENARIA ATRICA C. L. Koch {saeva Blackw., gigantea Ch. & Iv.)
Brit. Isl., Eur. (widely) E to S Russ. f Brit. Col.

TEGENARIA DOMESTICA CI. (derhami Scop.)
Almost cosmopolitan.

Icel., Faeroes, Brit. Isl., Eur. (generally), f U.S.A. & S Can. (widely).
N. Afr., N. Zeml., Sib. E to Kamch.,

Japan. t C & S Amer.

India, Austral., Hawaii.

TEGENARIA LARVA Sim. {praegrandts Fox)

Brit. Isl., C & S Eur. (probably over- f Distr. of Columbia,
looked).

TEGENARIA PAGANA C. L. Koch (simplex Bryant, antrias Ch. & Iv.)
Brit. Isl., Eur., N. Afr., Hither Asia. f Calif., Tex., Alab.

Lycosidae

ALOPECOSA (tarentula) aculeata CI. {beuni Em.)

N & C Eur. (not Brit. Isl.), N & - Alas. & Oreg.-Maine, Nfld.

C Asia E to Kamch.

ARCTOSA (lYCOSA) ALPIGENA Dol.

Icel., Brit. Isl., Eur. (boreomontane), Alas.-Labr., Nfld., S to Ariz.; Greenl.
WSib.

PARDOSA (lycosa) palustris L. (tarsalis Thor., andersoni Gertsch)

Icel., Faeroes, Brit. Isl., Eur. (gener- Alaska,
ally), N. Zeml., N & C Asia E to Kamch.
& Kuril.

PARDOSA (lycos.'v) saltuaria L. Koch (incl. hyperborea Thor.)

Icel., Eur. (boreoalpine), E to E. Prussia Alas. -Maine, Labr., Nfld.; Greenl.
& Kola Penins.

I04

CHAPTER I

PIRATA iNSULARis Em. (ptccolo Dahl)

N & C Eur. (not Brit. Isl.), E to C E U.S.A., W at least to Wyom., Utah &

Russ.' (possibly overlooked). Color.; Nfld.

PIRATA PIRATICUS CI.

Icel., Faeroes, Brit. Isl., Eur. Alas.-Nfld., N U.S.A.; Greenl.

(generally), N. Afr., W Sib.

TROCHOSA (lycosa) terricola Thor.

/. tvp. sbsp. orophila Ch. & sbsp. pratensis Em.

Gertsch
Faeroes, Brit. Isl., Eur. Alas., W Can., Rocky E Can., N & NE U.S.A.,

(generally), Sib. E to Mts. Nfld.

Kamch.

Gnaphosidae

GNAPHOSA MUSCORUM L. Koch {conspersa Thor.)
N & C Eur. {not Brit. Isl.), W Sib. Alas.-Nfld., N U.S.A.

GNAPHOSA ORITES Chamb. (humilis Holm, holmi TuUgr., labradorensis Fox)
N Scandinavia. Alas.; New Engl., Queb., Labr.

HAPLODRASSUS (drassodes) signifer C. L. Koch

Icel., Faeroes, Brit. Isl., Eur. Alas.-Nfld., S to Ariz.; Greenl.

(generally), Hither Asia, W. Sib.

HERPYLLUS (SCOTOPHAEUS) BLACKWALLI Thor. {pillS Chamb.)

Brit. Isl., W & C Eur. (mostly indoors), f Wash. -Calif., Louisiana.
Macaron.

SOSTOGEUS (scoTOPHAEUs) LORiCATUS L. Koch (zygethus Ch. & Gertsch)

C & E Eur. {not Brit. Isl.), Turcest., f Conn., lUin., Wise, Utah.

Sib. E to Yenisei.

ZELOTES (drassyllus) rusticus L. Koch {femoralis Banks, razumoivskii Pav.)

Brit. Isl., C & S Eur., N. Afr., Cauc, f Wash.-Calif., Ariz.; Conn., Illin.,
W Sib.; Tongking. Kans.

ZELOTES SUBTERRANEUS C. L. Koch

Eur. (widely, but not Brit. Isl.), W & N Alas.-Nfld., S to Ariz.
Asia E to Kamch.

List of land and freshwater animal species 105

Clubionidae

CHEIRACANTHIUM MILDEI L. Koch

C & S Eur., Mediterr., Cauc. f NE U.S.A., Utah.

CLUBiONA KULCZiNSKii De Less. (tntermontana Gertsch)
Eur. (boreoalpine); Kamch. Brit. Col.-New Engl., S to Wyom.

CLUBIONA LUTESCENS Westr.

Brit. Isl., Eur. (widely), W Sib.; Japan, f Wash.

CLUBIONA NORVEGiCA E. Str. {humlda Jacks., calif ornica Fox)
Brit. Isl., N Scand. Alas.-Calif., Nfld.

CLUBIONA PALLiDULA CI. {holosericea L.)
Brit. Isl., Eur. (widely), Cauc, W Sib. f Brit. Col.-Wash.

CLUBIONA TRiviALis C. L. Koch {obtusa Em.)
Faeroes, Brit. Isl., Eur. (boreoalpine). Alas.-Maine, S to Utah.

MiCARiA PULICARIA Sund. {montana Em.)

Brit. Isl., Eur. (widely). Sib. E to (t) N U.S.A., W to Oreg., S to Ariz.,

Kamch. Can. (widely), Nfld.

Xysticidae

CORIARACHNE VERSICOLOR Keys.

t SW France (i ex.). E U.S.A. & E Can., W to Manit. &

Rocky Mts., Nov. Scot., Nfld.

MISUMENA VATiA CI. {calycina L.)

Brit. Isl., Eur. (generally), Cauc, Sib. Alas.-Nov. Scot., Nfld., S to Flor. &
E to Amur, Sakhal., Japan. Texas.

OXYPTILA GERTSCHi Kurata
Sweden. Alberta, Ontario.

OXYPTILA PRATICOLA C. L. Koch

Brit. Isl., Eur. (except the east). f Wash.

Io6 CHAPTER I

XYSTlcus LABRADORENSis Keys, {deichmatini Sor., albidus Grese)

N Fennoscand., N Russ., NW Sib. Alas.-Labr., S to Calif. & Color.;

Greenl.

Philodromidae

FHILODROMUS CAESPiTicoLis Walck. (aureolus CI. in part, canadensis Em.)

Brit. Is!., Eur. (widely), Cauc, Sib. E N U.S.A. & Can. (widely),
at least to Yenisei.

PHILODROMUS DISPAR Walclc.

Brit. Isl., Eur. (widely), Cauc. f Wash.

PHILODROMUS RUFUS Walck. (pictus Em.)

Brit. Isl., Eur. (except the north), N. Alas.-Nfld., U.S.A. (widely).

Afr., W Sib.; Japan.

THANATUS COLORADENSIS Keys, {alpinus Kulcz.)
C Eur. Rocky Mts. of U.S.A. & Can.

THANATUS FORMiciNUS CI. {canadensis Gertsch)

Brit. Isl., Eur. (generally), N. Afr., Sib. Can. & N U.S.A. (widely), Nfld.
E to Kamch.

THANATUS STRIATUS C. L. Koch {zvalteri Gertsch)
Brit. Isl., Eur. (except the east). Alas., Idah.-Utah., NE U.S.A.

TiBELLUS MARiTiMus Menge

Brit. Isl., Eur. (widely), N. Afr., Sib. Alas.-Nfld., U.S.A. (widely).
E to Kamch.

TIBELLUS OBLONGUS Walck. (parallelus Sim.)

Brit. Isl., Eur. (generally), N. Afr., Alas.-E Can., U.S.A. (widely).

Cauc, Sib. E to Kamch.

Salticidae

HASARIUS ADANSONI Aud.

Cosmopolitan in warm regions.

t Brit. Isl., Finl., France, S Eur. (hot- f U.S.A. N to Wise. & New York,
houses).

List of land and freshwater animal species 107

MENEMERUS (mARPISSA) BIVITTATUS Duf.

Almost cosmopolitan in warm regions,
f S Engl., France t Flor. -Texas.

NEON RETICULATUS Blackw.

Brit. Isl., Eur. (generally); Baical. Alas.-Oreg., Rocky Mts. S to Ariz.

PELLENES (evarcha) falcata CI. {flammata CI., hoyi Peckh.)

Brit. Isl., Eur. (widely), N. Afr., Sib. Alas.-Nfld., N U.S.A.
E to Amur & Kamch.

PHLEGRA fasciata Hahn {leopardus Peckh.)
Brit. Isl., N, C & S Eur., W Sib. E U.S.A., S to Texas, W to Kansas.

PLEXIPPUS PAYKULLI Aud.

Cosmopolitan in the tropics,
t S Eur. (accidentally). f Southern-most U.S.A.

SALTICUS SCENICUS L.

Brit. Isl., Eur. (generally), N. Afr., W f Wash.-Nfld., S to Calif.; Greenl.

Sib.

t Iceland.

siTTicus TRUNCORUM L. {pubescens F.)

Brit. Isl., Eur. (generally, usually in- f New Engl,

doors), C Asia, Amur.

Atnaurobiidae

AMAUROBIUS FEROX Walck.

Eur. (except Brit. Isl., the east & f E Can. & E U.S.A. (widely),

north).

Dictynidae

DiCTYNA ANNULIPES Blackw. {miiraria Em., mitis Thor.)
Scandinavia. Alas., Can. & N U.S.A. (widely).

DICTYNA ARUNDINACEA L. (voltita Gertsch & Iv.)

Brit. Isl., Eur. (generally), W & N Color., Ontario.

Asia E to Amur & Kamch.

I08 CHAPTER I

DICTYNA MAJOR Menge {hamifera Thor., vincens Chamb.)
Brit. Isl., N & C Eur.; Kamch. Alas.-Calif., E at least to Mont.; Greenl.

TITANOECA NIVALIS Sim. {sUvicola Ch. & Iv.)
Eur. (boreoalpine). Alas., Rocky Mts. of Can. & U.S.A.

Oecobiidae

OECOBius ANNULiPES Luc. {parietalis Hentz)
Cosmopolitan in the tropics.

t S Eur., Mediterr., Cauc. f Calif. -Flor., N to New York.

t Hawaii.

Chilopoda — Centipedes

The distribution in North America was derived mainly from Chamberlin (1925,
&c.), Bailey (1928), and Palmen (1954). For Europe, Attems (1926, 1929-40),
Brolemann (1932), VerhoefF(i937), Brade-Birks (1939), and others, were consulted.
The distribution of Centipedes (as well as of Millipedes and Woodlice) in northern
Asia, unfortunately, is quite imperfectly known. Mr. H. Lohmander, Goteborg,
has kindly looked through the lists of Chilopoda, Diplopoda, and Isopoda, Valu-
able information on the two first-named groups were obtained from Dr. O. Schu-
bart {in litt.).

CRYPTOPS HORTENSis Leach

Brit. Isl., Eur. (generally), Macarones. f Utah, New York.
Isl.

CRYPTOPS PARISI Brol.

Brit. Isl., Eur. (widely; in the north f Nfld. (greenhouse),

restricted to greenhouses).

GEOPHILUS ELECTRICUS L.

Brit. Isl., N & C Eur., N to C Scand., f Nfld.
E to Rouman.

GEOPHILUS (necrophloeophagus) longicornis LeacH

Brit. Isl., Eur. (generally), N. Afr., f Mass., Ontar., Nfld.

Hither Asia.

List of land and freshwater animal species 109

HAPLOPHILUS (STIGMATOGASTER) SUBTERRANEUS LEACH

Brit. Isl., Eur. (widely); of Mediterr. f Nfld.

origin, in the north usually restricted to
greenhouses.

LAMYCTES FULVICORNIS Mein.

Almost cosmopolitan.

Icel., Faeroes, Brit. Isl., Eur. (widely), f Alas.-NE U.S.A., Nfld.
N to C Scand., N. Afr., Macarones.
Isl., Cauc.

t E Afr., Austral.

LITHOBIUS FORFICATUS L.

Brit. Isl., Eur. (generally), Cauc, N. f Canada & U.S.A., W to Rocky Mts.,

Afr. Nfld.i

t S. Amer., St. Helena, Austral.

LITHOBIUS MELANOPS Newp.

Icel., Faeroes, Brit. Isl., Eur. (widely), f Nfld.
N Scand.

LITHOBIUS (monotarsobius) microps Mein. {dubosqiii Brol.)

Brit. Isl., N & C Eur. (in the north f Nfld,

synanthropic).

pachymerium ferrugineum C. L. Koch
Almost cosmopolitan.

Faeroes, Eur. (except Brit. Isl. & the f Alas.; E U.S.A., S to Flor., W to
north), N. Afr., Cauc, W & C Asia. Minnes.

Mexico,
f Pacific islands.

schendyla nemorensis C. L. Koch
Brit. Isl., Eur. (generally), N. Afr. t E Can., NE U.S.A., Utah; Nfld.

scolioplanes (linotaenia) acuminatus Leach (chtonophilus Wood)^

Brit. Isl., W & S Eur., N. Afr., Cauc. (f) Alas., & N. W. Terr.-Queb., N U.S.A.
(t) Bering Isl.

^ Vide map by Chamberlin (1922, fig. 3, p. 451; by mistake exchanged with fig. 4, p. 453).
^ According to Lohmander (in litt.), acuminatus Leach and crassipes C. L. Koch have been
largely confused and the distribution here given may need revision.

no CHAPTER I

THEATOPS ERYTHROCEPHALUS C. L. Koch

SEur.^ tW U.S.A.

Diplopoda — Millipedes

Records for North America were mainly obtained from Chamberlin (1921) and
Palmen (1952), for Europe from Attems (1929-40), Schubart (1934), Blower
(1952), and other papers mentioned among the references (p. 126 a.f.).

BLANIULUS GUTTULATUS BOSC.

Brit. Isl., N & C Eur., E to S Russ. f NE U.S.A., E Can., Nov. Scot., Nfld.

(synanthropic in the north).

f St. Helena.

BRACHYDESMUS SUPERUS LatZ.

Faeroes, Brit. Isl., Eur. (widely), E to f Ohio, Nov. Scot., Nfld.
SW Russ., N. Afr.

BRACHYIULUS LiTTORALis Vcrh. {pusUliis Chamb.)

Brit. Isl., N & C Eur. E to Polon., S f CaHf.; E U.S.A., S to N. Carol., W

to Alban. to Mississ.; Nov. Scot., Nfld.

CHONEIULUS PALMATUS Nemcc

Brit. Isl., N & C Eur. E to Russ. f Nov. Scot., Nfld.

CYLINDROIULUS BRITANNICUS Verh.

Almost cosmopolitan.

Faeroes, Brit. Isl., Eur. (widely), E to f Nfld.
S Russ.

t S. Amer., S. Afr., S Asia.

CYLINDROIULUS FRisius Verh. (oiveni Bollm., luscus Mein.)
Almost cosmopolitan.

Faeroes, Brit. Isl., Eur. (S Scand.-N f Calif.; C & E U.S.A., Nov. Scot.,
France), E to S Russ. Nfld.

t C & S Amer., Juan Fern., S. Afr., St. Paul (S Ind. Ocean).

CYLINDROIl'LUS SILVARUM Mein

Brit. Isl., N & C Eur., E to NW Polon. f Nfld.

List of land and freshwater animal species m

CYLINDROIULUS TEUTONicus Poc. (londtnensis C. L. Koch)

Brit. Isl., N & C Eur., E to W Russ. f U.S.A. (widely), E Can., Nov. Scot.,

Nfld.

CYLINDROIULUS TRUNCORUM Silv.

t Brit. Isl.i, Eur. (widely; mainly in f Wash., Color, (greenhouses), Nfld.

greenhouses), E to S Russ.

N. Afr.

NOPOIULUS VENUSTUS Mein. (pulchellus & minutus Chamb.)

Brit. Isl., Eur. (widely), E to S Russ., f E U.S.A., Nfld.
Asia Minor.

t C & S Amer., Juan Fern., St. Helena.

OPHIODESMUS ALBONANUS LatZ.

Brit. Isl., W Eur., Sweden. f Nfld.

OPHYIULUS FALLAX Mein. (pilosus auctt., longabo Chamb.)

Brit. Isl., N & C Eur. E to Polon. f NE U.S.A. W to Ohio, E Can., Nov.

Scot., Nfld.
t Austral.

ORTHOMORPHA GRACILIS C. L. Koch

Cosmopolitan.

f Brit. Isl., Eur. (widely; mainly in f U.S.A. (widely, greenhouses), Nfld.

greenhouses), E to Russ. (greenhouse); Bermudas.

t S. Afr., S & E Asia, Austral., Pacific S. Amer. (probably indigenous).

Islands.

POLYDESMUS DENTICULATUS C. L. Koch

Brit. Isl., N & C Eur. E to Ural. f Nfld.

POLYDESMUS INCONSTANS Latz. {coriaceus auctt., nee Porat, testi BoUm.)
Brit. Isl., N & C Eur., E to Volga. f Indiana, Ohio, lUin.; Nfld.

POLYDESMUS RACOVITZAI Brol.

Pyrenees Orientales. f Wash.

PROTEROIULUS FUSCUS a. St.

Icel., Brit. Isl., N & C Eur., E to W f E U.S.A., Nov. Scot., Nfld.

Russ.

^ Not published from the British Isles but recorded by Palmen {in litt.) from Bristol.

112 CHAPTER I

Isopoda Terrestria — Woodlice

Van Name's excellent monograph (1936, 1940) of the American fauna was supp-
lemented by Hatch (1947) and Palmen (1951). European records were obtained
from Wachtler (1937), Collinge (1943), Palmen (1946), &c.

ANDRONISCUS DENTIGER Verh.

Brit. Isl., S, C & W Eur. t Ontar., Nfld. (greenhouses).

ARMADILLIDIUM NASATUM B.-Lund

Brit. Isl., Eur. (except the north), E to f Brit. Col., E Can., U.S.A. (widely; in

Polon. (in Scand. usually in green- the north in greenhouses).

houses).

ARMADILLIDIUM VULGARE Latr.

Almost cosmopolitan.
Brit. Isl., Eur. (widely), N. Afr., f Brit. Col., U.S.A. (generally), Bermu-

W Asia. das.

t C & S Amer., Austral., Pacific Isl.

CYLISTICUS CONVEXUS DeG.

Brit. Isl., Eur. (generally). Hither Asia, t Brit. Col.-New Brunsw., Nfld.,

U.S.A. (widely), S to N. Carol. & Color,
t C & S Amer.

HAPLOPHTHALMUS DANICUS B.-Lund

Brit. Isl., Eur. (except the north); f E U.S.A., W to Utah, E Can., Nfld.

originally Mediterranean (regularly in

greenhouses).

HYLONiscus RiPARius C. L. Koch {vtvtdus C. L. Koch)

Brit. Isl., Eur. (except the north), E f Nfld.

to Polon.

ONISCUS ASELLUS L.

Faeroes, Brit. Isl., Eur. (generally). f Brit. Col.-Oreg.; E U.S.A., E Can.,

Nov. Scot., Nfld.
t W. Indies.

PHILOSCIA MUSCORUM Scop.

Brit. Isl., Eur. (except the east), N. fWash.

Afr.

f C Amer.

List of land and freshwater animal species 113

PLATYARTHRUS HOFFMANSEGGI Brandt

Brit. Isl., S, W & C Eur., SE to Dal- f Connect, (myrmecophilous).
matia.

PORCELLio DILATATUS Brandt

Icel., Brit. Isl., S & W Eur. t Brit. Col.-Ariz.; Nfld.

t Australia.

PORCELLIO LAEVIS Latr.

Brit. Isl., Eur. (generally), N. Afr., W f U.S.A. (widely) W to Calif., E Can.
Asia.

t C & S Amer., Pacific isl.

PORCELLIO (PROPORCELLIO) QUADRISERIATUS Verh.

S Eur., Hither Asia. f Texas (greenhouse).

PORCELLIO SCABER Latr.

Cosmopolitan.

Icel., Faeroes, Brit. Isl., Eur. (generally), f Alas.-Labr., Nfld., U.S.A. (generally),
t C & S Amer., S. Afr., New Zeal., Pacific isl., Japan, Kamch.

PORCELLIO SPiNicoRNis Say {pictus Brandt)
Almost cosmopolitan.
Brit. Isl., N & C Eur. E to Russ. f E Can., E U.S.A. W to Mich.

PORCELLiONiDES (metoponorthus) pruinosus Brandt
Cosmopolitan.

Brit. Isl., Eur. (widely), E to C Russ., f Brit. Col.-E Can., U.S.A. (generally).
N. Afr.

t C & S Amer., Madag., S Asia, Pacif. isl., Japan.

trachelipus (tracheoniscus) rathkei Brandt
Almost cosmopolitan.

Icel., Brit. Isl., Eur. (except the east), f Brit. Col., NW U.S.A.; E U.S.A., E
Cauc. Can., Nov. Scot., Pr. Edw. Isl., Nfld.

t C & S Amer.

trichoniscoides sarsi Pat.
Brit. Isl., W & C Eur., N to Norway. f Nfld.

8 — 565597 Lindroth

114

CHAPTER I

TRICHONISCUS PROVISORIUS RaCOV.

Brit. Isl., C Eur., N Eur. (in green- f Nfld.

houses).

TRICHONISCUS PUSILLUS Brandt, /. parthenog. (? demivirgo Blake)

Icel., Faeroes, Brit. Isl., N & C Eur. f Wash.; NE U.S.A., E Can., Nov.

Scot., Nfld.

TRICHONISCUS PYGMAEUS G. O. Sars

Brit. Isl., N & C Eur. (in the north in f New York, lUin., Nfld.
greenhouses).

TRICHONISCUS (CORDIONISCUS) STEBBINGI Pat.

Brit. Isl., W Eur. to S Swed. (Loh- f Oreg., Mass. (in greenhouses),

mander, in litt.).

TYLOS LATREILLI Ad. & Sav.

Coasts of S Eur., N to C France, Asia (f) Florida; Bermudas.
Minor, N. Afr., Macaron. Isl. C Amer., W. Indies.

Mollusca, Gastropoda — Land- and freshwater Molluscs

The main sources of information on Mollusca were Ehrmann (1937), Taylor
(1902-21), Ellis (1926), and Licharev & Rammelmayer (1952), for Eurasia, Pilsbry
(1939-48) and La Rocque (1953), for North America. Other references may be
found in the bibliography (p. 126 a.f.). The nomenclature follows Pilsbry (I.e.).
Valuable information was obtained from Mr. H. W. Walden, Stockholm.

Terrestrial or amphibious species closely connected with the seashore are not
considered.

CEPAEA (helix) hortensis O. F. Miill. (fig. 29)

Icel., Faeroes, Brit. Isl., Eur. (except the (t)Mass.-Queb., N. S., Nfld., ?Labr.;
southern peninsulae), E to Leningrad ? Greenl.

& Vitebsk.

t New Zeal.

cepaea (helix) nemoralis L.

Brit. Isl., Eur. (widely), E to Balticum, f NE U.S.A. S to Tenn., E Can., An-
E. Pruss., Bohcm., & Bosnia. ticosti.

List of land and freshwater animal species 115

HELICELLA (THEBA, EUPARYPHA) PISANA O. F. Miill.

Brit. IsL, W & S Eur. N to N France, f Calif.
E to Bulg.; N. Afr., Hither Asia.

t S. Afr.

HELICOGONA (aRIANTA) ARBUSTORUM L.

Icel., Faeroes, Brit. IsL, Eur. (except the f Nfld. (old record).

southern peninsulae), E to Balticum &

Kiew.

HELIX aspersa O. F. Mull.

Brit. IsL, S & W Eur. N to HolL, f Calif, (generally); Louis., S. Carol,

isolated on Crimea; N. Afr., Asia Minor.

t C & S Amer., S Afr., Austral.

Jacostidae

COCHLICELLA VENTROSA Fer.

S Eur., N. Afr. f Calif.; S. Carol.

t Bermudas.

HYGROMIA (tRICHIA, TROCHULUS) HISPIDA L.

Brit. IsL, Eur. (except the extreme f NE U.S.A., E Can., Nov. Scot., Pr. Edw.

south), E to Volga; Cauc, N. Afr. IsL

HYGROMIA (trichia, trochulus) striolata Pfeiff.

Brit. IsL, W & C Eur. N to S Swed., f NE U.S.A., E Can., Labr., Nov. Scot.,
E to Czechosl. . Nfld.

jacosta (helicella, trochoidea, xeroclivia) elegans Gmel.

Mediterr.: S Eur., N. Afr. f S. Carol,

t Brit. IsL

MONACHA (theba) cantiana Mont.

Brit. IsL, C & W Eur., S Russ., Hither f E Can. (Ont., Queb.).
Asia.

Achatinidae

cecilioides acicula O. F. Mull.

Brit. IsL, Eur. (widely) E to Ucraina & f Florida, Penns.
Crimea; Cauc, W. Turcest.

t Bermudas, New Zeal.

Il6 CHAPTER I

LAMELLAXIS (OPEAS) CLAVULINUS Pot. & Mich.

t Brit. Isl. (greenhouses). t Penns.

Islands of Ind. Ocean,
t Hawaii, Japan.

LAMELLAXIS (OPEAS) MAURITIANUS Pfeiff.

t Irel., Holl., Germ, (greenhouses). f E U.S.A. (greenhouses).

Mauritius.

OPEAS PUMILUM Pfeiff.
t Brit. Isl., W Eur. Florida. f E U.S.A.

(greenhouses). C & S Am. (greenhouses).

RUMINA (STENOGYRA, BULIMUS) DECOLLATA L.

S Eur., N. Afr., Hither Asia. f Texas-N. Carol,

t Brit. Isl. (greenhouses). t W. Indies.

SUBULINA OCTONA Brug.

f Brit. Isl., W. Eur. (greenhouses). S & C Amer. N to Florida.

t Afr., S Asia, &c.

Testacellidae

TESTACELLA EUBOPAEA Roissy {haliotidea Drap.)

Brit. Isl., S & W Eur. N to Belg., E f Calif.; NE U.S.A., Nov. Scot,

to Dalmat.; N. Afr. (greenhouses).

Zonitidae

EUCONULUS FULVUS O. F. Miill. {trochiformis Mont.)
Icel., Brit. Isl., Eur. (generally), N. Alas.-Labr., S to Middle Atl. States;

Afr., N Asia E to Kamch. Nfld., Greenl.

t S Asia, Austral.

HAWAIIA MINUSCULA Binn.
t Brit. Isl., Holl. (greenhouses). Alas.-E Can., Nfld.; U.S.A. (generally)

E Sib. (S parts of Marit. Prov., S to C Amer.; Bermudas,

probably indigenous).

t Corea, Japan, Formosa, Pacific islands.

oxYCHiLus (hyalinia) ai.ll\rius Mill.
Icel., Faeroes, Brit. Isl., W & C Eur. fBrit. Col.-Calif.; Ont.-N. Y.
E to Leningrad; N. Afr. f Greenl.

t S. Afr., Austral.

List of land and freshwater animal species 117

oxYCHiLus (hyalinia) cellarius O. F. Mull.

Brit. Isl., W & C Eur. E to Leningrad; f Oreg., Calif.; E U.S.A., E Can.,
N. Afr., Cauc, Hither Asia. Nov. Scot.

t S. Amer., S. Afr., Austral.

OXYCHILUS (hyalinia) draparnaldi Beck {lucidus Drap.)

Brit. Isl., Eur. (not indig. N of Holl.) f Wash.-Calif.; E U.S.A., E Can., Nfld.
E to C Russia; N. Afr., Hither Asia.

OXYCHILUS (hyalinia) helveticus Blum
Brit. Isl., W & C Eur. N to Belg. f Calif.

RETiNELLA (PERPOLiTA, HYALINIA) HAMMONis Stroem {radiatula Aid.)

/. typ. sbsp. electrina Gould^

Icel., Faeroes, Brit. Isl., Eur. (except Alas.-Labr., S to New Mex. & Ariz.,

the south), N Asia E to Kamch., Nov. Scot., Nfld.
Japan.

ZONITOIDES ARBOREUS Say

t Brit. Isl., C & W Eur. E to C Russ. Brit. Col.-Labr., Nfld., entire U.S.A.

C Amer., W. Ind.

Kamch. (probably indig.).

t Japan, S. Afr., Austral.

ZONITOIDES NITIDUS O. F. Miill.

Brit. Isl., Eur. (generally), N. Afr., Alas.-E Can., Nfld., S to Maryl. &

Cauc, C & N Asia, Japan. Utah.

t Austral.

Limacidae

DEROCERAS CARUANAE Poll.

S & W Eur., N. Afr. t Calif.

(Original patria?)

DEROCERAS (agriolimax) laeve O. F. Mull., SBHsu lato {campcstrc Binn.)

Icel., Faeroes, Brit. Isl., Eur. E to C Alas.-Nfld.-S Baffin L.; S to S U.S.A.

Russia, Hither Asia (forms from N
Asia possibly conspecific).

t C & S. Amer., S. Afr., Austral.

^ Often regarded as distinct species {vide Pilsbry, 1946, p. 258).

Il8 CHAPTER I

DEROCERAS (agriolimax) reticulatum O. F. MviU. {agrestis auctt. nee L.)

Icel., Faeroes, Brit. Isl., Eur. E to C f Brit. Col., Calif.; E Can., Labr.,

Russia. Nfld.; U.S.A. (disjunct but widely),

f S. Amer., S. Afr., E Asia, Austral.

LI MAX FLAVUS L.

Brit. Isl., Eur. E to S Russia (origi- f U.S.A. (generally).

nal patria).

N. Afr., Hither Asia, Cauc.

t S. Amer., S. Afr., Austral., E Asia.

UMAX (lehmannia) marginatus O. F. Miill. (arborum Bourch. & Chant.)

Icel., Faeroes, Brit. Isl., Eur. E to W f U.S.A. (disjunct)i, Nfid.
Russia and Greece, N. Afr.

t S. Amer., Austral., Atl. & Pacif. islands.

LIMAX MAXIMUS L.

Brit. Isl., Eur. E to S Russia, Cauc; t Brit. Col., Calif.; E Can., Nfld.; U.S.A.

N. Afr., Asia Minor. (widely but disjunct),

t S. Amer., S. Afr., Austral., Hawaii.

MILAX GAGATES Drap.

Brit. Isl., S & W Eur. N to Belg.; f Wash.-Calif.; E U.S.A. (usually in

N. Afr. greenhouses).

t S. Afr., Austral., Juan Fern.

Endodontidae

DISCUS (goniodiscus) rotundatus O. F. Miill.

Brit. Isl., W & C Eur. E to S Russ.; f Mass., Nfld.

N. Afr.

HELicoDiscus PARALLELUS Say {litieatus Say)

t Brit. Isl., Holl. Can. W to Manit., Nova Scot., Nfld.,

E & C U.S.A.

Arionidae

arion ater L. coll. {empiricorum Fer.)

Icel., Faeroes, Brit. Isl., C, W & N Eur. f Brit. Col.-Oreg.; E Can., Nfld.; NE
E to Latvia & Hungaria. U.S.A.
t New Zeal.

^ The mainland form of North America, according to information from H. W. Wald^n,
Stockholm, belongs to the European-Mediterranean L. poirieri Mab.

List of land and freshwater animal species 119

ARION CIRCUMSCRIPTUS Johnst. (incl. fasciatus Nilss.)

Icel., Faeroes, Brit. Isl., W, C & N f Brit. Col.-Calif.; NE U.S.A. W to

Eur. E to C Russia. Wise; E Can., Nfld.

ARION HORTENSIS Per.

Icel., Faeroes, Brit. Isl., S, W & C f Wash.-Calif.; E U.S.A., E Can.,

Eur., E to W & NW Russia. Nov. Scot., Nfld.

ARION iNTERMEDius Norm.

Icel., Faeroes, Brit. Isl., W, C & N f Calif.

Eur. E to C Russia.

t S. Afr., New Zeal.

ARION SUBFUSCUS Drap.
/. typ. sbsp. sibiricus Simr.

Icel., Faeroes, Brit. Isl., f NE U.S.A., E Can., Altai, E Sib.

C & N Eur. E to Ural. Nov. Scot., Nfld.

t New Zeal.

Pupillidae

COLUMELLA (sphyradium) edentula Drap. {simplex Gould)

Icel., Faeroes, Brit. Isl., Eur. (gener- Alas.-Labr., Nfld., S to Iowa & Penns.

ally), N Asia E to Kamch., Sakhal.,
Kuril., Japan.

PUPILLA MUSCORUM L.

Icel., Brit. IsL, Eur. (generally), N. Alas.-E Can., Nfld., S to New Mex.

Afr., W, C & N Asia E to Amur &

Sakhal.

VERTIGO alpestris Alder

/. typ. sbsp. oughtoni Pilsb.

Brit. Isl., W, N. & C f Nfld. Huds. Bay (W Ontario),

Eur., Sib. E to Amur. Baffin L.

VERTIGO modesta Say

(incl. arctica Wallenb., borealis Morel., kraussena Reinh., &c.)
Icel., Eur. (boreomontane), Siberia, Alas.-Labr., Nfld., NE U.S.A., Baffin

Commander Isl. L., Greenl.

vertigo pygmaea Drap.

Brit. Isl., Eur. (generally), Cauc, N. (f) E Can., Nov. Scot., NE U.S.A. S to

Afr., N & C Asia. Virg., W to Ohio.

I20 CHAPTER I

Valloniidae

VALLONIA COST ATA O. F. Miill.

Brit. Isl., Eur. (generally), N. Afr., Alberta-Queb., NE U.S.A. S to Virg.

Sib. E to Amur.

t Austral.

VALLONIA EXCENTRICA Sterki

Brit. Isl., Eur. (excl. the north). (f) E Can., Nov. Scot., Nfld.; NE

U.S.A., locally elsewhere (for inst.
Oreg. & Calif.),
t Mexico, S. Afr.

VALLONIA PULCHELLA O. F. Mull.

Brit. Isl., Eur. (generally), N. Afr., N. Amer. E of Rocky Mts., N to Nov.

Sib. E to Amur. Scot., S to Kentucky.

t Calif., Texas,
f Makarones., Austral., &c.

zooGENETES (acanthinula) harpa Say

Eur. (boreoalpine), Cauc, N Asia E to Alas.-Labr., Nfld., S to Color. & New
Amur, Kamch., Sakhal. & Bering Isl. Hampsh.

Cionellidae

cionella (cochlicopa) lubrica O. F. Mull.

Icel., Faeroes, Brit. Isl., Eur. (gener- Alas.-Labr., Nfld., S to Ariz. & north,

ally), N. Afr., N & C Asia E to Kamch. Mexico.
& Japan.

t New Zeal.

Carychiidae

CARYCHIUM MINIMUM O. F. Miill.

Brit. Isl., NW & C Eur., entire Russia, f Massach. (greenhouse).
Cauc, Sib.

Lymnaeidae

(Vide Hubendick, 1951)

lymnaea (radix) auricularia L. (s. str.)

Brit. Isl., Eur., N. Afr., N & C Asia S Alaska (probably indigenous).

E to Amur, Kamch., Japan. f U.S.A.

List of land and freshwater animal species I2i

LYMNAEA (sTAGNICOLA) CATASCOPIUM Say

(t) Scotl. (Kevan, 1943). Great Slave Lake-Nov. Scot.; NE

U.S.A.

LYMNAEA (pseudosuccinea) COLUMELLA Say {peregrina Cless.)

t Brit. Isl., W & C Eur. E U.S.A., SE Can.

(mainly in hothouses). f Oreg.-Calif.

t C & S Amer.; S. Afr.

LYMNAEA (gALBA) CUBENSIS Pfeiff.

t Irel., Germ, (hothouses). SE U.S.A., C & S Amer.

LYMNAEA (STAGNICOLA) PALUSTRIS O. F. Miill.

Brit. Isl., Eur., N. Afr., C & N Asia Alas.-Queb., Nfld., S to Arizona.

E to Amur & Kamch.

LYMNAEA (rADIX) PEREGRA O. F. Miill.

Icel., Faeroes, Brit. Isl., Eur. (gener- Nfld.

ally), N. Afr., Asia E to Baikal.

LYMNAEA STAGNALIS L.

Brit. Isl., Eur., N. Afr., Asia E to Alas.-Nov. Scot., S to Arizona.

Amur & Kamch.

LYMNAEA (fOSSARIA) TRUNCATULA O. F. Miill.

Icel., Faeroes, Brit. Isl., Eur., Afr. Aleut. Isl., S Alas., C Canada.

(widely), C & N Asia E to Amur &

Kamch.

Physidae

aplexa (aplecta) HYPNORUM I..

Brit. Isl., Eur. (generally), W & N Alas.-E Can., S to Color.

Asia E to Kamch.

PHYSA ANCILLARIA Say

t Germany (hothouses). NE U.S.A., E Can. W to Manitoba.

PHYSA GYRINA Say

t Brit. Isl. (Cardiff). Alas.-E Can., S to Alabama, Nfld.

122 CHAPTER I

PHYSA HETEROSTROPHA Say

t Genp., Czechosl., Holl., Denm. Manit.-Nfld., U.S.A. E of Mississ.

(mainly hothouses).

Planorbidae

GYRAULUS (aRMIGER) CRISTA L.

Brit. Isl., Eur. E to C Russia, N. Afr., Alas.-Ont., NE U.S.A.
WSib.

GYRAULUS GREDLERI Bielz (incl. borealis Loven, syn. arcticus Beck)

Eur. (boreoalpine), N & C Asia E to Alas.-Labr.; Greenl.

Kamch. (Glacial subfossils in the Brit.

Isl.)

GYRAULUS (torquis) PARVUS Say^
t Denmark (aquarium). Alas.-E Can., S to Florida, Nfld.

helisoma duryi Weth. (incl. eudisciis Pilsb.)
t Engl, (aquarium). Florida.

HELISOMA (PIEROSOMA) TENUIS Phil.

t Denmark (aquarium). Calif., Mexico.

HELISOMA (pierosoma) trivolve Say
f Moscow (aquarium). Alas.-E Can. & Maine, S to Missouri.

Valvatidae

valvata piscinalis O. F. Miill.

Brit. Isl., Eur. (generally), Cauc, f NE U.S.A., E Can., Nov. Scot.

Hither Asia, W Sib.

Amnicolidae

BULIMUS (biTHYNIA) TENTACULATUS L.

Brit. Isl., Eur. (generally), N. Afr., (f) NE U.S.A., E Can., W to Wise*

Cauc, W Sib.

^ Sometimes regarded as a sbsp. of the Palaearctic G. laevis Aid.

'^ Baker (1928, p. 81) described a new and possibly indigenous sbsp. of Bulimus tenta-
ciilatus from N. America but its state seems doubtful (La Rocque, 1953).

List of land and freshwater animal species 123

Oligochaeta, Lumbricidae — Earthworms

Great interest in the zoogeography of OHgochaeta was shown by the late W.
Michaelsen {vide 1903) but only the terrestrial or amphibious members of the
Lumbricid family, the true earthworms, seem suited to our purpose. Valuable in-
formation has also been obtained from Smith (1917), Ude (1929), and Cernosvitov
& Evans (1947), as well as from a few additional papers listed in the bibliography
(p. 126 a.f.).

ALLOLOBOPHORA (hELODRILUS) CALIGINOSA Sav.

Cosmopolitan.!

Icel., Brit. Is!., Eur. (generally), N. f U.S.A. (generally), E. Can.

Afr., Hither Asia.

f S. Amer., S. Afr., S & E Asia, Austral., Hawaii.

ALLOLOBOPHORA (HELODRILUS) CHLOROTICA Sav.

Brit. Isl., Eur. (generally), Hither Asia, f Brit. Col., W & E U.S.A., E Can.,

Bermud.; Greenl.
t C & S Amer.

ALLOLOBOPHORA (HELODRILUS) LONGA Ude

Brit. Isl., Eur. E to S Russ. f E Can., Maine, Indiana.

BIMASTUS (HELODRILUS) BEDDARDI Mich.

t Ireland. U.S.A. (widely).

f Tibet, Hawaii.

BIMASTUS (HELODRILUS) TENUIS Eis. {constrlctus Rosa)

Icel., Brit. Isl., Eur. (generally), Sib. Alaska (probably indigenous).

(t) Brit. Col.-Calif.; E U.S.A.
t C & S Amer., India, N. Zeal., Hawaii.

DENDROBAENA (hELODRILUS) OCTAEDRA Sav.

Icel., Brit. Isl., Eur. (generally), N. f Color., Nfld., Greenl.

Zeml., Cauc, Sib. E at least to Baical. f Mexico.

DENDROBAENA (hELODRILUS) SUBRUBICUNDA Eis.

Icel., Brit. Isl., Eur. (generally), N. f Calif., Color., Illin.; E Can., Nfld.

Afr., S Sib.

f S. Amer., India, Austral.

^ "The commonest species of earthworm in the world "(Cernosvitov & Evans, 1947).

124 CHAPTER I

EISENIA CAROLINENSIS Mich.

t Germ. (Hamburg; Smith, 191 7) • N. Carolina.

EISENIA FOETIDA Sav.

Almost cosmopolitan.
Icel., Brit. Isl., Eur. (generally), t Brit. Col., U.S.A. (generally), E Can.,

Hither Asia, Sib., Japan. Bermud.

t C & S Amer., S. Afr., S Asia, Austral., Hawaii.

EISENIA ROSEA Sav.

Almost cosmopolitan.
Icel., Brit. Isl., Eur. (generally), N. f Calif., Ariz., E U.S.A., E Can.

Afr., W & N Asia.

t C & S Amer., S. Afr., Austral.

EISENIA VENETA Rosa ("var. hortensis Mich.")
Brit. Isl., Eur. (except the north), f Calif.

Hither Asia (/. typ.).

t S. Amer., S. Afr.

EISENIELLA TETRAEDRA Sav.

Almost cosmopolitan.
Icel., Brit. Isl., Eur. (generally), f Wash. -Calif.; C & NE U.S.A., E Can,

Hither Asia, Cauc.

t S. Amer., S. Afr., S Asia, Austral.

LUMBRICUS CASTANEUS SaV.

Faeroes, Brit. Isl., Eur. (widely), Sib. t NE U.S.A., E Can.

LUMBRICUS FESTIVUS Sav.

Brit. Isl., Eur. N to S Sweden. f E Can.

LUMBRICUS RUBELLUS Hffm.

Icel., Faeroes, Brit. Isl., Eur. (generally), f Wash.-Calif.; Mich., E Can., Nfld.
Cauc, Sib. E at least to R. Lena.

f S Asia, Austral.

LUMBRICUS TERRESTRis L. {herculeus Rosa)

Icel., Brit. Isl., Eur. (generally). f W & E U.S.A., E Can., Nfld.

t Mexico, Falkland Isl., Macaron. Isl.

OCTOLASIUM LACTEUM Oerl.

Brit. Isl., Eur. (except the north) E f W & C U.S.A.

to C Russia, N. Afr., Hither Asia.

t C & S Amer., S & E Asia, Austral.

List of land and freshwater animal species

125

TABLE I . Summary of species common to Europe and North America among animal

groups treated above. ^

Mammalia

Aves

Pisces

Coleoptera (in part) .

Carabidae

Elateridae

Coccinellidae ....

Scarabaeidae ....

Ceranibycidae . . .

Chrysomelidae . . .

Rhynchophora (in part)
Lepidoptera (in part)

Diurna

Sphinges

Bombycimorpha . .

Noctuae

Geometrae

Isolated families . . .
Diptera (in part) . .

Culicidae

Tabanidae

Hymenoptera (in part)

Bombinae

Crabronidae ....

Vespinae

Formicidae

Odonata

Orthoptera Saltatoria

Araneae

Chilopoda

Diplopoda

Isopoda Terrestria .

Mollusca

Lumbricidae ....

DiflF.

sbsp.

17
63

3

27

17
2

5

41

14

2

6

15

Iden-
tical

3

44

241

74

10

8

25
12
27
85
94
15
I

7
38
29

4
24
20

4

35
2

5

7

141

13
17
22
70
17

Introduced species

Total Eur. to

N. Am.

20
107

3
267

91
12
12
25
14
28

85

135
29

3
13
46

39

5

25
21

4
50
4
8
6
32
8

7
146

13
17
22

71
17

162-170

37-38

3-5

5

23

7

19-21

68-71

15-17
2
I
3
2-4
4
3

4-7

2-3
I

1-3

2
30-46
12-13

15
21-22
32-36

15

N. Am.
to Eur.

3-4

1-2
1-4

1-2
0-2

13-14

2

In

both
contin.

Total
introd.

3

o

o

174-183

41-42

3-5

5

24

8

20-22

73-77

16-21

4-5
2-6

4
3
I
I
o
21-24
o

2-3
I

18-20

o

4
43-59
12-13

17
21-22

48-53
17

Total

15

65-68
45-46
25-42

38

96

57

71-79

86-91

12-16

7

33
31-38

4-13

10

60

4

5

42-48

25-38

17
56-63

57
29-40
93-100

100

95-100

66-73

94

Total 177

733

908 309-344

= 33-38%

27-34
= 3-4 %

39

= 40

375-417

41-46

^ The following five species stated or believed to occur in North America in part as in-
digenous, in part introduced from Europe, are counted twice on calculating the percentage
of introduced species and, for the subspecifically distinct Coccinella and Vertigo, also in the
two first columns: Amara hmicolbs (Col. Car.), Coccinella ii-punctata (Col. Cocc), Vertigo
alpestris (Moll.), Lymnaea auricularia (Nioll.), Bimastus tenuis (Lumbr.).

Bibliography of Chapter I.

Ander, K., 1950. Zur Verbreitung und Phanologie der boreoalpinen Odonaten der
Westpalaarktis. — Opusc, Ent. 15. Lund, p. 53-71.

1951- Vastpalearktis' och Nordamerikas orthopterfauna, en jamforelse. —

Ent. Medd. 26. Copenhagen, p. 155-156.

Anderson, R. M., 1946. Catalogue of Canadian recent Mammals. — Bull. Nation.

Mus. Can. 102 (Biol. Ser. 31). Ottawa, p. 1-238.
Attems, C, 1926. Chilopoda. — Handb. d. Zool. (W. Kiikenthal). IV: i. Berlin &

Leipzig, p. 239-402.

1929-40. Myriapoda. 1-3. — Das Tierreich. 52. 54, 68-70, Berlin & Leipzig.

Backlund, H. O., 1949. Oligochaeta. i. Lumbricidae. — Zool. of Icel. 2. Copen-
hagen, p. 1-15.

Bailey, J. W., 1928. The Chilopoda of New York State with notes on the Diplopoda.

—Bull. N.Y. State Mus. 276. Albany, N.Y., p. 1-50.
Baker, F. C, 1928. The fresh water Mollusca of Wisconsin. L — Bull. Wise. Geol.

& Nat. Hist. Survey. 70. Madison, Wise, pp. I-XX, 1-507.

1945. The MoUuscan family Planorbidae. — Univ. Illin. Press. Urbana, 111.,

pp. I-XXXVI, 1-530.

Bartenef, a., 1935. iJber die Gattung Libellula und besonders iiber ihre palaark-
tischen Arten. — Zeitschr. wissensch. Zool. (Arch. f. Naturgesch.) (N.S.) 4. Leip-
zig, p. 274-290.

Berg, L. S., 1933. Ubersicht der Verbreitung der Siisswasserfische Europas. ^
Zoogeogr. i. Jena, p. 107-208.

1948-49. Ryby presnych vod SSSR i sopredelnych stran (Freshwater fishes

of SSSR and adjacent countries). 1-3. — Opredel. Fauna SSSR. 27. 29. 30.
Moscow & Leningrad, p. 1-1382.

Berland, L., 1928. Hymenopteres vespiformes. II. — Faune de France. 19. Paris,
p. 1-208.

1932. Les Arachnides. - Encycl. Ent. 16. Paris, p. 1-485.

Blake, C. H., 1931. Distribution of New England Wood Lice. — Occ. Papers

Boston Soc. Nat. Hist. 5. Boston, p. 349-355.
Blatchley, W. S., 1920. Orthoptera of Northeastern America. — Indianapolis,

p. 1-784.
Blatchley, W. S. & Leng, C. W., 1916. Rhynchophora or Weevils of North Eastern

America. — Indianapolis, p. 1-682.
Blauvelt, Helen H., 1936. The comparative morphology of the secondary sexual

organs of Linyphia and some related genera, including a revision of the group. —

Festschr. f. Embr. Strand. 2. Riga, p. 81-171.
Blower, G., 1952. British Millipedes with special reference to Yorkshire species.

— The Naturalist (Quarterly Journ. Nat. Hist. N. Engl.). London, p. 145-157.
BoETTGER, C. R., 1932. Die Besiedelung neu angelegter Warmhauser durch Tiere.

— Zeitschr. Morph. & Okol. d. Tiere. 24. Berlin, p. 394-407.

Bowman, K., 195 i. An annotated list of the Lepidoptcra of Alberta. — Can. Journ.
Zool. 29. Ottawa, p. 121-165.

Bibliography 127

Brade-Birks, S. G., 1939. Notes on Myriapoda XXXVI. Sources for description

and illustration of the British fauna. — Journ. S.-E. Agric. Coll., Wye. 44. London,

p. 156-179.
Brolemann, H. W., 1930. Elements d'une faune des Myriapodes de France. — Faune

de France. 25. Paris, p. 1-405.
Brown, W. J., 1935. American species of Ludius; the cruciatus and edwardsi groups

(Coleop.). — Can. Ent. 67. Orillia, Ont., pp. 1-8, 125-135, 213-221.
1936. Notes on some species of Elateridae (Coleoptera). — Ibidem. 68. p.

246-252.

1940. Notes on the American distribution of some species of Coleoptera

common to the European and North American continents. — Ibidem. 72. p. 65-78.

1943- The American species of Phytodecta Kby. (Coleoptera, Chrysomelidae).

— Ibidem. 74. p. 99-105.

1946. Some new Chrysomelidae with notes on other species (Coleoptera). —

Ibidem. 78. p. 47-54.

1950. The extralimital distribution of some species of coleoptera. — Ibidem.

82. p. 197-205.

1951- The American species of Phratora Chev. (Coleoptera: Chrysomelidae).

— Ibidem. 83. p. 121-130.

Bryk, F., 1942. Zur Kenntnis der Grossschmetterlinge der Kurilen. — Deutsch.
Ent. Zeitschr. "Iris". 56. Dresden, p. 3-90.

Buchanan, L. L., 1927. Synonymical notes on several Otiorhynchid Weevils. — •
Can. Ent. 59. Orillia, Ont., p. 183-184.

Burt, W. H. & Grossenheider, R. P., 1952. A field guide to the Mammals. — Bos-
ton, p. 1-200.

Carpenter, S. J. & La Casse, W. J., 1955. Mosquitoes of North America. — Berk-
eley & Los Angeles, p. 1-360.

Cernosvitov, L. & Evans, A. C, 1947. Lumbricidae (Annelida). With a key to
the common species. — Synops. Brit. Fauna (Linn. Soc). London, p. 1-36.

Chamberlin, R. v., 1921. The Julidae and Isobatidae in North America. — Proc.
Biol. Soc. 34. Washington, D.C., p. 81-84.

— — 1925. The genera Lithobius, Neolithobius, Gonibius, and Zinapolys in Ame-
rica north of Mexico. — - Bull. Mus. Comp. Zool. 57. Cambridge, Mass., p. 441-

504-

1946. On the Chilopods of Alaska. — Ann. Ent. Soc. Amer. 39. Columbus,

Ohio, p. 177-189.

Chamberlin, R. V. & Ivie, W., 1947. The Spiders of Alaska. — Bull. Univ. Utah.
37. Salt Lake City, p. 1-103.

Charitonov, D. E., 1932. Katalog der Russischen Spinnen. — Ann. Mus. Zool. 32,
Beilage (Ac. Sci. URSS). Leningrad, p. 1-206.

1936. (Supplement of the Catalogue.) — Scient. Mem. Gorky State Univ.

2:1. Perm, p. 167-225.

Check-List of North American Birds, 1931. 4. ed. — Amer. Ornith. Union. Lan-
caster, Pa., p. 1-526. (With supplements in The Auk, 61-71, 1944-54.)

Chittenden, F. H., 1923. Notes on the distribution and habits of North American
Phyllotreta (Coleop.). — Proc. Ent. Soc. 25. Washington, D.C., p. 131-139.

Coleopterorum Catalogus, 1910-40. (Junk & Schenkling). 1-170. — Berlin & Haag.

128 CHAPTER I

COLLINGE, W. E., 1943. The distribution of the woodlice of Great Britain. — N. W.

Naturalist. 18. Arbroath, p. 69-73.
CoLWfiLL, W. E., 1943. Some common alfalfa troubles. — Cornell Extension Bull.

616. Ithaca. N.Y., p. 1-8.
CoMSTOCK, J. H. & Gertsch, W. J., 1948. The Spider Book. — Ithaca, N.Y., pp.

I-XI, 1-729.
Creighton, W. S., 1950. The ants of North America. — Bull. Mus. Comp. Zool.

104. Cambridge, Mass., p. 1-585.
Darlington, P. J., Jr., 1927. Aegialia arenaria Muls. in New England, with local

records for other species. — Psyche. 34. St. Albans, Vt., p. 98-99.
DjAKONOV, A., 193 1. Die Geometridenfauna von Kamtshatka. (Russian.) — Ann.

Mus. Zool. Ac. Sci. URSS. 32. Leningrad, p. 385-410.
Dobrzhanskij, T., 1926. Die palaarktischen Arten der Gattung Coccinella L. —

Revue Russe Ent. 20. Moscow, p. 16-32.
DoNiSTHORPE, H. S. J. K., 1927. British Ants. Their life-history and classification.

2. ed. — London, pp. I-XVI, 1-436.
Dyar, H. G., 1928. The Mosquitoes of the Americas. — (Carnegie Inst.) Wash-
ington, D.C., p. 1-616.
Eason, E. H., 1951. Notes on the Chilopoda (Centipedes) of Warwickshire and

Worcestershire. — Ann. Mag. Nat. Hist. (12) 4. London, p. 257-268.
Ehrmann, P., 1937. Mollusca. — Tierwelt Mitteleur. II. Leipzig, p. 1-210.
Ellerman, J. R. & MoRPlsoN-ScOTT, T. C. S., 1951. Checklist of Palaearctic and

Indian Mammals. — • (Brit. Mus.) London, p. 1-810.
Ellis, A. E., 1926. British snails. A guide to the non-marine Gastropoda of Great

Britain and Ireland, pliocene to recent. — Oxford, p. 1-275.
Fall, H. C, 1924. The blueberry leaf-beetle and some of its relatives. — Maine

Agric. Exp. Stat. 319. Orono, p. 81-91.

1929. On the genus Phaedon (Coleoptera). — Pan-Pac. Ent. 5. San Francisco,

p. 145-152.

1934- On certain North American Elateridae, new and old. — Journ. N. Y.

Ent. Soc. 42. New York, p. 7-36.
Ferguson, D. C, 1955. The Lepidoptera of Nova Scotia, i. Macrolepidoptera. —

Bull. N. S. Mus. Sci. 2. Halifax, p. 1-375.
Flerow, C. C, 1934. On the geographical distribution and systematics of Elks or

Moose (Alces Gray). — C. R. Ac. Sci. SSSR. (N. S.) 2. Leningrad, p. 141-143-

1952. Mammalia. 1:2. — Fauna SSSR. 55. Moscow & Leningrad, p. 1-256.

Forbes, W. T. M., 1923, 1948, 1954. The Lepidoptera of New York and neighboring,

states. 1-3. — Mem. Cornell Univ. Agric. Exp. Stat. 68. 274. 329. Ithaca, N. Y.,

pp. 1-729, 1-263, 1-433.
Ford, E. B., 1946. Butterflies. — The New Naturalist. London, pp. I-XIV, 1-368.
Franz, H., 1947. Beitrage zur Curculioniden-Systematik. i. Revision der europa-

ischen Arten der Gattung Miarus Steph. — Ann. Naturh. Mus. 55. Wien, p .210-

264.
Gertsch, W. J., 1934. Notes on American Lycosidae. — Amer. Mus. Nov. 693. New

York, p. 1-25.

1939- A revision of the typical crab-spiders (Misumeninae) of America north

of Mexico. — Bull. Amer. Mus. Nat. Hist. 76. New York, p. 277-442.

Bibliography 129
1949- American Spiders. — (The New Illustrated Naturalist.) New York,

pp. I-XIII, 1-285.
Gibson, A., 1920. Lepidoptera. — Rep. Can. Arct. Exp. 3. Ottawa, p. 1-58.
Hackman, W., 1954. The spiders of Newfoundland. — Acta Zool. Fenn. 79. Hel-

singfors, p. 1-99.
Handbook of British birds, 1945. By H. F. Witherby, F. C. R. Jourdain, N. F. Tice-

hurst, and B.W. Tucker. I-V. 3. impr. London, pp. 1-326, 1-352, 1-387, 1-461,

1-381.
Handbuch der Pflanzenkrankheiten (P. Sorauer), 1949-54. IV: 1-2. V: 1-2. 5. ed.

Berlin & Hamburg.
Harrington, W. H., 1891. Notes on a few Canadian Rhynchophora. — Can. Ent.

23. London, Ont., p. 21-27.
Hartert, E., 1903, 1912-21, 1921-22, 1932-38. Die Vogel der palaarktischen Fauna.

I. II. III. Erganzungsband. Berlin, pp. 1-2328, 1-602.
Hatch, M. H., 1947. The Chelifera and Isopoda of Washington and adjacent regions.

— Univ. Wash., Publ. Biol. 10: 5. Seattle, Wash., p. 157-274.

1953- The beetles of the Pacific Northwest. I. Introduction and Adephaga.

— Ibidem. 16. p. 1-340.

1955- Bradycellus harpalinus Serv. in North America. — Coleopt. Bull. 9.

Rochester, N. Y., p. 10.
Henriksen, K. L., 1937. Insects collected on the fifth Thule expedition. — Rep.

5th Thule Exp. 1921-24. II: 8. Copenhagen, p. 1-34.
Heydemann, F., 1943. Die Bedeutung der sogenannten Dualspecies (Zwillingsarten)

fiir unsere Kenntnis der Art- und Rassenbildung bei Lepidopteren. — Ent.

Zeit. 104. Stettin, p. 116-142.
Heyden, L. von, 1880-98. Catalog der Coleopteren von Sibirien. With 3 suppl. —

(Deutsch. Ent. Gesellsch.) Berlin, pp. I-XXIV, 1-224, 1-2 17, 1-84, 1-24.
Holdhaus, K., 1954. Die Spuren der Eiszeit in der Tierwelt Europas. — Abh.

Zool.-Bot. Gesellsch. 18. Wien, p. 1-493.
Holdhaus, K. & Lindroth, C. H., 1939. Die europaischen Koleopteren mit boreo-

alpiner Verbreitung. — Ann. Naturh. Mus. 50. Wien, p. 123-293.
Holland, W. J., 1903 (1917, 1934). The Moth Book. — New York, pp. I-XXIV,

1-479.
Holm, A., 1950. Studien iiber die Spinnenfauna des Tornetraskgebietes. — Zool.

Bidr. 29. Uppsala, p. 103-213.
Holzapfel, Monika, 1932. Die Gewachshausfauna des Berner Botanischen Gartens.

— Rev. Suisse Zool. 39. Geneve, p. 325-374.

Hopping, R., 1932. Anoplodera planata S. & H. — Can. Ent. 64. Orillia, Ont., p. 72.
Hovanitz, W., 1950. The biology of Colias butterflies, i. The distribution of the

North American species. — Wasm. Journ. Biol. 8. San Francisco, p. 49-75.
Hubendick, B., 1 95 1. Recent Lymnaeidae. Their variation, morphology, taxonomy,

nomenclature, and distribution. — K. Vet. Ak. Handl. (4) 3. Stockholm, p.

1-223.
Jacobi, a., 1931. Das Rentier, eine zoologische Monographic der Gattung Rangifer.

— Zool. Anz. 96 (Erganzungsband). Leipzig, p. 1-264.

Jacobson, G. G., 1905-15. Zuki Rossij. (The Beetles of Russia, &c.) i-io. (Rus-
sian.) — Leningrad, p. 1-864.

9 — 565597 Lindroth

130 CHAPTER I

Jawlowski, H., 1930. On European Diplopoda introduced to North America. —

Fragm. Faun. Mus. Zool. Pol. 1:7. Warszawa, p. 181-185.
JoHANSEN, F., 1926. On the woodlice (Oniscoidea) occurring in Canada and Alaska.

— Can. Field-Natur. 40. Ottawa, p. 165-167.
Johnson, C. W., 1906. On the distribution of Helix hortensis Mueller, in North

America. — Nautilus. 20. Philadelphia, p. 73-80.
Johnston, E. C, 1950. Lepidoptera of the Pribilof Islands, Alaska. — Lep. News.

4. New Haven, Conn., p. 27-30.
Jordan, D. S., Evermann, B. W., & Clark, H. W., 1930. Check list of the fishes

and fishlike vertebrates of North and Middle America, &c. — Rep. U.S. Comm.

Fisheries. 1928: 2. (Dept. of Commerce.) Washington, D.C., p. 1-670.
JouRDAiN, F. C. R., 1933. On the Palaearctic element in the A. O. U. 'Check-List'

(4th edit.). — The Auk. 50. Lancaster, Pa., p. 201-204.
JuLiN, E., 1949. De svenska daggmaskarterna. (The Swedish Earthworms.) — Ark.

f. Zool. 42 A. Stockholm, p. 1-58.
K.^STON, B. J., 1948. Spiders of Connecticut. — Bull. State of Conn. Geol. & Nat.

Hist. Surv. 70. Hartford, Conn., p. 1-874.
Kennard, a. S. & Woodward, B. B., 1926. Synonymy of the British non-marine

MoUusca. — (Brit. Mus.) London, pp. I-XXIV, 1-447.
Kevan, D. K. McE., 1943. Study of an introduced North American freshwater

Mollusc, Stagnicola catascopium (Say). — Proc. R. Soc. B 61. Edinburgh, p.

430-461.
Kloet, G. S. & Hincks, W. D., 1945. A check list of British insects. — Arbroath,

pp. I-LIX, 1-483.
Klots, a. B., 1 95 1. A field guide to the Butterflies of North America, east of the

Great Plains. — Cambridge, Mass., pp. I-XVL i~349-
Kozhantshikov, L B., 1937. Noctuidae, Agrotinae. — Fauna SSSR. 13: 3. Moscow

& Leningrad, pp. I-XVI, 1-675.

1950. Orgyidae. — Ibidem. 12. p. 1-582.

Kraepelin, K., 1 90 1. Uber die durch den Schiffsverkehr in Hamburg eingeschlepp-

ten Tiere. — Jahrb. Hamb. Wissensch. Anst. 18 (1900). Hamburg, p. 185-209.
Krogerus, H., 1954. Investigations on the Lepidoptera of Newfoundland. I. Macro-

lepidoptera. — Acta Zool. Fenn. 82. Helsingfors, p. 1-80.
KusNEZOV, N. J., 1925. Some new Eastern and American elements in the fauna

of Lepidoptera of Polar Europe. — C. R. Ac. Sci URSS (A). Leningrad, p. 119-

122.
Leclercq, J., 1954. Monographie systematique, phylogenetique et zoogeographique

des Hymenopteres Crabroniens. — Liege, p. 1-37 1.
Lehmann, H. C. & Klinkowski, M., 1942. Zur Pathologic der Luzerne, i. Die

schadlichen Russelkafer (Curculionidae). — Ent. Beihefte. 9. Berlin-Dahlem, p.

1-78.
Leng, C. W., 1920, 1927-48. Catalogue of the Coleoptera of America, north of

Mexico. With 5 suppl. — Mount Vernon, N. Y.
Leonard, M. D., 1926. A list of the insects of New York. With a list of the spiders

and certain other allied groups. — Mem. Cornell Univ. Agric. Exp. Stat. loi.

Ithaca, N. Y., p. 1-1121.
Lepidopteroriim Catatogus, 191 1-39. (Ed. F. Bryk.) — i-94- Berlin & Haag.

Bibliography 131

LiCHAREV, I. M. & Rammelmayer, E. S., 1952. Terrestrial Mollusca of SSSR.

(Russian.) — Opredel. Fauna SSSR. 43. Moscow & Leningrad, p. 1-5 12.
LiNDROTH, C. H., 1954. Carabidae common to Europe and North America. — Co-

leopt. Bull. 8. Washington, D. C, p. 35-52.

1955- The Carabid beetles of Newfoundland, including the French islands

St. Pierre and Miquelon. — Opusc. Ent., Supp). XII. Lund, p. 1-168.

Locket, G. H. & Millidge, A. F., 1951-53. British spiders. I. II. — (Ray Soc.)

London, pp. 1-3 10, 1-449.
Lowrie, D. C. & Gertsch, W. J., 1955. A list of the spiders of the Grand Teton

Park Area, &c. — Amer. Mus. Nov. 1736. New York, p. 1-29.
Lutz, F. E. & CocKERELL, T. D. A., 1920. Notes on the distribution and biblio-
graphy of North American bees, &c. — Bull. Amer. Mus. Nat. Hist. 42. New

York, p. 491-641.
Mader, L., 1926-37. Evidenz der palaraktischen Coccinelliden und ihrer Aberra-

tionen in Wort und Bild. — Zeitschr. Ver. Naturbeob. Wien; Ent. Anz. Wien;

Ent. Nachr.-Blatt Troppau. pp. 1-412, I-XII.
Malkin, B., 1953. New records of Arachnida from Alaska. — Pan-Pac. Ent. 29.

San Francisco, p. 205-206.
Mandahl-Barth, G., 1938. Land and freshwater Mollusca. — Zool. of Icel. 4.

Copenhagen, p. 1-3 1.
Marshall, J. F., 1938. The British Mosquitoes. — (Brit. Mus.) London, pp.

I-XII, I-34I.
Mc DuNNOUGH, J. H., 1938. Check list of the Lepidoptera of Canada and the

United States of America, i. — Mem. S. Calif. Ac. Sci. i. Los Angeles, p. 1-272.

1949- Revision of the North American species of the genus Eupithecia (Lepi-
doptera, Geometridae). — Bull. Amer. Mus. Nat. Hist. 93. New York, p. 537-
728.

1950. Species of Euxoa of eastern North America, &c. — Ibidem. 95: 6. p.

355-408.
Meeuse, a. D. J. & Hubert, B., 1949. The Mollusc fauna of glasshouses in the

Netherlands. — Basteria. 13. Liege, p. 1-30.
Michaelsen, W., 1903. Die geographische Verbreitung der Oligochaeten. — Berlin,

p. 1-186.
MoNTSHADSKY, A., 1 95 1. Culicid larvae. (Russian.) — Opred. Fauna SSSR. 37.

Moscow & Leningrad, p. 1-290.
Mozley, a., 1935. The fresh-water and terrestrial Mollusca of Northern Asia. — ■

Trans. R. Soc. 58 (1936). Edinburgh, p. 605-695.
1938. The fresh-water Mollusca of sub-arctic Canada. — Can. Journ. Research.

16 D. Ottawa, p. 93-138.
Muesebeck, C. F. W., Krombein, K. V., & Townes, H. K., 1951. Hymenoptera of

America north of Mexico. Synoptic catalog. — U.S. Dept. Agric, Agric. Monogr.

2. Washington, D. C, p. 1-1420.
MuLLER, G., 1949-53. I Coleotteri della Venezia Giulia. II. — Publ. Centro Sperim.

Agric. Forest. 4. Trieste, p. 1-685.
MuNROE, E. G., 1 95 1. The geographic variation of Dasyuris polata (Duponchel) in

North America (Lepidoptera: Geometridae). — ■ Can. Ent. 83. Ottawa, p. 290-

294.

132 CHAPTER I

Natvig, L. R., 1948. Contributions to the knowledge of the Danish and Fennoscan-
dian Mosquitoes. Culicini. — Norsk Ent. Tidsskr., Suppl. i. Oslo, pp. I-XXIII,

1-567.
Needham, J. G. &WESTFALL, M. J., Jr., 1955. A manual of the Dragonflies of North

America (Anisoptera). — Berkeley & Los Angeles, p. 1-6 15.
NuNENMACHER, F. W., 1 934- Studies among the Coccinellidae. 7. — Pan-Pac. Ent.

10. San Francisco, p. 11 3-1 14.
Oekland, F., 1925. Die Verbreitung der Landgastropoden Norvvegens. — Skr.

Norske Vid.-Ak., Mat.-Naturv. Klasse. 8. Oslo, pp. I-VIII, 1-168.
Olsoufiev, N. G., 1937. Tabanidae. — Fauna SSSR. 7: 2. Moscow & Leningrad,

pp. I-XIII, 1-434-
OuGHTON, J., 1948. A zoogeographical study of the land snails of Ontario. — Univ.

Toronto, Biol. Ser. 57. Toronto, pp. I-XI, 1-126.
Palmen, E., 1946. Die Landisopoden Finnlands. — Ann. Zool. Soc. Zool. Bot.

Fenn. Vanamo. 11. Helsingfors, p. 1-35.

1948. The Chilopoda of Eastern Fennoscandia. — Ibidem. 13. p. 1-45.

1951. A survey of the Oniscoidea (Isopoda Terr.) of Newfoundland. — Ibidem.

14. p. 1-27.

1952. Survey of the Diplopoda of Newfoundland. — Ibidem, 15. p. 1-31.

1954. Survey of the Chilopoda of Newfoundland. — Arch. Soc. Zool. Bot.

Fenn. Vanamo. 8. Helsingfors, p. 131-149.
Peters, J. L., 193 1-5 1. Check-list of birds of the world. I-VII. Cambridge,

Mass.
Petersen, B., 1954. Some trends of speciation in the cold-adapted Holarctic fauna.

— Zool. Bidr. 30. Uppsala, p. 233-314.
Peterson, R. L., 1952. A review of the living representatives of the genus Alces. —

Contr. R. Ont. Mus. Zool. & Paleont. 34. Toronto, p. 1-30.
Philip, C. B., 1947. A catalog of the blood-sucking fly family Tabanidae (Horseflies

and Deerflies) of the Nearctic region north of Mexico. — Amer. Midi. Nat. 37.

Notre Dame. Ind., p. 257-324.
Pierce, W. D., 1919. Contributions to our knowledge of the weevils of the super-
family Curculionoidea. — Proc. Ent. Soc. 21. Washington, D. C, p. 21-36.
Pilsbry, H. a., 1939-48. Land Mollusca of North America (north of Mexico).

I: 1-2. II: 1-2. — Ac. Nat. Sci., Monogr. 3. Philadelphia, pp. I-XVII, 1-994;

I-VI, 1-1113.
PiTTiONi, B,, 1942-43. Die boreoalpinen Hummeln und Schmarotzerhummeln. —

Mitt. Kgl. Naturw. Inst. 15-16. Sofia, pp. 155-218, 1-78.
Plavilstshikov, N. N., 1936, 1940. Cerambycidae. 1-2. — Fauna SSSR. (N. S.) 7.

22. Moscow & Leningrad, pp. 1-6 12, 1-784.
Rausch, R., 1953. On the status of some Arctic Mammals. — Arctic. 6: 2. Ottawa.

p. 91-148.
Reimoser, E., 1919. Katalog der echten Spinnen (Araneae) des Palaarktischen Ge-

bietes. — Abh. Zool. Bot. Gesellsch. 10: 2. Wien, p. 1-280.
RocQUE, A. La, 1953. Catalogue of the recent Mollusca of Canada. — Bull. Nation.

Mus. Can. 129. Ottawa, pp. I-IX, 1-406.
ROEWER, C. F., 1942, (1944) 1954. Katalog der Araneae. 1-2. Bremen, pp. 1-1040.

1-160; Bruxelles, p. 1-1751.

Bibliography 133

Sahlberg, J., 1880. Bidrag till Nordvestra Sibiriens insektfauna. Coleoptera, &c. —

K. Svensk. Vet. Ak. Handl. 17. Stockholm, p. 1-115.
Schmidt, A., 1922. Aphodiinae. — Das Tierreich. 45. Berlin & Leipzig, p. 1-614.
ScHUBART, O., 1934. Diplopoda. — Tierwelt Deutschl. 28. Jena, p. 1-3 18.
Seitz, a. (with co-operators) 1909-24. Die Gross- Schmetterlinge der Erde. 1-160.

Stuttgart.
Shadin, W. I., 1935. tJber die okologische und geographische Verbreitung der

SiisswassermoUusken in der USSR. — Zoogeogr. 2. Jena, p. 495-554.

1952. Freshwater MoUusca. (Russian.) — Opred. Fauna SSSR. 46. Moscow

& Leningrad, p. 1-376.

Simon, E., 1914-37. Les Arachnides de France. 6: 1-5. Paris, p. 1-1298.
Simpson, G. G., 1945. The principles of classification and a classification of Mam-
mals. — Bull. Amer. Mus. Nat. Hist. 85. New York, pp. I-XVI, 1-350.
Skorikov, a. S., 1937. Die gronlandischen Hummeln im Aspekte der Zirkumpolar-

fauna. — Ent. Medd. 20. Copenhagen, p. 37-64.
Smith, F., 1917. North American earthworms of the family Lumbricidae in the

collection of the United States National Museum. — Proc. U. S. Nat. Mus. 52.

Washington, D. C, p. 157-182.
Smith, J. B., 1893. Catalogue of the Lepidopterous superfamily Noctuidae found

in Boreal America. — Bull. Smiths. Inst. 44. Washington, D. C, p. 1-424.
Smith, M. R., 195 i, vide Muesebeck, Krombein, & Townes.
Stackelberg, a. a., 1937. Culicidae, Culicinae. (Russian.) — Fauna SSSR. 3: 4.

Moscow & Leningrad, pp. I-X, 1-258.
Stegmann, B., 1938. Principes generaux des subdivisions ornithogeographiques de

la region palearctique. — Fauna SSSR (N.S.). 19. Moscow & Leningrad, p.

I-I57-

Stephenson, J., 1930. The Oligochaeta. — Oxford, pp. I-XVI, 1-978.

Stitz, H., 1939. Ameisen oder Formicidae. — Tierwelt Deutschl. 37. Jena, p. 1-428.

Taylor, J. W., 1902-21. Monograph of the land & freshwater Mollusca of the British
Isles. II-IV. — Leeds, pp. 1-3 12, 1-522, 1-160.

Titus, E. G., 1911. The genera ■ Hypera and Phytonomus (Coleoptera, Family
Curculionidae) in America, north of Mexico. — Ann. Ent. Soc. Amer. 4. Colum-
bus, Ohio, p. 383-473.

Urquhart, F. a. & Beaudry, J. R., 1953. A recently introduced species of European
grasshopper. — Can. Ent. 85. Ottawa, p. 78-79.

Valle, K. J., 1933. Die Lepidopterenfauna des Petsamogebietes. — Ann. Zool.
Soc. Zool. Bot. Fenn. Vanamo. i. Helsingfors, pp. I-VII, 1-262.

1952. Die Verbreitungsverhaltnisse der ostfennoskandischen Odonaten. — -

Acta Ent. Fenn. 10. Helsingfors, p. 1-87.

1955. Odonata from Newfoundland. — Ann. Ent. Fenn. 21. Helsingfors, p.

57-60.
Vandel, a., 1945. La repartition geographique des Oniscoidea. — ■ Bull. Biol.

France et Belg. 79. Paris, p. 221-272.
Van Name, W. G., 1936. The American land and fresh-water Isopod Crustacea. —

Bull. Amer. Mus. Nat. Hist. 71. New York, p. 1-535.
1940- A supplement to the American land and fresh-water Isopod Crustacea.

— Ibidem. 77. p. 109-142.

134 CHAPTER I

Verhoeff, K. W., 1937. Diplopoda u. Chilopoda. — Tierwelt Mitteleur. II. Leipzig,

p. 1-120.
Wach'tler, W., 1937. Isopoda. — Tiervvelt Mitteleur. II. Leipzig, p. 225-317.
Walker, E. M., 1927. The woodlice or Oniscoidea of Canada (Crustacea, Isopoda).

— Can. Field-Natur. 41. Ottawa, p. 173-179.

1953- The Odonata of Canada and Alaska, i. — Toronto, p. 1-292.

W.^rner, Rose E., 1952. Another European weevil, Pentarthrum huttoni Woll. in
North America. — Coieopt. Bull. 6. Washington, D. C, p. 51-52.

Weyrauch, W., 1937. Zur Systematik und Biologic der Kuckuckswespen Pseudo-
vespa, Pseudovespula und Pseudopolistes. — Zool. Jahrb., Abt. Syst. 70. Jena,
p. 243-290.

W^iLCOX, J., Mote, D. C. & Childs, L., 1934. The root-weevils injurious to straw-
berries in Oregon. — Bull. Agric. Exp. Stat., Oreg. State Agric. Coll. 330. Cor-
vallis, Oreg., p. 1-109.

Williams, C. B., 1950. American moths in Britain. — Lep. News. 4. New Haven,
Conn., p. 62.

Williams, S. R. & Hefner, R. A., 1928. The Millipedes and Centipedes of Ohio.

— Ohio State Univ. Bull. 33. Columbus, Ohio, p. 91-147.

Wilson, E. O., 1955. A monographic revision of the ant genus Lasius. — Bull.

Mus. Comp. Zool. 113. Cambridge, Mass., p. 1-205.
Wilson, E. O. & Brown, W. L., Jr., 1955. Revisionary notes on the sanguinea and

neogagates groups of the ant genus Formica. — Psyche. 62. Watertown, Mass., p.

108-129.
Wynne-Edwards, V. C, 1952. Freshwater vertebrates of the Arctic and Subarctic.

— Bull. Fish. Research Board Can. 94. Ottawa, p. 1-28.
0KLAND, vide Oekland.

Chapter II

THE HUMAN TRANSPORT OF ANIMALS ACROSS THE
NORTHERN ATLANTIC

The five criteria of an introduced species

JVlan has purposely carried several animal species, and not only domesticated
ones, across the Atlantic in either direction, and many of them have become per-
manently established (instances, vide above, p. i6). By far more numerous are
the animal and plant species which were carried over unintentionally by all kinds
of ships. Of course only a small fraction of them were able to find and colonize
suitable ground in the new continent and the successful event, with very few ex-
ceptions, was not apparent until long afterwards when it was impossible to re-
construct the procedure accurately. Consequently, in trying to sort out the foreign
elements of recent introduction from the indigenous fauna of a continent, we are
usually restricted to second hand evidence: certain features of distribution, abund-
ance, ecology, &c., shown by a species in the country where it is believed to have
been introduced. There are reasons to presume that such characteristics of an
immigrant into North America from Europe, or vice versa, as a rule have not been
completely blotted out during the comparatively short period of, at most, 460 years.
The foremost criteria of an introduced species are:

I. THE HISTORICAL CRITERION. This includes all cases in which we are able
to trace by historical evidence, if not the very landing of a species, which is a
rare exception, at least parts of the following expansion. It is true that the period
open to scientific survey is too short to control the normal changes of a species'
area, but the recent immigrants often show a surprising power of rapid dispersal
during the interval between first establishment and balanced distribution. Thus
the introduction, temporary extermination, re-introduction, and final victorious
progress of the Colorado-Beetle {Leptinotarsa decemlineata Say) in Europe (fig. 3),
as well as the history of the Gypsy Moth {Lymantria dispar L.) in North America,
were followed in detail from year to year.

Similar instances are provided by Passer domesticus L. (the English sparrow) and
Sturnus sturnus L. (the Starling; fig. 4), which are known to be introduced into North
America, as well as the Muskrat {Fiber or Ondatra zibethicus L.) in Europe (fig. 5),
and by Pieris rapae L. (the Cabbage Butterfly), Phytonomus posticus Gyll. (the

136

CHAPTER II

FIG. 3. Spread of the Colorado Beetle {Leptinotarsa decem-
lineata Say) in Europe until World War II, starting from
Senejac near Bordeaux about 1920. The present eastern
limit is tentative only.
(According to Alfaro, 1941, i943, and O. Ahlberg, in lift.)

(Photo P. Ardo.)

Alfalfa Weevil; fig. 6), the ground-beetle Carahus nemoralis Miill. (fig. 7)^ and many
others, which are supposed to be European introductions, in part just because of
their rapid expansion. Another ground-beetle, Pterostichus melanarius 111. {vulgaris

^ The statement made by Evans (1952, p. 217-218), that Carabus nemoralis had been
purposely introduced into eastern Canada as a general predator, is due to a misinterpreta-
tion of Coscns (1923, p. 10).

The human transport of animals across the Northern Atlantic

137

FIG. 4. Spread of the Starling (Sturnus sturnus L.) in North
America since its introduction into New York in 1890.

(After Wing, 1943.)

(From Rosenberg, Faglar i Sverige.)

auctt.), and a small clover-feeding weevil, Tychius picirostris F., are spreading fast
over the northern parts of the American continent at this very time.

Other examples of recent dispersal, in Newfoundland and adjacent regions, are
mentioned below (p. 151).

138

CHAPTER II

(From Burt & Grossenheider,
1952.)

FIG. 5. Spread of the Muskrat, Ondatra {Fiber) zibethica L., in Central Europe since
its introduction 40 km. SSW of Prague, in 1905.

(After Ulbrich, 1930.)

On the other hand, introduced species may very well be conservative and remain
for a long time restricted to the port of introduction or its immediate vicinity.
This, as suggested by Brown (1950, p. 197), is probably due to the fact that these
species are often confined to open ground, the surrounding forests forming an
efficient barrier against further distribution. The click-beetle Agriotes lineatus L.,
found in Newfoundland as early as about 1840, is still restricted to the Avalon
Peninsula of the southeast where it was first introduced.

There are also instances, at least in Europe, of animals showing a rapid tem-
porary expansion of area not due to human influences, for example certain aquatic
birds, the Carabid beetle Amara majuscula Chaud. (Lindroth, 1949, p. 625), and
the Click-beetle Corymbites cupreus F. (Holdhaus & Lindroth, 1939, pp. 184,
260), in Scandinavia and Finland. In these cases, however, the migration has taken
place from a larger continent (Asia and SE Europe) towards the periphery (At-
lantic Europe), never in the opposite direction.

2. THE GEOGRAPHICAL CRITERION. The distribution of an introduced species
is often "immature": the area is more or less restricted, usually to some coastal
district, and "unnatural", in the sense that its limits can hardly be co-ordinated
with any external factor (climate, soil, vegetation, &c.) which could explain this
restriction. A disjunct (broken-up) area is especially suspicious; for instance if
a species, otherwise not coast-bound, occurs in North America only in the North-
east and in the Pacific Northwest. Thus the ground-beetle Bembidioti lampros
Hbst. is restricted to British Columbia and the very town of St, John's, New-

1

The human transport of animals across the Northern Atlantic

139

(From EssiG, 1931.)

FIG. 6. Spread of the Alfalfa Weevil, Phytonomiis posticus Gyll. (variabilis Hbst.)

in Central and Western U.S.A. since its introduction in Utah about 1902.

Black area = before and incl. of 1910

Densely hatched area = before and incl. of 1919

Thinly hatched area = ,, ,, ,, ,, 1952.

Since 1952 the beetle has appeared also in the eastern states.

(According to TiTUS, 191 1, Reeves, 1927, and information from Bureau of Agriculture,
Washington, D.C.)

foundland. Other Carabid beetles showing almost as wide a gap in their North
American distribution are Carabus granulatus L. (fig. 8), C. nemoralis Miill. (fig. 7),
Agonum miilleri Hbst., Clivina fossor L., and Pterostichus melanarius 111. (for further
examples, vide p. 144). This pattern of distribution in introduced species holds also
true for terrestrial Molluscs (Pilsbry, 1948, p. 522). An "immature" distribution
as a sign of introduction in Newfoundland is shown by Bembidion tetracolum Say
(ustulatum auctt.), Clivina fossor L. (maps, figs. 12-13), and many others.

3. THE ECOLOGICAL CRITERION. The ecology of a species may indicate its
character of a foreigner. It is most striking to a European biologist that in North
America the "culture steppe", the open, dry (artificially drained) land in and
around ports and other settlements, especially along the Atlantic coast, is inhabited
by a flora and fauna of pronounced European character. Certain indigenous North
American species may invade such spots too, but a plant or an animal restricted to
this habitat is very likely to be a recent introduction.

An instance is given by the ground-beetle genus Amara. With very few excep-
tions, its members are confined to dry, open ground, some of them inhabiting

140

CHAPTER II

FIG. 7. Distribution of Carabus nemoralis O. F. Miill., intro-
duced into North America.

(From Faune de France.)

The human transport of animals across the Northern Atlantic

141

FIG. 8. Distribution of Carabus granulatus L., repeatedly
introduced into North America. The figured specimen is of
forma typica from Sweden.

Black dots = forma typica. Pointed dots ~ sbsp. hibernicus
Lth.

(Photo P. Ardo.)

142 CHAPTER II

the alpine region, others moorland and sandflats at a lower altitude, or native
meadows, including the epilittoral zone of lakes or the sea. In the maritime pro-
vinces of Canada, however, a notably high number of Amara species is restricted
to cultivated ground, waste places, &c., to the "culture steppe". From Labrador,
Newfoundland, and Nova Scotia, 24 species of Amara are known altogether; 13
of these inhabit ground more or less untouched by man, as just described, only
one, Amara fulva DeG., being a recent introduction. The remaining 11 are bound
to waste ground or arable land or at any rate are clearly favoured by human culture;
of these 6 are European introductions {A. aenea DeG., apricaria Payk., aulica
Panz., hifrons Gyll., familtaris Dft., lunicoUis Schio.) and only 5 indigenous to
North America {A. avida Say, impuncticolUs Say, latior Kby., pallipes K.hy.,pat-
ruelis Dej.).

4. THE BIOLOGICAL CRITERION. The natural history of a species may preclude
any idea of regarding it as indigenous. This evidence is very strong in plant-feeders
and parasites monophagously bound to a single host. The Colorado beetle {Lep-
tinotarsa decemlineata Say) was unable to live in Europe before the cultivation of
potatoes was started in the late i6th century and the same applies to several pests
of Citrus trees in North America before the introduction of their hosts. The weevils
[Curcidionidae) and other insects feeding on various species of clover (Trifolium)
in eastern Canada and northern New England had no indigenous host at their
disposal until the transatlantic trade started.

5. THE TAXONOMic CRITERION. This is especially valuable in cases where a
species occurs on a continent partly as indigenous, partly as a result of recent in-
troduction. In certain cases the two forms may be taxonomically separate, even
if the distinguishing characters are of less than subspecific value. The form of the
lady-bird Cocciriella ll-punctata L. occurring on the Pacific coast of North America
(as far north as Alaska) has been separated as a subspecies or variety, menetriesi Muls.,
and is probably indigenous. The same species on the Atlantic coast is not distin-
guishable from the European forma typica and is no doubt a late introduction. A
taxonomic separation may also be possible between one indigenous and one intro-
duced form of the Carabid beetle Amara lunicoUis Schio.

Due to subspecific difi"erences it can also be proved that Carahus granulatus L.
(fig. 8) has been introduced at least twice into the North American continent. Nova
Scotia and New Brunswick are inhabited by the sbsp. hibernicus Lth. which in
Europe seems to occur in unmixed populations only in Ireland (Lindroth, 1955b),
and it therefore certainly originated from this island. In other parts of eastern
North America, as well as in the Pacific Northwest, granulatus is represented by

The human transport of animals across the Northern Atlantic 143

its typical form which must have been brought over independently from parts of
Europe other than Ireland, though possibly from Great Britain.

The distribution of long-winged (macropterous) and short-winged (brachy-
pterous) individuals of Notiophilus biguttatus F. in Newfoundland, the only part
of North America where it has been found, clearly demonstrates its introduction
to and subsequent dispersal from the Avalon Peninsula (fig. 14).

Westward transport

From where did they come?

Most of the animal species stated or believed to be introduced into North America
are widespread in Europe and great difficulties are therefore encountered in
deciding on that basis alone from which country they once emigrated. As will be
shown in the following chapters, it is easier to find out the possible directions of
transport by studying the North Atlantic trade during the past three or four cen-
turies. It may anyhow be worth while to analyze the distribution in western Europe,
and also in North America, of the Carabid beetles in the list above (p. 40 a.f.)
regarded as introduced into the latter continent.

A summary of table 2 shows that the British Isles are the only part of the
European west coast where all 40 species of Carabidae introduced from Europe
to North America occur, though two of them (Carabus auratus L. and cancellatus
111.) are occasional only and certainly reached North America from some other
European country. The same can be said about Amara anthobia Villa, quite re-
cently discovered in Britian, and Trechus obtusus Er., occurring there in its short-
winged (brachypterous) form only, whereas in North America (the Pacific North-
west) it is constantly long-winged (macropterous), a form otherwise restricted to
southern Europe, north to SW France.

The remaining 36 species of table 2 may very well have been introduced from
the British Isles and it is interesting to observe that all of them have been found
in southwestern England, 22 species even on ballast-places or localities in the
old ports where such are supposed to have been situated (besides the 20 species
treated on p. 193 a.f., below, Clivina collaris Hbst. diudfossor L. which, according
to specimens in the Bristol Museum, have been taken on the Avon River banks).

The 23 Carabid species introduced into Newfoundland all occur in southwestern
England, as do the 5 additional species found on the mainland of eastern Canada
[Bembidion properans Steph., Carabus granulatus L., Clivina collaris Hbst., Stomis
pumicatus Panz., and Trechus discus F.)

^\ '^^ y^J

144

CHAPTER II

TABLE 2. Distribution of Carabid beetles regarded as European introductions in

North America. »

Three cosmopolitan species {Perigona nigriceps Dej., Plochionus pallens F., and
Somotrichus elevatus F.) are excluded.

Acupalpus meridiamis L. . .
Agonum miilleri Hbst. . . .

A. riificorne Gze

Atnara aenea DeG

A. anthobia Villa

A. apricaria Payk

A. aulica Panz

A. bifrons Gyll

A. familiaris Dit

A. fulva DeG

A. liinicollis Schio.^ ....
Anisodactyliis binotatus F. .
Asaphidion flavipes L. . . .
Bembidion bruxellense Wesm.

(rupestre auctt.) ....

B. lampros Hbst

B. properans Staph

B. stephensi Crotch ....

B. tetracoliim Say (ustulatimt
auctt.)

Bradycelliis harpalinus Serv.
Calathus fuscipes Gze. . . .
Carabus aiiratiis L

C. cancellatiis III

C. gramdatus L

C nemoralis Miill

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^ The European records were compiled from Blair (1931), Fuente (19 18-21), Jeannel
(1941-42), Johnson & Halbert (1902), Lindroth (1945), Palmer (1946), Pearce (1926-27), &c.
Several unpublished British records from the museums of Bristol, Exeter, Plymouth, and
the British Museum, are included. Valuable information was received from Mr. G. H. Ashe,
Colyton (S. Devon), and Mr. R. S. George, Gloucester.

* Vide map, fig. 16.

* Amara liinicollis is probably introduced in NE North America only.

'' Bembidion properans was not kept separate from lampros by Pearce (1926-27) but both
are doubtless widely spread in Dorset.

The human transport of animals across the Northern Atlantic

H5

Clivina collaris Hbst

C. fossor L

Harpahis offinis Schrk. (aenetis auctt.)
H. rufipes DeG. {pubescens Miill.) . . .

Licinus punctatulus F

Nebria brevicollis F

Notiophilus biguttatiis F

Pristonychiis complanatus Dej

P. terricola Hbst

Pterostichus melatiorius III. (vulgaris

auctt.)

Pterostichus strenuus Panz

Stomis puniicatus Panz

Tachys parvulus Dej

Trechus discus F

T. obtusus Er

T. rubens F

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Conditions are similar as regards weevils [Curculionidae). Of 69 species regarded
as European introductions in North America, 61 belong to the British fauna. The
31 Newfoundland species of the same provenience all occur there.

37 species of " Iso-Myriapods"- {Chilopoda, Diplopoda, Isopoda terrestria) and
14 species of non-marine Gastropoda in the fauna of Newfoundland are regarded
as introduced and all of them are known from the British Isles; of the other 10
European species of Iso-Myriapods found on the North American mainland, 5
are British (for explanation, vide p. 168 a.f.).

The fauna of Newfoundland as an illustration
The European species

It is a well-known fact that the flora of Newfoundland contains a notably large
number of plants in common with Europe. The recent "Enumeratio" by Rouleau^

^ Rouleau's list (1949) apparently includes species originally cultivated in Newfoundland
and later escaped and naturalized.

10—565597 Litidroth

146 CHAPTER II

(1949) lists 1107 species of flowering plants (Phanerogams) from the island, of
which almost exactly half, 556 species, grow in Europe (incl. the British Isles,
but excl. Iceland) though in some cases as a different subspecies. The floristic rela-
tionship is quite striking in certain families: in Boraginaceae all 12 Newfoundland
species are "Europeans", in Cruciferae 35 out of 50, in Leguminosae 18 of 27, in
Lahiatae 12 of 19, in Scrophulariaceae 25 of 40.

The European introductions in the Newfoundland flora can easily be extracted
from records given in "Gray's Manual" (Fernald, 1950). Of the 556 "Europeans"
mentioned, 227 species are regarded as such in North America, that is no less
than 21 per cent of the whole flora of the island. ^ Only 4- of these are lacking in
the British Isles (Clapham, Tutin & Warburg, 1952).

The lower terrestrial fauna of Newfoundland shows similar features though,
unfortunately, only a restricted number of taxonomical groups has so far been
subjected to a modern revision, including a detailed comparison with European
specimens. Table 3 therefore represents a rather artificial selection.

In the first place it is notable how variable is the European element within
the different groups treated in table 3, consisting (next-to-last column) of from
100 per cent identical species among terrestrial Isopods, to 14 per cent among
the Odonata. As demonstrated by the last column, this is mainly due to the different
influence of introduced species. This, in its turn, is easily explained by the utterly
different suitability for passive, anthropochorous transport of the selected groups
of animals, as will be more closely elucidated in the following pages (p. 198 a.f.)

In spite of the unequal importance of an introduced European element in different
taxonomic groups, its striking dominance in the Newfoundland fauna, as compared
with that of the adjacent mainland, is quite clear (cf. maps, figs. lo-ii). Against

^ This figure is no doubt too low. Fernald apparently overstimated the age and the original
peculiarity of the Newfoundland flora. At least the following 30 species, regarded by Fernald
as indigenous, should probably be removed to the group of European introductions:

Alchemilla filicaiilis Gnophalium sylvaticiim P. sylvatica

A. minor Jiincus acntiflorus Polygonum hydropiper

A. pastoralis J. bidbosus P. Rati

Cardamine flexuosa Latliyrus pratensis Potentilla anglica

Cirsium paliistre Litiuni catharcticiim P. erecta

Cochlearia danica Liizida campestris P. sterilis

Festuca capillata L. pallescens Scrophidaria nodosa

Fragaria vesca Moutia rkndaris Sieglingia deciimbens

Galium aparine Nardus stricta Speraularia rubra

G. saxatile Pedicularis paliistris Veronica officinalis

If these species are included, the percentage of European introductions in the flora of
Newfoundland is increased to 23.

* Alopecurus ventricosus Pers., Centaurea nigrescens Willd., Geranium ibericum Cav.,
Lappula echinata Gilib., and possibly 2 or 3 micro-species oi Hieracium. A few other species
grow in the British Islands only as cultivated, escaped, or casual adventives.

The human transport of animals across the Northern Atlantic

147

(Photo the author, 4.VII. 1949.)

FIG. 9. St. Fintans, SW Newfoundland. A patch of dry meadow along the railway
embankment. The vegetation is dominated by Marguerites {Chrysanthemum leuc-
antheryutm L.) and Red Clover {Trifolium pratense L.), both introduced from Europe.

the 14 per cent introduced species of Carabid beetles in the fauna of Newfound-
land (23 species of 165), Nova Scotia (Lindroth, 1954c) possesses 11 per cent
(20 of 186 species) and the total number of European species is there 34 (18 per
cent). The European Carabids are more numerous in Labrador (Lindroth, i954d),
contributing no less than 39 per cent (32 species of 82), but of these only 4 per cent
(3 species) may be regarded as introduced.

It can be stated, without any exaggeration, that Nezvfoundland more than any
other part of North America has received an introduced element of animals and
plants from Europe. It is an important task to explain why.

Another interesting feature is that the introduced European species show an
unequal, in the individual species often disjunct, distribution within the limits of
Newfoundland, with a marked concentration in the southeast, on the Avalon Penin-
sula, the centre of trade and population. On the eastern coast of Avalon 19 in-
troduced species of Carabid beetles have been found, the highest number shown
by any part of North America {vide maps, figs. lo-ii).

148

CHAPTER II

TABLE 3. The European element in the fauna of Newfoundland as illustrated by

some sufficiently known groups of terrestrial (and limnic) animals. Arrangement of

groups from largest to smallest European influence.

Isopoda (Palmen, 1951). . . -

Total

In common with Europe

Total

Different
sbspp.

Identical

Regarc

introc

from E

ed as
uced
urope

12

12 "

100%

12 =

100%

12 =

100%

Diplopoda (Palmen, 1952) . .

18

16-

89%

16 =

89%

i6 =

89%

Chilopoda (Palmen, 1954). . •

12

9

75 %

9 =

75%

9 =

75%

Curculionidae (Leng, 1920;

Brown & Palmen, in litt.) . .

65

35 =

53%

1 = 2%

34 =

52%

29 =

44 %

Scarabaeidae (Landin, in litt.) .

14

6 =

43%

6 =

43%

5 =

36%

Mollusca (Brooks, 1936, 1940)-

581

33 =

57%

1 = 2%

32 =

55%

14^ =

24%

Chrysomelidae (Brown & Pal-

men, in litt.)

30'
15

8 =

27%

1 = 3%

7 =

23 %

6 =

20%

Coccinellidae (Chapin, in litt.) .

7 =

47%

1 = 7%

6 =

40%

3 =

20%

Carabidae (Lindroth, 1955) . .

165

53 =

32%

12 = 7%

41 =

25%

23 =

14%

Elateridae (Becker, in litt.) . .

25

5 =

20%

1 = 4%

4 =

16%

2 =

8%

Araneae (Hackman, 1954). . .

220

54 =

25%

54 =

25%

II =

5%

Macrolepidoptera (Krogerus,

lO^J.)

280

61 =

22%

19 = 7 %

42 =

15%

4 =

I %

Odonata (Walker, I953; Need-

ham & Westfall, 1955; Valle,

locc)

29

5 =

17%

1 = 3%

4 =

14%

■■ VjD/

All groups together

944

285 =

30%

37 = 4 %

259 =

28%

134 =

14%

^ The Lymnaeidae of Brooks' list (1940, p. 75) have been revised according to Hubendick

(1951)-

^ The Brooks' (1936, 1940) are very restrictive in regarding only 6 of the Newfoundland
Molluscs (one, Deroceras reticulatum Mull., even doubtfully) as introduced from Europe.
To my mind, at least the following 8 species should be added: Arion circuniscn'ptus Johnst.,
Cepaea hortensis Miill., Hygrornia striolata Pfeiff., Limax marginatus Miill., L. maximus L.,
Oxychiliis draparnaldi Beck, Vallonia excentrica Sterki, Vertigo alpestris Aid. For Lymnaea
peregra Miill. a transport with birds is perhaps equally probable {vide below, p. 251).

^ The revision of the Newfoundland Chrysomelidae is not finished and the figures here
given are preliminary only.

This is a feature of general validity. In the "Iso-Myriopods" (Palmen, 1951,
1952, 1954) 42 species have been stated in Newfoundland, 37 of which are re-
garded as European introductions, and of these no less than 20 are known from
the Avalon Peninsula only.

In a foregoing section of this chapter (p. 135 a.f.) the criteria of an introduced
species were formulated. Let us now try to test them in practice. The reader, of

The human transport of animals across the Northern Atlantic

149

FIG. 10. "Europeization" of the North American fauna, according to the number of
introduced Carabid beetles.

course, is by no means obliged to accept the verdict "introduced" for any species
without hearing the arguments. Carabid beetles, as usual, may provide the material.

Carabid beetles common to Europe and Newfoundland

t = regarded as introduced from Europe
Agonum bogemanni Gyll. f A. miilleri Hbst.

A. consimile Gyll.
A. mannerheimi Dej.
(different subspecies)

A. qiiadripunctatum DeG.
t A. ruficorne Gze.
A. thoreyi Dej.

15°

CHAPTER II

FIG. II. "Europeization" of
the fauna of Newfoundland,
according to the number of
introduced Carabid beetles.

t Amara aenea DeG.

A. alpina Payk.

(different subspecies)
t A. apricaria Payk.
f A. aidica Panz.
t A. bifrons Gyll.

A. erratica Dft.
t A. familiar is Dft.
t A.fulva DeG.

A. hyperborea Dej.
t A. lunicollis Schio.

A. quenseli Schnh.

A. torrida 111.

t Bembidion bruxellense Wesm.
{rupestre auctt.)

B. grapei Gyll.

t B. lampros Hbst.

B. petrosum Gebl.

(different subspecies)

B. quadrimacidatum L.

(different subspecies)
t B. stephensi Crotch
t B. tetracolum Say (ustulatum auctt.)

B. tramparens Gebl.

Blethisa midtipimctata L.

(difTerent subspecies)

Calathus micropterus Dft.

(different subspecies)
t Carabiis nemoralis Miill.
t Clivina fossor L.

Dyschirius politus Dej.

Elaphrus ri par ins L.
t Harpaliis affinis Schrk. (aeneiis F.)

H. fidiginosus Dft.

H. nigritarsis C. R. Sahib.

(different subspecies)
t H. rufipes DeG. {piibescens Miill.)

Loricera pilicornis F.

Miscodera arctica Payk.

(different subspecies)
f Nebria brevicollis F.

N. gyllenhali Schnh.

(different subspecies)

A''. 7nvalis Payk.

(different subspecies)

Notiophilns aquaticus L.

The human transport of animals across the Northern Atlantic

151

(Photo P. Ardo.)

FIG. 12. Bembidion tetracolum Say {iistulatiim auctt.) in Newfoundland. An introduced
Carabid beetle with "immature" disjunct area, constantly fligthless on this island.

t N. bignttatus F.

Patrobus septentrionis Dej.

(different subspecies)

Pelophila borealis Payk.

(different subspecies)
f Pristonychus terricola Hbst.

Pterostichus adstrictiis Eschz.

P. brevicornis Kby.

(different subspecies)
t P. melanarius 111. (vulgaris auctt.)
t P. streniius Panz.
t Trechus rubens F.

Trichocellus cognattis Gyll.

Species regarded as introduced are 23 in number. This has been decided on
the basis of "the five criteria" (above, p. 135 a.f.), in the following way:

I. On HISTORICAL evidence. Three species, Amara aulica, A. bifrons, and
Harpalus rufipes, easy to collect and now common in the southwest of New-
foundland, were not captured during the intense collecting carried out in this
region in the period 1905-15 by several skilled collectors (Lindroth, 1955a, p. 8).
They are probably the result of later introduction. On the other hand, Amara fulva
and Bembidion bruxellense were taken there by P. G. Bolster in 1905 and 1907,
respectively, the first North American records. Other first-records were made
outside the area: Carabus nemoralis in New Brunswick, 1870; Bembidion stephensi at

152

54'

SCALE OF MILES
25 25 50 75 100

I I I I — I

CHAPTER II

52'

52°

50'

FIG. 13. Clivina fossor L. in Newfoundland. An introduced Cara-
bid beetle with "immature" disjunct area. Both long- and short-
winged specimens occur on this island.

(From Faune de France.)

The human transport of animals across the Northern Atlantic

153

(Photo P. Ardo.)

FIG. 14. Distribution of long-winged (white) and short-winged (black) specimens
of the Carabid beetle Notiophilus biguttatus F. in Newfoundland, its single occurrence
in North America. The area of the circles is in proportion to the number of individuals
investigated.

Short wing is a dominant; consequently, long-winged specimens are homozygotes
and a preponderance of them indicates late colonisation. This species is a European
introduction, spreading from the Avalon Peninsula in the southeast towards the west
and north.

Ottawa, 1891; Clivina fossor at Montreal, 1915. These have since been obviously
enlarging their area and so, with unusual rapidity, has Pterostichus melanariuSy
first found in Nova Scotia in 1926.

2. On GEOGRAPHICAL evidence. Nebria brevicollis, Bembidion lampros, and
Pterostichus strenuus are known in eastern North America each from one quite
small area only, the Nebria on the French island Miquelon, the remaining two on
the eastern shore of the Avalon Peninsula. A pronouncedly "unnatural", disjunct
area in Newfoundland is occupied by Agonum ruficorne, Bembidion tetracolum
(fig. 12), Carabus nemoralis, Clivina fossor (fig. 13), and Trechus rubens. A large-
scale disjunction, suggesting introduction into North America on the whole, is
characteristic of Agonum miilleri, Amara familiaris, Bembidion lampros and tetra-
colum, Carabus nemoralis (fig. 7), Clivina fossor, Harpalus af finis, and Pterostichus

154 CHAPTER II

melanarius, which have an isolated, late occurrence in the Pacific Northwest
(Hatch, 1953).

3. On ECOLOGICAL evidence. The following European species occur in New-
foundland only on ground strongly influenced by human culture, mainly around
ports and other communities: All Amarus of the list above, except fulva (cf. p. 142),
Bembidion lampros and tetracolum, Carabus nemoralis, Clivina fossor, Harpalus
rufipes, and Trechus rubens. Still more pronounced is Pristonychus ferricola, a true
indoor insect.

4. BIOLOGICAL evidence of introduction does not apply to Newfoundland
Carabidae but is valid for several Curculionids (Weevils), especially those confined
to Trifolium as food-plants (Phytonomus meles F., nigrirostris F., punctatus F.,
Sitona lepidus Gyll., &c.), a plant genus not represented by indigenous species
on the island.

5. On TAXONOMic evidence, Notiophihis biguttatus, due to the geographical
distribution of its macro- and brachypterous forms, must be regarded as introduced
(fig. 14). Pterostichus stretmus, in Europe dimorphic with respect to the hind- wings,
has apparently been introduced into Newfoundland in the pure (homozygotic)
brachypterous form, which possibly explains its occurrence only within a restricted
area on the eastcoast of Avalon.

It is also worth while to point out that the characters referred to by Lapouge
(1908, p. 19) in constituting the populations of Carabus nemoralis from Newfound-
land and New Brunswick as belonging to a distinct, indigenous North American
subspecies (also referred to by Okland, 1927, p. 355), are quite imaginary and
provide no facts, which contradict the opinion that this species is a European emi-
grant.

The Newfoundland trade
Historical review

The trade between Newfoundland and Europe is so intimately connected with and
so deeply stained by the peculiar political relations to the mother country. Great
Britain, during more than four centuries, that a brief summary of early New-
foundland history seems appropriate. The most important sources have been
Prowse (1895), Rogers (1911), Newton (1930), Harris (1930), Cochrane (1938),
and Parkinson (1948).

The discovery of Newfoundland by the Europeans took place at a very early
date. It is now regarded as a historical reality that Norsemen starting from Green-

The human transport of animals across the Northern Atlantic 155

land in the year 1000 A.D. reached the coast of North America and even erected
a settlement in "Vinland", though this lasted some few years only. By several
earlier authors Vinland was identified either with Newfoundland or, more likely,
with Nova Scotia or New England {vide Gathorne-Hardy, 1921, p. 224 a.f.) but,
though Newfoundland was no doubt visited by the Vikings, modern authors have
felt inclined to place Vinland farther south— according to Naess (1954), who based
his opinion on the astronomic observations from the journeys mentioned in the
Icelandic "Sagas", not north of Chesapeake Bay. The method of Naess seems, in
spite of its somewhat unexpected result, to be more reliable than that of Lowe
(195 1), who, according to the plants mentioned in the old travel account, located
Vinland somewhere between Maine and Long Island.

It has been suggested by Iversen (1938) that plants such as Sisyrinchium mon-
ianum Greene (angustifolium auctt., nee Mill.) were brought back to Greenland
from North America by the Norsemen and that other species may have been carried
in the opposite direction. Even if this possibility is admitted^ there seems to exist
no case of animal distribution requiring a similar explanation.

The definite incorporation of Newfoundland in the European trade was started
by John Cabot's rediscovery of the island in 1497, only five years after the famous
first voyage of Columbus. Cabot was sent out from Bristol and for more than three
centuries Newfoundland's closest naval connection was with southwestern Eng-
land. Though Cabot had discovered a new land, it was the surrounding sea that
was exploited, not only by English, but in the earlier part of the i6th century
mainly by French and Iberian fishermen. The island itself was used as nothing
but an immense fishing-depot, with ports actively frequented in the summer
months but left practically uninhabited during the winter. From the very beginning
the Avalon Peninsula in the southeast was the predominant centre of this trade.

No wonder that the colonization of Newfoundland went on very slowly. Only
occasionally a few crews may have been left behind over winter for preparatory
work in the ports. The first known serious attempt to establish a settlement was
made by John Guy from Bristol, who in 1610, with his "Company", chose Cupids
on the Conception Bay of Avalon as a perrnanent residence. This small colony
lasted only a few years. A more successful settlement was established at Ferryland
on the eastcoast of Avalon in 1621; this was inhabited, with interruptions, for
more than 50 years.

However, any attempt to colonize the island was met with heavy opposition from
the fishermen and their customers (merchants and shipowners) in the ports of
southwestern England. "From the very beginning of the sixteenth century they

^ This opinion has been opposed by Bocher (1948, p. 19-24). Cf. also p. 250.

156 CHAPTER II

had conducted a profitable business in organizing the annual fishing expeditions
to Newfoundland. They were opposed to any permanent settlement with its
consequent laws and regulations which would interfere with their authority. It
suited them better to send their fleets westward each spring to take possession of
the stages, flakes, and cook-rooms which they had left at the end of the preceding
season, to catch and cure fish there during the summer, then to abandon the place
in the autumn. If there were any settlers, they would occupy the harbours and
coves which the merchants had been accustomed to use and so interfere with their
business. Accordingly they tried hard to prevent any settlement, or at least to
keep it as low as possible" (Cochrane, 1938, p. 56). In 1633, on an influential
petition from the southwestern ports, the Privy Council issued an order, called the
"Western Charter", on which all subsequent regulations concerning Newfound-
land were based for more than a century and a half. Among the rules laid down here
was one giving the jurisdiction of every port to the "Fishing Admiral", that is the
captain of the ship arriving first in the spring. Another, especially directed against
settlement: "All owners of ships trading to Newfoundland forbidden to carry
any persons not of ships Company or such as are to plant or do intend to settle
there." A complementary rule, issued in 1637, deprived settlers the right to live
less than six miles from the shore (!)

These and other obstacles contributed to keep the number of resident settlers
very low for a long time. The approximate figures of Prowse (1895, p. 698 a.f.) give
a total permanent population of Newfoundland during the last quarter of the 17th
century not exceeding 3000 men. In summer they were greatly outnumbered by
the crews of fishing- vessels; already in 1644 the English alone exceeded 10,000
(Prowse, I.e., p. 190). French settlements seem not to have existed before 1662,
at Placentia (westside of Avalon), and about 1670, on St. Pierre, Not until the later
half of the i8th century did the total permanent population of Newfoundland
exceed the seasonal fishermen in number.

The extraordinary conditions prevailing in Newfoundland during a period of
almost three centuries were bound to stamp the trade with the mother country.
The large fishing-fleets leaving the ports of southwestern England every spring
were destined for a poor, almost uninhabited country, a bad market for goods
of any kind. The crews' own supplies gave no full cargo. The ships sailed in
ballast. At the end of the fishing season they returned fully loaded, as a rule not
directly home but to the foremost consumers, the catholic countries in southern
Europe, and thence back to England in the late fall, often likewise in ballast. This
triangular traffic was carried on to an almost unchanged extent as long as sailing-
vessels ruled the sea, that is to the middle of the 19th century. The first steam-line
calling at Newfoundland, connecting it with Halifax, was opened in 1842 (Harris,

The human transport of animals across the Northern Atlantic 157

1930, p. 431). Already at that time several introduced European insects had be-
come established on the island (below, p. 215).

The history of the French islands St. Pierre and Miquelon has much in common
with that of Newfoundland proper. Though already colonized about 1670 by
the French, they later stood under British supremacy during the main part of
two long periods: 1713-63, 1778-1815. I was told by Mr, Mathews, of Poole, that
this Dorset port upheld an intermittent direct trade with St. Pierre, probably
about 1800. From the report of the astronomer Cassini, in 1768 (Prowse, 1895,
p. 570), we learn that at that time the French fishing-fleets arriving at St. Pierre
every spring belonged to many different ports, from Honfleur on the Seine to Bay-
onne on the Bay of Biscay. However, European animals hitherto observed on St.
Pierre-Miquelon are not necessarily introduced from France; the Ground-beetle
Nehria hrevicollis F., the Lamellicorn beetle Aegialia rufa F., and the Weevil
Trachodes hispidus L., unknown elsewhere in North America, may as well have
arrived from the British Isles. Not even the period of prohibition in U.S.A., when
St. Pierre served as a too well-known staple of French spirits, certainly packeted
in nice straw cases, seems to have had particular importance for the introduction
of European animals. This is a confirmation of the view stressed in this chapter,
that transport of ballast is superior to any kind of cargo as an instrument for
synanthropous dispersal.

The ballast-traflfic

There are many records from Newfoundland as well as from England indicating
the regular use of ballast on board the sailing-vessels of the North Atlantic trade.

When, in 161 1, John Guy, the founder of Newfoundland's first permanent
settlement, published "Certaine orders for the ffishermen", he gave them the
form of 8 rules and \h& first of these ran (cited from Prowse, 1895, p. 99): "Ballast
or anything hurtful to Harbours not to be throwne out but be carried ashore—
Penalty £ 5 for every offence".

This instruction was repeated as point 2 of the "Western Charter" of 1633:
"No ballast to be thrown out to prejudice of harbor", and as number i of the
"Lawes, Rules, and Ordinances whereby the Affaires and fishery of Newfound-
land are to be governed untill the Parlam^ shall take further order", of 1653:
"That noe Ballast, Prest stones nor anything else hurtfuU to the Harbours bee
throwne out to the prejudice of said Harbours, but that it be carryed ashore and
layd where it may not doe annoyance".

Even in 171 2 the appointed Governor of Newfoundland, Sir Nicholas Treva-
nion, included, as No. 13, a point of similar content into his "Fishery Scheme".

158

CHAPTER II

Among records given for 1618 by Sir Richard Whitbourne, acting as Commis-
sioner of Vice- Admiralty, in order to illustrate various disorders committed in the
Newfoundland ports, was: "Harbours frequented by English near 40 in number,
almost spoiled by casting out their balast and presse stones into them".

The quotations above show sufficiently that ballast was brought to Newfoundland
in great quantities , at least in the 17th and 18th centuries, and that as a rule it
was delivered on the shore. It is easily understood that this procedure involved
the best imaginable chances for fruits and seeds of plants, as well as for all kinds
of animals associated with the soil, to be carried across the Atlantic.^ This has
already been clearly realized by W. J. Brown (1940, 1950).

Those harbours of Newfoundland in the first line affected by the conditions
just described were of course the oldest ones, which in a remarkable way were
concentrated to the eastern coast (map, fig. 15). Apparently St. John's, the present
capital of the country, predestined by its sheltered position, was the most frequented
port even in the 16th centjry {vide for instance Prowse, 1895, pp. 70,72, 113;
Rogers, 1911, pp. 19, 23, 25, 26) though probably it was not the first place to be
permanently settled. Fishing-trade and settlement were almost exclusively con-
centrated to the Avalon Peninsula, including the first French colony, Placentia, on
its western shore. This lead of the Avalon has been kept ever since and at present
it houses about 45 per cent of the population. In accordance herewith the fauna
and flora of this part of the island contains a larger European element than any
other district in North America. As already mentioned above (p. 147), this is
evident in the case of Carabid beetles (map, fig. 11), of which 19 introduced species
occur on the east coast of Avalon; four of them are confined to this part of the island
[Bembidion lampros Hbst., Pristonychus terricola Hbst., Pterostichus melanarius 111.,
P. streuuus Panz.). In the Iso-Myriapods, 20 introduced species (54 per cent of this
element in Newfoundland) are restricted to the Avalon Peninsula.

Two isolated, early colonized parts of Newfoundland are the twin islands St.
Pierre-Miquelon, settled by the French about 1670 and still in French possession,
and the islands Twillinggate and Fogo in the northeast, colonized about 1700.
Both areas show faunal vestiges of a direct connection with Europe: on Fogo-
Twillinggate the isolated occurrence of the Carabid beetles Bembidion tetracolum
Say {ustulatmn auctt.) (fig. 12) and Clivina fossor L. (fig. 13), both in the short-

^ It seems reasonable to assume that the isolated occurrence of certain European Molluscs
on small islands off the coast of Avalon can be explained by the fact that sailing-vessels
arriving from Europe, especially at low tide, used to discharge their ballast here before
entering the port of destination on the mainland of the Peninsula. Brooks & Brooks (1940,
p. 58 a. f.) regarded these species as old relicts on the small islands in question, a rather
adventurous hypothesis, which does not explain why the species concerned occur in
Europe.

The human transport of animals across the Northern Atlantic

159

FIG. 15. The most frequented
ports in the Newfoundland
trade before the 19th cen-
tury.

Trepossey

winged form, incapable of flight; on St. Pierre-Miquelon the three beetles men-
tioned above (p. 157).

The western parts of Newfoundland, the climatically most favoured, served for
centuries as a base for the French fishery and had no permanent settlement at all.
In the period after 1764 the French were downright forbidden to stay over winter
(Prowse, 1895, p. 318; Newton, 1930, p. 144). Not until 1877 ^^^^ the colonization
of the west coast started by the English and shortly afterwards some French lobster
factories were established, the first one at Port-au-Choix in the northwest (Harris,
1930, p. 673 a.f.). In spite of this the European faunal element is quite pronounced
on the west coast and includes even species [Amara aulica Panz., A.fulva DeG.,
Harpalus rufipes DeG., among Carabid beetles) which are lacking in the east.
Possibly the active trade of modern times with Nova Scotia has contributed to the
Newfoundland fauna some "second-hand" European species, first arrived in the
Halifax region or on Cape Breton Island (the three species mentioned all occur in
Nova Scotia). A good proof of late appearance is, as mentioned above (p. 151), that
Amara aulica and Harpalus rufipes, now widely distributed and abundant, were
not captured by any of the entomologists who made intensive collections in
western Newfoundland between 1905 and 191 5. The former was first taken in
Nova Scotia in 1929 (Fall, 1934), the latter on Prince Edward Island in 1937, in
Nova Scotia in 1938, in New Brunswick in 1939.

l6o CHAPTER II

At least it seems more reasonable to suppose that these species have arrived in
Newfoundland from the adjacent mainland, either carried by ship, or flying
(all three of them are capable of flight), than that they should have been introduced
directly from Europe, for instance to Corner Brook on the west coast.

If the non-resident French fishermen of earlier centuries had been partly res-
ponsible for the introduction of European species into western Newfoundland,
this would be supposed to have resulted in the appearance there of species un-
known elsewhere in the Maritime Provinces of Canada, such as the beetles Nebria
brevicoUis F., Aegialia rufa F., and Trachodes hispidus L., on St. Pierre- Miquelon.

In 195 1 I was told by an old fisherman of St. Pierre that even in later years
sailing-vessels sometimes went in ballast from the west coast of Newfoundland,
at least from the Port-au-Port region oflF St. George's Bay, to St. Pierre. The
isolated occurrence on the French islands (in part also on the Burin Peninsula
situated just opposite) of such westcoast species as the Carabid beetles Amara
avida Say, Harpalus rufipes DeG., and Patrobus longicornis Say, all favoured on
cultivated ground, may possibly be explained thereby.

Some of the earliest introductions in Newfoundland, for instance the ground-
beetles Agonum mulleri Hbst. and Bembidion bruxellense Wesm. {rupestre auctt.),
which are more abundant and widespread there than anywhere else in North
America, may very well have used this island as an accumulator from which waves
of air-borne emigration started and reached the mainland.

In order to understand the eff'ect of the animal transport with man here suggested,
it is also necessary to get a clear idea of where and how the ballast was taken on the
European coast. Both the composition of the introduced faunal element in New-
foundland and the history of the North Atlantic trade have shown that English
ships and English ports have played the foremost role. Unfortunately, published
English records on the ballast -traffic seem to be very scarce. The brief description
given here has therefore been compiled mainly from information generously
supplied by persons living in or closely connected with the southwestern ports:
in Poole by Mr. Edwin F. J. Mathews and the late Mr. Horace P. Smith, respect-
ively present and former borough historian; in the Bideford district by the late
Mr. Vernon C. Boyle of Westward Ho!, who was deeply interested in the matter;
in Barnstaple by Mr. Alfred E. Blackwell, Head Librarian; in Bristol by Miss
Elizabeth Ralph, City Archivist. Especially profitable was the study of Poole
harbour journals, from 181 3 and some years thereafter, liberally put at my disposal
by the present owner, Mr. E. E. Kendall, aged 84, former Harbour Comissioner.

The human transport of animals across the Northern Atlantic

i6i

FIG. 1 6. The fore-
most British ports
of the Newfound-
land trade before
the 19th century.

The British ports regularly mentioned in every treatise on the history of New-
foundland (e.g. by Prowse, 1895), all situated in southwestern England (in Devon
and Cornwall, in addition Poole in Dorset, and Bristol), are marked on the map,
fig. 16. Their trade with Newfoundland goes back to the early i6th century, as
described above, and did not lose its outstanding importance until steamers out-
ranged sailing-vessels towards the end of the last century.

Sometimes the ships went to Portugal or Spain on their way out to load salt for
the fishery but as a rule they sailed directly to Newfoundland. Published records
give very little information concerning to what extent they then used to go in
ballast but Braddick (1953, p. 30) mentions an example from Topsham showing
that in the winter season (Sept. -March) 1763-64 every second ship passing through
from Exeter, with unknown destination, was in ballast. Miss Ralph informs me
that as early as the 17th century a "ballast master" was appointed in Bristol, for
whom special rules, orders, and instructions were established.

Far more detailed records were obtained from the unpublished "Lists of Arrivals
and Sailings for the Port of Poole", placed at my disposal by Mr. Kendall and cover-
ing a short period from and including the year 1813 (fig. 17).

Diagram i summarizes the Newfoundland trade from Poole during two years.
In the first of these, counted from September ist 1813 to August 31st 1814 (re-
cords for the remaining four months of 18 14 being lost), 38 sailing-vessels left
Poole for Newfoundland, 21 of them partly or fully loaded with ballast amounting
to more than 1,100 tons altogether. In 1815 (January ist to December 31st) the
figures were: 57 ships, 17 of them partly or fully loaded with ballast to a total amount

1 1 — 565597 Lindroth

1 62 CHAPTER II

of 1, 1 80 tons. An estimation of the tonnage of British soil exported to Newfound-
land from Poole alone during the course of four centuries would give an imposing
figure indeed!

Poole seems on the whole to have played a very important role, underestimated
by Prowse (1895) and others, in the Newfoundland trade. According to an act
passed in 1778, permitting the export of limited quantities of cereals "for the use
of the fisheries at Newfoundland, Nova Scotia, and Labrador" (Gribble, 1830, p.
607), Poole was allowed to send greater quantities than any other British port.
The maximum intensity of the Poole trade with Newfoundland (according to
Mr. Mathews) lasted from the later decades of the i8th century to about 18 15
{vide also Parkinson, 1948, p. 244). Not until about 1800, did Liverpool catch up
with Poole in importance to the Newfoundland trade (Parkinson, 1948, pp. 243,
248). The direct traffic finished in the i88o's.

The ports visited in the first place by Poole sailors of this time, according to
information from Mr. Mathews, were Brigus, Carbonear and Placentia on the
Avalon Peninsula, Burin in the south. Trinity and Catalina on the Bonavista
Peninsula, and Fogo and Twillinggate in the northeast (fig. 15).

There is no reason to believe that the Newfoundland sailing trade of other south-
western ports differed essentially from that of Poole, but details such as the ballast
procedure have been lost, due to lack of published records. For the concrete pic-
ture gathered from the old harbour papers of Poole, we are entirely obliged to the
piety and foresight of a man, Mr, Kendall, who saved them from the fate of similar
documents of other ports.

From a biological point of view it is also worth observing that, according to the
diagrams i and 2, the export of ballast reached its maximum in March and April,
with a second, less pronounced, in the fall. The spring maximum was no doubt
still more evident before the permanent settlement of Newfoundland was started.
This may have contributed to the selection of the animal and plant species which
managed to cross the Atlantic as stowaways (cf. below, p. 210).

As to the kind of vessels taking part in this kind of transatlantic traffic, Mr.
Boyle informed me that they were usually schooners of about 80 to 120 tons
(figs. 18-19) ^^^ that 24 days would be a good passage for a ship of this kind from
the North Devon coast to Newfoundland.

FIG. 17. A page from "Lists of Arrivals and Sailings for the Port of Poole", April
1 8 14. Of the 9 ships mentioned, all but one are destined to Newfoundland and 7 of
these carried ballast besides the cargo.

The bottom item runs: — "In the John & Mary, [Captain] Collins, for New-
foundland. [Cargo] 60 tons. Ballast 10 [tons]."

Published by kind permission of the owner, Mr. E. E. Kendall, Poole.

The human transport of animals across the Northern Atlantic

i6'

164

CHAPTER II

DIAGR. I. Ships (white) and ballast (black) from Poole, Dorset, destined to New-
foundland during two years, i.ix. 1813-31.VIII. 1814, and 181 5. (The records of
September-December 18 14 are missing.)

The next important task is to find out from where the ballast was taken in the
English ports. The kind of soil or other materials used of course determined the
kind of living organisms which may be supposed to have found their way on board.

As far as Poole is concerned, I was told by Mr. Kendall that the ballast heaps on
the quay, visible in fig. 21, consisted partly of material brought in by ships arriving
in ballast from other ports, for instance from the Iberian Peninsula, and partly
of stones, bricks, mortar, &c., from the town (for instance remnants from demo-
lished houses). A similar case was related from Bideford through Mr. Boyle:
The schooner "Katie", in 1905, took in 20 tons of builders' rubbish (broken bricks,
mortar and stone) in Exeter on her way home to Appledore. It seems justifiable
to regard this as the normal kind of ballast. The simplest way for a vessel to get
ballast before sailing was to have it taken on board from the same wharf where
she had discharged her load and the stuff most usually available there was no
doubt builders' rubbish. For Bristol Miss Ralph speaks of stones and rubble ob-
tained from the banks of River Avon. Later, after 1880, coastal vessels, at least,
were using limestone as ballast, an economical method of carrying lime to the
farmers (according to information from Mr. John Keast, Streatham, London).

A different kind of ballast was regularly used by vessels from Bideford, including
Appledore, and Barnstaple, as described by Mr. Boyle: —"The ballast was al-
ways from the beaches, and sand was liked best. The vessels would be laid on
Sandridge at Appledore and given a list, so that when the tide was out the crew
could jump out and heave the sand up over the side with shovels. About 40 tons
would be enough."— This locality, "The Crow" (figs. 23, 25) north of Appledore,
was investigated in 1954 (below, p. 178).

The human transport of animals across the Northern Atlantic

165

II III IV

VI VII VIII IX

1823

XI XII

DIAGR. 2. Outward ballast from Poole in 1823, regardless of destination. Total
4,298 tons.

Both diagrams (1-2) based on unpublished records from "Lists of Arrivals and
Sailings for the Port of Poole", generously made available by Mr. E. E. Kendall,
former Harbour Comissioner, Poole.

However, the kind of ballast used in the Bideford-Barnstaple district seems to
have been an exception. Above all the ships would not as a rule take it from the
tidal zone because the soil there is almost constantly mixed in with mud, which is
difficult to clean away from the hold afterwards, and also because it is moist,
requiring too much work in proportion to its weight after it has dried out.

A vivid picture of how the ballast was obtained in past days comes out from
interviews made by Mr. Boyle in 1952 with a number of old sea captains from
North Devon, of which the following are cited: —

Mr. Samuel Daniel, sailor, aged 85, says: "In about 1892 I sailed in a new schooner,
the Snowflake, 120 tons, to Newfoundland. We were in ballast. In Runcorn and
Liverpool you cannot get gravel from beaches, so we had stone and earth and rubble
brought in carts from builders' demolitions, or deads from excavations. Another
cargo from Runcorn she took out was coal for Harbour Grace [Avalon Peninsula].
I am not sure but I believe that river mud was never used as ballast in our vessels."

Captain W. J. Slade, Master of wooden schooners and ketches, Bideford: "Some
vessels would sail without ballast, some with little. I once took a cargo of coal to
Court Macsherry in Ireland for a farmer. The onus of getting ballast for me to come
away was on him. He got me some tons from the site of a tumble-down shed^,
stones, broken slates, mortar and rubbish. Then he went off up the lanes and got
a few cartloads from a hedge. It was earth and grass. When I got back here I
tipped it overside in this river."

Captain William Quance, aged 83: "In October 1888 I sailed in the wooden
barquentine Fanny from Appledore to St. John's, Newfoundland, in ballast. The
ship had been built in Barnstaple in 1878. We were loaded with gravel ballast got

^ An excellent opportunity for the indoor Carabid Pristonychus terricola Hbst. to get on
board!

1 66

CHAPTER II

= h.m-

FIG. i8. "Little Minnie", two-masted topsail schooner, built 1866 at Padstow on
the north coast of Cornwall. Finally belonging to J. Stephens of Fowey.

Dimensions: 82.0 ft x 21.2 ft x 10.2 ft. Tonnage: 98 net register.

She was in the Newfoundland trade and was lost in the ice off the island and the
captain was killed, prior to 1899.

Published by due permission of Mr. H. Oliver Hill, through Mr. David R. Mac-
Gregor, London.

from those banks of gravel close inside the Taw-Torridge Bar [i.e. 'The Crow';
vide below, p. 178]. Barges came alongside (30 tons each) [vide fig. 25] to give it
to us. This gravel is in great demand by builders to make mortar, so when we got to
St. John's 23 days later, the gravel was all taken away at the quay in carts."

Captain J. R. Pile, aged 65, coasting trade (not Newfoundland): "When I was a
boy in a ketch we were over at Clonakilty [south of Cork, Ireland] and to get ballast
we had to row about the harbour in the ship's boat and pick up stones on the
beaches."

Mr. Mathews informs me that Irish ports, such as Waterford, Limerick, and
Youghal in Cork, were sometimes called at by Poole vessels on their way to New-

The human transport of animals across the Northern Atlantic

167

-i±4-

>*v«4j }^p^^ J^j^dt,

FIG. 19. A Poole brig, possibly the "Carbonear" (belonging to Harrison, Slade &
Co.), off Henley Island, Chateau Bay, Strait of Belle Isle, southern Labrador. Ships
of this type, about 100 tons, were commonly used in the Newfoundland fishing trade.
Drawn in 1857 by William Gosse of Poole (a brother of Philip Henry Gosse) and
published in his "Sketches of Newfoundland & Labrador" (Ipswich, about 1858).
— Made available by Mr. E. F. J. Mathews, Poole.

foundland. In the i8th century there was also periodically a very important
direct trade between Ireland and Newfoundland (Prowse, 1895, pp. 283, 345) and
ballast-transport westward from Ireland occurred frequently (Parkinson, 1948,
p. 211). Ballast of the varied content described by Captain Slade from Court
Macsherry may therefore very well occasionally have been carried to Newfound-
land.

1 68 CHAPTER II

The triangle traffic, England-Newfoundland-Iberian Peninsula-England, de-
scribed above (p. 156) may also have resulted in the introduction of plants and
animals' with ships from southern Europe returning to England in ballast. Mr.
G. H. Ashe, Colyton, Devon, told me that he suggested the Weevil Sitona
gemellatus Gyll., restricted on the British Isles to southern Devon where it feeds
on Ononis arvensis, to have been introduced in that way. Can this possibly be
applied to the occurrence of another southern Weevil, Otiorrhynchus auropunctatus
Gyll., in Ireland?^ It would be wise not to neglect this point of view altogether,
when defining the so-called "Lusitanian element" of the British fauna and flora.

Application to other parts of North America

The conditions of trade and the use of ballast may have developed in a similar
way in other parts of the Maritime Provinces of Canada and especially in Nova
Scotia, the history of which in many points is similar to that of Newfoundland.
The permanent colonization of Nova Scotia was started in the 1630's, in the be-
ginning mainly by the French. The English did not become predominant until more
than a century later and the last French colony, Louisburg on Cape Breton Island,
was surrendered in 1758. The first settlement at Halifax (British) took place in

1749-

Nova Scotia was frequented by British vessels of very much the same pro-
venience as Newfoundland, originating from the southwest of England. For
instance, as Mr. Mathews informed me, there was periodically an intense but
publicly underestimated traffic between Poole and Nova Scotia.

In connection with the discovery of the European weevil Barynotus sqiiamosus
Germ, (schonherri Zett.), Harrington (1891, p. 22) mentions the ballast heaps at
Sidney, Cape Breton Island, "formed by vessels discharging their ballast of stone,
earth, etc., before loading coal, and many species of introduced plants are found
on, or about them." The ground-beetle Carabus granulatus hibernicus Lth., re-
stricted in North America to Nova Scotia and New Brunswick, has no doubt
arrived with ballast taken in Ireland. This and Bembidion properans Steph. are
the only European introductions among Carabid beetles in Nova Scotia, unknown
in Newfoundland (table 2). Among Weevils (Curculionids) the same applies to
Barynotus moerens F., Otiorrhynchus scaber L., Tropiphorus obtusus Bonsd. and
tomentosus Mrsh., and Tychius picirostris F.

^ Carpenter (1895, p. 215) writes about Otiorrhynchus auropunctatus: "It is certainly
remarkable that so comparatively large an insect should have been overlooked by the older
naturalists; not a specimen is to be found in the collection of that prince of Irish entomolo-
gists, the late A. H. Haliday." The species is not parthenogenetic; the late Mr. E. O'Mahony
informed me that he had seen mating couples several times.

The human transport of animals across the Northern Atlantic 169

Bideford (incl. Appledore) and Barnstaple were connected by direct trade also
with Prince Edward Island, especially in the period from 1840 to 1890, according
to Mr. Boyle. It is worth mentioning, then, that the introduced Carabid beetle
Harpalus rufipes DeG. was first found on Prince Edward Island in 1937, and is
now rapidly spreading.

Brown (1950, p. 197) has drawn attention to the fact that during the Napoleonic
Wars British ships were seeking timber at ports in the Maritime Provinces of
Canada and that "large quantities of ballast" were landed when they arrived there.
As reported by Fowler (1901) St. Andrew's in New Brunswick seems to have been
an important centre for this export of lumber and, consequently, "probably no
locality of equal area in Canada can boast of a larger percentage of foreign plants
in its Flora than that which flourishes on the streets and in the neighborhood of
St. Andrew's".

The ground-beetles Carahus granulatus L. and nemoralis Miill., and the click-
beetle Agriotes sputator L. were first observed in North America in New Bruns-
wick (Brown, 1940).

New England served as a gateway for European introductions in a more restricted
way {vide map, fig. 10). The first colonization and early history of this region
were largely different. As early as the beginning of the 17th century it had the
character of a permanent settlement and, though England tried to monopolize
the trade, this was always more diversified than in the Canadian East and gained
full independence in 1776. It is therefore easily understood that in part other ani-
mals and plants were introduced into northeastern U.S.A. For instance, the ground-
beetles Carabus auratus L. and cancellatus 111., of which at least the first mentioned
has become established in New England, are not permanent inhabitants of the
British Isles and must have arrived from the European mainland; likewise the
Centiped Pachymerium ferrugineum C. L. Koch, widely distributed in Europe
but not recorded from the British Isles.

Similar, more numerous instances are to be found among plant-feeding insects^
apparently introduced with their hosts, such as the following Weevils (Curcu-
lionidae), introduced into the northeastern United States but unknown in Canada:
Elleschus scanicus Payk., Gymnetron netum Germ., Polydrosus impressifrons Gyll.,
Stomodes gyrosicollis Boh., all lacking in the British Isles. One species at least, the
Alfalfa Weevil {Phytonomus posticus Gyll.), has been imported directly to the Middle
West, to Utah, possibly "in the straw packing about fragile imported packages"
(Howard, 1930, p. 115), from where it is spreading rapidly in all directions (fig. 6).

Apparently, due to the early permanent colonization, there was little need for
ballast in ships destined to New England, which in itself was a good market for
all kinds of European products. However, the Lamellicorn beetle Aegialia arenaria

lyo CHAPTER II

F., inhabiting the sand-dunes of the coast and known in North America only from
Massachusetts (DarUngton, 1927), was no doubt a ballast passenger. It was found
on "The Crow", one of the investigated ballast-places of North Devon (below,
p. 178).

All kinds of cattle had to be imported from Europe. It requires only moderate
imagination to guess how slow was the procedure of loading these animals on board
the old sailing-vessels. Meanwhile, of course, various dung-feeding insects were
flying on board and as the rich supply of suitable food was perpetually renewed,
they had no reason to leave the ship during the voyage. It is anyhow almost
surprising what a rich assortment of for instance European dung-beetles, genus
Aphodius, was carried across: 14 of the 41 species occurring in the British Isles
are now well established in the United States, 10 of them also in Canada. All
3 British species of the likewise dung-feeding Hydrophilid genus Sphaeridium were
also introduced into North America, both on the Atlantic and the Pacific coast
(Brown, 1940, p. 70-71; Hatch, 1946, p. 78).

A third important reception area for European introductions was the Pacific
Northwest. It seems that many of the introduced species of that region arrived
comparatively late {vide list of beetles, Hatch, 1953, p. 25-29) and probably in
part as secondary adventives from original centres of introduction in eastern North
America. This may also be true in cases where, to our present knowledge, the
Middle West constitutes an interruption of the area of a species. Even if it is actually
lacking there, the climatic conditions alone may be responsible.

There is, however, quite a series of European species, notably among insects,
occurring in North America exclusively on the Pacific side of the continent, for
instance the following Coleoptera:

Carabidae:

Acupalpus meridianus L. * Calathus fuscipes Gze.

* Amara anthobia Villa * Pristonychus complanatus Dej.

* Anisodactylus binotatus F. Tachys parvulus Dej.
Bradycellus harpalinus Serv. * Trechus obtusus Er.

Curculionidae:
Ceuthorrhynchus assimilis Payk. Otiorrhynchus jneridionalis Gyll.

Otiorrhynchus cribricollis Gyll. Sitona Uneatus L.

* = taken in greenhouses (partly or exclusively).

For these a direct importation from Europe must be assumed (except for
Otiorrhynchus cribricollis which may have arrived from its secondary centre in
Australia, and perhaps the almost cosmopolitan Pristonychus). The late arrival of
most European species into the Pacific Northwest, compared with eastern U.S.A.,

The human transport of animals across the Northern Atlantic 171

is at least partly due to the fact that the Panama Canal was not opened until
1914.

Hatch (1949) has made intense investigations of the greenhouse fauna in Wash-
ington, Oregon, and British Columbia, and found it surprisingly rich. Not only
the obligatory terrestial Isopods (Wood-lice) but also insects, especially Coleoptera,
were strongly represented, in part by species not regularly confined to this kind
of habitat, among these several Europeans, as marked in the list above. Hatch
(I.e., p. 162) is certainly right in his conclusion: "To the extent to which green-
house species are introduced, it is probable that the shipment of nursery stock and
other plant materials have played an important if not exclusive role in their
dispersal. And some of these species may first have found their way into this
country through the medium of greenhouse shipments" (cf. also below, p. 216).
This applies also to a surprisingly large number of European indoor Spiders
(genus Theridium and others) found exclusively in the Pacific Northwest [vide list,
p. 94 a.f.).

However, there are also other European introductions in the Northwest,
seemingly without any connection with greenhouses. Most of them occur also in
northeastern North America (for examples, vide p. 138) and are there regarded
as having arrived with ballast. Some of them may have reached the Pacific coast
by subsequent transcontinental dispersal, either actively or with man, but others
apparently are completely isolated in the Northwest. It seems reasonable to sup-
pose that at least the majority of them has reached also this region by ballast
transport and probably, in spite of the distance, directly from Europe.

In order to get an idea of to what extent ballast was used in this trade, I contacted
the officials of the foremost ports of the Pacific Northwest and recieved two answers
of considerable interest.

The first was from Tacoma, Wash., forwarded by Miss Elfriede Gudelius of
the Public Library, indicating that in sailing ship days, when big square-riggers
arrived on Puget Sound from foreign ports to load wheat and lumber, they were
often in ballast.

The second reply was from Portland, Oreg., and sent by the General Manager
of the port, Mr. J. J. Winn, Jr., who, referring to a letter from his predecessor in
office, quotes as follows:

"In the old days, ending with World War I, sail vessels came in ballast to load
wheat (mostly) for Europe. They all discharged earthen ballast— usually rocky
stuff or sand which the stevedore companies removed and dumped into low lands
abutting on the river. For many years the 'ballast' dock was in the vicinity of the
present Terminal I; later it was in the Linton area in the vicinity of the present
Gasco plant, as I recall."

172 CHAPTER II

It thus seems clear that an introduction of foreign plants and lower animals in
ballast may have played a considerable role also on the Pacific coast of North
America. Apparently in the same way, the Carabid beetle Colpodes buchanani Hope
was introduced into Oregon from SE Asia (Malkin & Hatch, 1953) and it is per-
haps only surprising that countries facing the Pacific, especially on corresponding
latitudes of E Asia, have contributed so little to this immigration. Probably ships
arriving from there used to carry full load.

Animals of a pronounced synanthropic type may of course have been carried
to the Pacific coast already with the earlier settlers, as exemplified by Essig
(1934)-

Investigations on ballast-places in Southwestern England

Several facts mentioned in the foregoing part of this chapter, in animal distribu-
tion as well as in the history of the Atlantic trade, suggest the British Isles and
above all the Southwest of England as the main region of departure for animals
which have been unintentionally introduced from Europe into North America.
It was therefore quite natural that I should wish to make a direct field study of
the insect fauna around the old ports of this region, especially on the ballast-
places, if these could still be located. These investigations were carried out in 1954,
between May 21st and June ist. On April 7th the same year I had already had the
opportunity of a few hours collecting on the Avon banks in Bristol.

Most of the English ports mentioned in the old records of the Newfoundland
trade {vide Prowse, 1895, pp. 37, 40, 56, 81-82, &c.) are situated from Poole,
Dorset, in the south, along the southwestern peninsula to Bristol on the west coast,
the majority in Devon (map, fig. 16). I visited the following: Poole, Topsham,
Dartmouth, Plymouth, Bideford (Appledore), Barnstaple (incl. Fremington), and
Bristol.

By means of preceding correspondence with initiate persons in every port and
inquiries on the spot after my arrival, I tried to find out the exact position of the
place where in old days sailing-vessels used to take on ballast, and in this respect,
thanks to most generous information from Mr. Mathews and the late Mr. Boyle,
both mentioned above, was especially successful at Poole and Bideford. In case
no exact information was available, the collecting was made as close to the oldest
parts of tlie port as possible, or on spots which for other reasons seemed most
likely to have been used as ballast-places.

The terricolous fauna was the main object of collecting and I tried to obtain
complete lists of the following groups of animals: Coleoptera, Hemiptera Heteroptera;

The human transport of animals across the Northern Atlantic 173

Araneae; Chilopoda, Diplopoda, Oniscoidea {Isopoda terrestriaY; and shell-bearing
Mollusca. Quantitative figures were pursued only for Carahid beetles; of these
every observed specimen was captured. The predominating or otherwise charac-
teristic plants were noted in order to describe the nature of the locality in a simple
way.

The identification of animal and plant species was carried out by myself, with
the exceptions accounted for below.

Account of collecting

I. Poole (Dorset), 21-23.V. 54.— The actual site of the old ballast-place is still
known and, according to the late Borough Historian, Mr. H. P. Smith, was
marked on older maps as "Ballast Quay". It was situated on the eastern tip of the
Hamworthy Peninsula (map, fig. 20) and the ballast-heaps were reproduced on an
engraving from the year 1833 (fig. 21), generously put at my disposal by Mr.
Edwin F. J. Mathews, the present Borough Historian. The intensity of ballast
transport from Poole to Newfoundland has already been described (p. 162 a.f.).
The actual spot has now become industrialized and built over with wharfs and
stores but there is a small open patch of grassland left, less than 100 metres south
of the previous ballast-heaps, and here, as well as on similar ground inside the
new power station, collecting was carried out.

a— Tip of Hamworthy Peninsula, immediately inside the concrete quay. Hard,
dry, open soil (artificial filling of gravel and clay, with pebbles and pieces of brick).
Vegetation (according to Mr. Mathews similar to that of the true Ballast Quay
during the first decade of this century):

Bellis perennis Sarothamnus (Cytisus) scoparitis

Cirsiiim arvense Trifolium campestre (prociimbens)

Dactylis glomerata T. pratense

Lotus corniculatus Ulex europaeus

Poa pratensis Vicia angustifolia

Plantago coronopus V. cracca

P. lanceolata Vulpia bromoides

P. media

The collected animal species (vide list, p. 186 a.f.), very few in number due to
the extreme dryness of soil, were distributed among the following groups (num-
ber of "emigrant" species^ in brackets):

^ For Chilopoda, Diplopoda, and Oniscoidea, the collective name " Iso-Myriapoda" has
been used.

^ "Emigrants" are here understood to mean only those species which are regarded as
European introductions in North America.

CHAPTER ir

^

FIG. 20. The old central part of
Poole. Cross = old ballast place,
a-c = author's collecting places.

Chilopoda 2 (2) species

Diplopoda o »

Oniscoidea 2 (2) »

Mollusca 2 (i) »

Specially considered groups i; 14 species, 8 "emigrants" = = 57 per cent.

Coleoptera
Carabidae
Ciirculionidae

Araneae

I'j (6) species

8 (3) »
o »

3 (o) »

b— Inside the power plant on the northern side of the Hamworthy Peninsula,
about 800 metres from the ballast-place (map, fig. 20). Artificial, open soil, of
the same kind as in the preceding locality but partly not so dry, in spots bare.
Vegetation:

Achillea millefolium

Anisantha (Bromiis) sterilis

Artemisia vulgaris

Ballota nigra

Calystegia (Convolvulus) sepium

Capsella hursa-pastoris (coll.)

Cerastium vulgatum

Cirsium arvense

C. vulgare (lanceolatum)

Coronopus didymus

Epilobium hirsutum

Equisetum arvense
Geranium dissectum
Holctis lanatus
Hordeum murinum
Hypochaeris radicata
Poa annua
P. trivialis
Plantago lanceolata
P. major

Ranunculus repens
Rtimex crispus

^ "Specially considered groups" here and in the following include: Carabidae, Cur-
culionidae, Dermaptera, Opilionida, "Iso-Myriapodn" and Mollusca.

FIG. 21. The Port of Poole in the year of 1833, seaside view.

Artist J. M. Gilbert. On stone by L. Haghe, Day & Haghe Lithographers. Pub-
lished by R. A. Grove, Lymington, Hants. — Inscription: "To the Merchants,
Shipowners & Inhabitants of Poole, This View of the Harbour, is most Respectfully
inscribed, by their obedient servant, Rich*^. And^Y. Grove." (Copy received by the
courtesy of Mr. E. F. J. Mathews, Poole.)

The arrows indicate two ballast heaps.

The human transport of animals across the Northern Atlantic

^IS

iy6 CHAPTER II

Senecio jacobaea T. pratense

S. vulgaris T. repens

Sisymbrium officinale Tussilago farfara

Trifolitim hybridum Vicia angustifolia

In addition Alopecurus sp., Crepis sp. (possibly capillaris), and Taraxacum sp.
In spots the cosmopolitan moss Funaria hygromeirica (det. Dr. R. Tuomikoski).
In a moist ditch Juncus effusus.

Collected animal species ("emigrants" in brackets):

Coleoptera

79 (27)

species

Araneae

14

(3) species

Carabidae

29 (13)

»

Opilionida

2

(i) »

Curculionidae

10 (3)

»

Iso-Myriapoda

8

(8) »

Hem. Heteroptera

7 (0)

»

Mollusca

4

(2) »

Dermaptera

I (i)

»

Specially considered groups: 54 species, 28 "emigrants" = 52 per cent.

c— Along a small salt-water pond connected with the sea by a ditch, situated
inside the power plant (map, fig. 20). Clayish soil, bare and covered by a carpet of
green algae close to the water, higher up with:

Artemisia vulgaris Matricaria maritima maritima

Hordeum murinum Spergularia salina

and Atriplex sp. (sterile).

Animals collected in the litoral zone (obvious salinity):

Coleoptera 8 species Hem. Heteroptera 2 species

Carabidae 5 » Isopoda i

None of these occur in North America.

II. Topsham (S. Devon), 25.V. 54.— This small town was formerly an important
harbour for the transatlantic trade. Even ships sailing from Exeter, through the
canal, were supplied with ballast at Topsham (Braddick, 1953, p. 30), especially
in the period 1760-1870 (according to Major A.B. Gay, of the R. Albert Mem.
Museum, Exeter). The actual site of the place where ballast was taken seems no
longer to be remembered. I collected on the inside of the quays and banks on the
eastern side of the mouth of the canalized River Exe, within the limits as well as
immediately north of the town. Hard, dry, open soil (mainly artificial: coke,
bricks, gravel, &c.). Vegetation, depressed and mostly thin:

The human transport of animals across the Northern Atlantic

177

Capsella biirsa-pastoris
Matricaria matricarioides
Plantago major
Pea annua
Ranunculus repens

Senecio vulgaris
Sisymbrium officinale
Stellaria media
Trifolium repens
&c.

Collected animal species:

Coleoptera

Carabidae^

Curculionidae
Hem. Heteroptera

Specially considered groups: 31 species, 18 "emigrants" = 58 per cent.

34 (16) species

Dermaptera

I (i) species

22 (10) »

Araneae

4 (0) »

I (i) »

Iso-Myriapoda

5 (5)

2 (0) »

Mollusca

2 (i) .>

III. Dartmouth (S. Devon), 27. V. 54.— Mr. Percy Russel, Hon. Curator of the
Dartmouth Borough Museum, called my attention to the place still marked
"Ballast Cove" on the new Ordnance Survey map (sheet 20/85). It is situated on
the eastern (Kingswear) side of the harbour, opposite the Royal Naval College,
and consists of a small, shallow salt-water bay, now separated from the shore by
the railway. By this means the shape of the cove has certainly been changed con-
siderably and so have the original flora and fauna of its shore, now stony and barren.
The latter consisted only of the Ground-beetle Agonum ruficorne Gze. and a
Bristle-tail, Petrobius sp. (immature). The railway embankment, with accompany-
ing quays, has a synanthropous vegetation of the usual weed-type, and an utterly
scarce fauna:

Coleoptera 13 (6) species Dermaptera

Carabidae 4 (2) ■ » Araneae

Curculionidae o » Iso-Myriapoda

Hem. Heteroptera 5 (o) » Mollusca

(i) species

(o)

(4)

(2)

Specially considered groups: 12 species, 9
is unreliable because of the poor material.

'emigrants" = 75 per cent. The figure

IV. Plymouth (S. Devon), 26.V. 54.— No records could be obtained of the places
where ballast was taken. On the advice of Mr. A. A. Cumming, Curator of the
City Museum & Art Gallery, I investigated the eastern side of the older (eastern)
of the two harbour basins, especially along the quays at its entrance, which has
not yet become fully industrialized but is largely covered with grass and weeds.
The soil is hard, dry and stony, mixed in with coke, &c.
Vegetation (mostly depressed, but more or less continuous):

^ In addition, Bembidion concinnum Steph., abundant in the tidal zone.

12 — 565597 Lindroth

178

CHAPTER II

Brassica nigra

Potentilla reptans

Dactylis glomerata (dom

.)

Reseda luteola

Daiicus^carota

Senecio vulgaris

Festiica ovina

Trifolium campestre

Kentranthus ruber

T. repens

Lotus corniculatus

Urtica dioica

Plantago lanceolata

In addition Crepis sp

• (P

ossibly capillaris) and Taraxacum sp.

Collected animals:

Coleoptera

17

(6)

species

Araneae

6 (2) species

Carabidae

10

(4)

»

Opilionida

I (i) »

Curculionidae

I

(0

))

Iso- Myriapoda

6 (6) »

Hem. Heteroptera

3

(0)

»

Mollusca

5 (2) »

Dermaptera

I

(I)

»

Specially considered

groups:

24 species, 15 "emigrants" = 63

per cent.

V. Appledore, pr. Bideford (N. Devon), 29-30. V. 54.— Concerning this place,
I obtained extremely valuable information from the late Mr. Vernon C. Boyle, of
Westward Ho!, a well-known expert of the old North Devon trade {vide above,
p. 160 a.f.). He told me that on leaving the estuaries of the rivers Torridge (Bide-
ford area) and Taw (Barnstaple area), sailing-vessels took on board ballast mainly
from the sandridge situated on the confluence of the two rivers' mouth, straight
N. of Appledore, and generally known as "The Crow". The ballast thus consisted
of sand, which was either directly heaved up with shovels at low tide or brought
alongside in small barges. The normal rise and fall of tide here is 20 feet.

a— "The Crow" (map, fig. 22; figs. 23, 25). The collecting was carried out on the
very tip of the peninsula, the only part accessible at any tide. The soil is pure sand,
in the upper tidal zone partly mixed with gravel. Higher up a pronounced chain
of low sand-dunes bound by Ammophila and, on their inside, a depressed vegeta-
tion, continuous only in spots:

Euphorbia paralias
E. portlandica
Ononis repens repens
Senecio jacobaea

Ammophila arenaria
Anagallis arvensis
Cirsium arvense
Cynoglossum officinale
Er odium cicutarium

Collected animals:

Coleoptera

34 (12) species

Araneae

12 (2) species

Carabidae

12 (4) »

Opilionida

I (i) »

Curculionidae

6 (I) »

Mollusca

I (0) »

Hem. Heteroptera

5 (0) »

Specially considered

groups: 20 species, 6 "emigrants"

= 30 per cent.

The human transport of animals across the Northern Atlantic

179

FIG. 22. The Bideford-Barnstaple district of North Devon.

Crosses indicate the author's collecting places; "V a" is "The Crow".

The broken line is the deepest channel of the rivers Taw and Torridge used by
ships.

Of particular interest are the Lamellicorn beetle Aegialia arenaria F. and the
weevil Philopedon plagiatus Schall. (fig. 26). The former has been introduced into
Massachusetts (Darlington, 1927), the latter on several localities in the Maritime
Provinces of Canada (Brown, 1940, p. 76; 1950, p. 202).

In the tidal zone north of the tip, on the east side of the peninsula, the soil was
heavily mixed with mud. The vegetation consisted of scattered Salicornia sp. and
thin crusts of green algae.

Recorded Coleoptera, all extremely abundant:

Bembidion (Cillemis) laterale Sam.
Dichirotrichus piibescens Payk.

Bledius spectabilis Kr.

None of these occurs in North America.

b — The quarry immediately south of Appledore close to the western bank
of River Torridge (fig. 24). Captain Schiller of Appledore, aged 75, told me that
sailing-vessels took gravel and stones as ballast from this place. The bottom of the
quarry, where the collecting was carried out, is a dry, hard, stony plane, with
scattered loose heaps of stones and gravels. The vegetation was discontinuous,
rich in species, and had a pronouncedly synanthropous character:

CHAPTER II

w-isippan»^nptll|[ii-

(Photo the author 30.V. 1954.)

FIG. 23. The Taw-Torridge Bar, called "The Crow", viewed from the south (Apple-
dore side) at low tide.

On the sand-flat (the same as in fig. 25) ballast was commonly taken by sailing
vessels in olden days.

Achillea millefolium

Alliaria petiolata

Bellis perennis

Carduus crispiis

Cerastium vulgatum

Chaerophyllum temulum

Chrysanthemum leucanthemum

Cirsium arvense

C. vulgare {lanceolatum)

Dactylis glomerata

Galium aparine

Geranium dissectum

G. robertianum

Holcus lanatus

Hypochaeris radicata

Lolium perenne

Lotus cornicidatus

Matricaria marititna inodora

Melilotus altissima

Myosotis arvensis

Plantago media

P. 7najor

Poa annua

P. trivialis

Ranunculus repens

Riimex crispus

Senecio jacobaea

Silene cucubalus (vtdgaris)

Solanum dulcamara

Sonchus oleraceus

Trifolium campestre (procumbens)

T. pratense

T. repens

Ulex europaeus

Urtica dioica

Vicia angustifolia

The human transport of animals across the Northern Atlantic

i8i

(Photo the author 30. V. 1954.)

FIG. 24. The quarry south of Appledore (locahty V b) seen from the river-side (River
Torridge).

Stones and earth from the quarry were said to have been used as ship's ballast.

In addition Agrostis ? stolonifera (sterile), Crepis sp., Rubus "plicatus", and
Scrophularia sp.

Collected animals:

Coleoptera

95 (29)

species

Araneae

25 (2) species

Carabidae

32 (11)

»

Opilionida

I (i) »

Curculionidae

15 (4)

»

Iso-Myriapoda

16 (13) »

Hem. Heteroptera

II (0)

»

Mollusca

10 (2) »

Dermaptera

I (I)

»

Specially considered groups: 75 species, 32 "emigrants" = 43 per cent.

Some of the species recorded above (including Stomis, Simplocaria semistriata,
Sitona lepidus, and those living on Ulex) were collected on a slightly moister
pasture between the quarry and the river bank, with Trifolium repens as dominating
plant.

c— Along a salt-water ditch inside the stone wall which follows the river
bank immediately below the quarry (loc. b). Muddy, soft marsh-soil, with Beta

The human transport of animals across the Northern Atlantic 183

vulgaris maritima, Glaux maritima, Puccinellia maritima (dom.), Triglochin
maritimum, &c.

Collected animals:

Coleoptera 9 species

Carabidae 4 » No "emigrants".

Via. Fremington, pr. Barnstaple (N. Devon), i.VI. 54 (map, fig. 22).— This
little port, situated about 2 miles west of Barnstaple, was selected on the advice
of Mr. Alfred E. Blackwell, Curator of the North Devon Athenaeum, his reason
being that this is the only place along the Taw estuary where in- and outgoing
ships to and from Barnstaple come close to land (cf. fig. 22) and thus would be
easily accessible to vessels requiring ballast. The collecting was carried out along
the quay, on dry and hard, artificially fiUed-out soil (mainly coke). Dominating
plants were the three Trifolium species, campestre (procumbens), pratense, and
repens.

Collected animals:

Coleoptera

49 (18)

species

Araneae

8 (i) species

Carabidae

15 (7)

»

Opilionida

I (i) »

Curcidionidae

13 (6)

»

Jso-Myriapoda

8 (8) »

Hem. Heteroptera

6 (0)

»

Mollusca

8 (3) »

Dermaptera

I (I)

»

Specially considered

groups: 46

species, 26 "emigrants"

=- 57 pei

cent.

b— Muddlebridge, east of Fremington. On saline clayish soil around upper tidal
limit, at the bottom of a small estuary. Among bunches of straw and leaf in dense
vegetation of Phragmites communis. The Carabid beetle Pogonus chalceus Mrsh.
under Halimione (Obione) pedunculata.

Collected animals:

Coleoptera 16 (2) species

Carabidae 1 1 (2) »

FIG. 25. Barges taking sand and gravel at low tide on "The Crow", north of Appledore
(visible in the background to the left), N. Devon.

In old days the stuff was then shovelled on board the larger vessels used in the
transatlantic fishing trade. At the time of the photo (about 1900), the gravel was
principally used for concrete work.

Photo taken by a Mr. Fox and published by permission of the National Maritime
Museum, London, through Mr. David R. MacGregor.

184 CHAPTER II

VII. Barnstaple (N. Devon), 31.V, 54. —According to Gribble (1830, p. 547),
"the exports [from Barnstaple], the earliest statement of which refers to 1742,
were pfincipally to Newfoundland". Actually, Barnstaple ships started the New-
foundland fishing trade much earlier; there is a record from 1593 (I.e., p. 622).
It is possible that ballast, when used by vessels leaving this port, was taken on
"The Crow" off Appledore {vide pp. 164, 178) and no records of a certain ballast-
place within the limits of the town are available. On the other hand, the woollen
goods &c. which seem to have constituted the main cargo for Newfoundland may
occasionally have contained seeds, living insects &c. and there is reason to believe
that ballast was sometimes taken on board even in Barnstaple proper.

On the advice of Mr. Blackwell, I chose as collecting-place the northern bank
of River Taw, immediately west of the mouth of the small River Ye, thus in the
westernmost part of the town. This is the part of the harbour where land is most
easily accessible to larger vessels, even at low tide. The selected place is situated
inside the quay and the railway embankment and is partly cultivated (allotment-
gardens). The vegetation otherwise consists of weeds, with Poa annua as completely
dominating plant.

Collected animals:

Coleoptera 56 (29) species Araneae 5 (o) species

Carabidae 25 (13) » Opilionida i (i) »

Curculionidae 3 (3) » Iso-Myriapoda 8 (6) »

Hem. Heteroptera 3 (o) » Mollusca 7 (3) *

Dermaptera i (i) »

Specially considered groups: 45 species, 27 "emigrants" = 60 per cent.

VIII. Bristol (Somerset), 6. IV. 54.— According to information kindly given by
Miss Elizabeth Ralph, City Archivist of the Council House, a "ballast master"
was appointed in Bristol from about the 17th century and the orders to be observed
by him included the establishment of a "ballast wharf" where ships might collect
and deposit ballast. The exact site of this wharf is not known but Mr. Walter
Minchinton suggests, in a letter to Miss Ralph, that the ballast came "from the
neighbourhood of the Avon Gorge and the Downs", that is from the banks of
River Avon in the northwestern part of the town, possibly from the quarries dug in
the steep rock.

A short collecting trip, in company with Mr. P. Ardo, was made on the left
(Somerset) side of the Avon Gorge on both sides of the CHfton Suspension Bridge,
around upper tidal limit, thus in more or less saline localities.^ The vegetation

^ Several species collected at Bristol are no true inhabitants of saline localities (for instance
Amara aenea DeG. and Agomim dorsale Pont, among Carabid beetles, many of the Staphy-

The human transport of animals across the Northern Atlantic

185

(Photo P. Ardo.)

FIG. 26. The Dung beetle Aegialia arenaria F. and the Weevil Philopedon {Cneor-
rhinus) plagiatiis Schall., both occurring on "The Crow" (fig. 23) and introduced into
North America. These sluggish, flightless insects may easily be swept down by
heavy wind on to the tidal flat where ballast was taken (fig. 25). Swedish specimens.

was not noted. Due to the early season the results were poor, for instance no spiders
were found.

Collected animals:

Coleoptera

Carabidae

Ciircidionidae
Hem. Heteroptera

27 (7) species Dermaptera

12 (3) » Iso-Myriapoda

1 (i) » Molhisca

2 (o) »

I (i) species

7 (5) »
6 (4) .)

Specially considered groups: 27 species, 14 "emigrants" = 52 per cent.

linids, &c.). Apparently due to the early time of year, they had not managed to make their
definite selection of habitat after hibernation.

i86

CHAPTER II

It should also be mentioned that, according to specimens preserved at the Bristol
Museum, the ground-beetles Clivma collaris Hbst. and fossor L. have been taken
on the Avon banks. Both of them are European introductions into North America
but they were not found by rne on the investigated ballast-places.

In the fauna of the investigated, stated or supposed, ballast-places, the importance
of species which have been carried over to North America, is quite obvious. Within
the specially considered groups {Carahid and Curculionid beetles, Dermaptera, Op-
ilionida, " Iso-Myriapoda" , and Mollusca), the said element amounts to between
43 and 75 per cent of the species on waste ground. On pure sand and on saline
localities it is less pronounced or even absent.

Lists of animals collected on the ballast-places in SW England

Genera and species in alphabetical order. — Locality numbers as given above
(p. 173 a.f.). — Abundance recorded for common species, (c), only,
f = occurring in North America, ff = in Newfoundland, and regarded as introduced.
*= ,, ,, ,, ,, ** = ,, ,, , and regarded as indigenous.

These symbols are placed in brackets in case only old records from North America
(not later than Leng, 1920) are available.

Coleoptera

Carabidae
83 species vide table 5 (p. 193).

Hydrophilidae
Helophorus brevipalpis Bed. Vc.
t Megasterniim boletophagum Mrsh.
Vb.

Ochthebius impressicollis Cast. (det.
V. Hansen) Vc.

O. marimis Payk. (det. V. Hansen)
Vc
ft Sphaeridium scarabaeoides L. Va.

Silphidae
Ablattaria laevigata F. Via, VII.
Silpha tristis 111. Va, b. Via.

Staphylinidae
t Aleochara bipustidata L. la.

tt Amischa analis Gr. Vb, Via.

A. decipiens Sharp lb, V b, VII.

Astenus longelytratus Palm {angus-

tatus auctt., nee Payk.) lb.

Astilbus canalicidatus F. Vc, Via,

VII, VIII.

Atheta (Dinaraea) angustida Gyll.

(contr. L. Brundin) Vb.

A. {Dimetrota) atramentaria Gyll. la.

A. (Acrotona, "Oxypoda") exigiia

Er. la.
(*) A. {Acrotona) fungi Gr. la, Vb, VII.

A. {s. str.) oblita Er. (det. L. Brun-
din) lb.
(t) A. {Acrotona) orbata Er. (contr. L.

Brundin) Va.

A. {s. str.) triangultim Kr. (contr. L.

Brundin) Va.

A. {Thinobaena) vestita Gr. Ic (c),

VIII (c).

The human transport of animals across the Northern Atlantic

187

Bledius spectabilis Kr. Va (c).

Conosoma lividum Er. lb.

C. testaceum F. (pubescens Gr.) I b, c.

Falagria obscura Gr. Ic, VI b.

Halobrecta flavipes Th. VIII.
(t) Hypocyptus longicornis Payk. Vb.

Lathrobiiim geminum Kr. lb.

Lesteva heeri Fauv. VI b.
(t) Mycetoporus splendidiis Gr. lb.

Myrmecopora iivida Er. Ic (c).
t Oxytelus tetracarinatiis Block Va.

Paederus ftiscipes Curt. Vc (c).

P. litoralis Gr. lb, II, III, Vb, Via,

VII, VIII.

Philonthiis (Gabrius) astutus Er.

(det. V. Hansen) Va.
t Ph. concinnus Gr. VII.
t Ph. cruentatus Gmel. lb.

Ph. fimetarius Gr. Vb, VIII.
t Ph. fuscipennis Mnh. lb, II, III,

IV (c), Va (c), VII.

Ph. laminatus Crtz. Vb.

Ph. marginatus Stroem VIII.
(t) Ph. {Gabrius) nigritulus Gr. Via,

VII.
t Ph. various Payk. (genital slide) II,

VII.
t Ph. varius Gyll. lb, Va, c, VII,

VIII.

Philorinum sordidum Steph. Vb.

Quedius curtipennis Bernh. (fuligi-

nosiis Britten; genital slide) VII.
t Q- fidiginosus Gr. {subfuliginosus

Britten; genital slide) Vb.^

Q. maritimus J. Sahib, (umbrinus

Er., p.p.; genital slide) III.
(tt) Q- picipennis Payk. {molochinus Gr.)

Vb.

Q. picipes Mnh. Vb.

Q. rufipes Gr. la, b, III, IV, Vb.

Q. schatzmayri Grid, (contr. E.

Gridelli) lb, IV, Vb, VII.

Q. tristis Gr. la, b, II, IV, Va, b,

Via, VII.

Sipalia circellaris Gr. lb, Vb.
f Staphylinus ater Gr. Via.
(t) S. caesareus Ced. II.
t S. olens Mull. la. III.

Stenus brunnipes Steph. lb, Vb,

Via.
** S. canaliculatus Gyll. VI b.

S. clavicornis Scop. Vb, VIb, VII,

VIII.

S. flavipes Steph. VIII.

S . formicetorum Mnh. (genital slide)

Vc.

S. fulvicornis Steph. lb, VII.

S. impressus Germ. VII.
** S. juno F. VI b.

S. nanus Steph., coll. Vb (2$).

S. picipes Steph. Vb,

S. similis Hbst. Vb.

S. subaeneus Er. (det. O. Renkonen)

Vb.

Stilicus orbiculatus Payk. VII.
(f) Tachinus rufipes DeG. lb. Via, VII.

T. subterraneus L. VIII.
ft Tachyporus chrysomelinus L. lb (c),

II, III, Vb, Via, VII, VIII.

T. hypnorian F. II, III, IV, Va, b,

Via, VII.
t T. nitidulus F. lb.

T. obtusus L. Vb, Via.

T. pusillus Gr. lb.

T. solutus Er. lb (c), VII.

Xantholinus angiistatus Steph. lb,

II, III, IV, Vb, Via, VII, VIII.

Zyras (Myrmedonia) limbatus Payk.

la.

Histeridae

t Hister purpurascens Hbst. lb, II,
VII.

Lampyridae
Lampyris noctiluca L. VIII (larva).

^ Hatch (1949), who reported Quedius fuliginosus Gr., new to North America, did not
consider curtipennis Bernh. The identification is therefore somewhat uncertain.

CHAPTER II

Cantharidae
ft Cantharis riifa L. Ill, Vb.
C\ rustica Fall. Vb, Via.
Malachim viridis F. Via.
Rhagonycha limbata Th. VII.

Elateridae
ft Agriotes lineatus L. VII.

Byrrhidae
** Cytilus sericeus Forst. lb, Vb.
ft Simplocaria semistriata F. lb, Vb,
Via.

Nitidulidae
(f) Meligethes aeneus F. lb, VII.

M.flavipes Sturm (det. O. Sjoberg)

lb.

M. picipes Sturm (det. O. Sjoberg)

lb.

Cryptophagidae
Atomaria fuscata Schnh. (det. O.
Sjoberg) Va.

Cryptophagus (Micrambe) vini Panz. ,
Bruce Vb (c).

Phalacridae
OUbriis aeneus F. II.
O. Uqiiidus Er. lb.
Stilbiis testaceus Panz. lb.

Lathridiidae
Corticarina {Melanophthalma) fiis-
cula Gyll. Vb.
(t) C. (M.) gibbosa Hbst. Vb (c)
Enicmus transversus Ol. Via.

Coccinellidae
Coccidula riifa Hbst. Ib.Vb.
tt Coccinella (Adalia) bipunctata L. I b.
C. (Thea) 22-piinctata L. lb.
C. (Propylaea) 14-piinctata L. lb.
C. (5. str.) /-punctata L,. Ia,b, Va,b.
Micraspis {Tytthaspis) i6-punctata
L. Vb (c).

Rhizobius litura F. lb (c), Vb, VII.
Subcoccinella 24-punctata L. Via.

Oedemeridae
Oedemera lurida Mrsh. I b, Vb, VI a.
Oe. nobilis Scop. Va, b.

Anthicidae
Anthicus humilis Germ. Ic.

Tenebrionidae
Melanimon tibiale F. Va.
Phylan gibbus F. Va (c).

Scarabaeidae
■f Aegialia arenaria F. Va.
ft Aphodius fimetarius L. Vb.
t A. granarius L. (det. B.-O. Landin)
Vb, VII.
(t) A. rufipes L. Vb.

Cerambycidae
Clytus arietis L. Vb.

Chrysomelidae

Chalcoides fulvicornis F. lb.

Chrysomela banksi F. Vb.
t Phaedon cochleariae F. VIII (c).

Ph. tumidulus Germ. Via.
t Phyllotreta undulata Ktsch. Va.

Curculionidae
Apion aestivum Germ. Via.
A. apricans Hbst. Via
A. assimile Kby. Via.
A.flavipes Payk. lb, Vb.
A. hooker i Kby. Vb.
A. nigritarse Kby. Vb, Via.
A. ononicola Bach Va.
A. ononis Kby. Va (c).
A. sanguineiim DeG. {miniatum
Germ.) lb, Vb, Via.
A. ulicis Forst. Vb (c).
A. violaceum Kby. lb.
A. virens Hbst. Vb.
t Ceuthorrhynchus assimilis Payk. VII.
C. hirtulus Germ. Va.

The human transport of animals across the Northern Atlantic

189

C troglodytes F. la, b.

Cionus alaiida Hbst. Vb.

C. scrophulariae L. Vb (c).

Cneorrhinus {Atactogenus) exarattis

Mrsh. lb.

Liosoma deflexiim Panz. lb.
ft Otiorrhynchiis ligneus Ol. Via.
ft O. ovatus L. Via.

f O. rugosostriatus Gze. lb.
ft O. sidcatus F. lb, II, IV, Vb, VIII.
ft Philopedon plagiatiis Schall. Va (c).

Phytonomus ononidis Chvr. Va.
f Ph. posticus Gyll. {variabilis Hbst.)

Via.

Polydrosus confliiens Steph. Vb.
Sibinia potentillae Germ. lb.
Sitona griseiis F. Va.
ft S. hispidulus F. Vb, Via.

f S. hiimeralis Steph. Via.
ft S. lepidus Gyll. (flavescens Mrsh.)
Vb.
f S. lineatus L. Vb (c), VII.
S. piincticollis Steph. Via.
S. regensteinensis Hbst. Vb.
S. sidcifrons Thunb. lb, Vb, Via.
f Tychius (Miccotrogus) picirostris F.
lb. Via, VII.

Hemiptera Heteroptera

(det. F. Ossiannilsson)

Arenocoris falleni Schill. Va, b.

Beosus tnaritimus Scop. III.

Chartoscirta cocksi Curt. Vb.

Coreus marginatus L. Via.

Dicranomerus agilis Scop. Va.

Dolycoris baccarum L. Vb.

Drymtis sylvaticus F. lb, II, IV, Vb,

Via.

Eiirygaster testudinaria GeofFr. Via.

Harpocera thoracica Fall. III.

Lygus riigidipennis Popp. II.

Megalonotus chiragra F. lb,- IV,

Vb, Via, VII.

M. praetextatus H.-S. Va.

Nabis myrmecoides Costa lb, Vb,

Via.

N. rugosiis L. II, Vb, VIII.
Palojuena prasina L. Vb.
Peritrechiis niibiliis Fall. Ic.
Piesma quadrata Fieb. Ic.
Podops inuncta F. VII.
Salda littoralis L. Vb.
Scolopostethus affinis Schill. lb, IV,
Via, VII.

S. decoratus Hahn Va.
S. puberulus Horv. III.
Stenodema calcaratum Fall. lb.
S. laevigatum L. lb, Vb.
Stygnocorus fuligineus Geoffr. lb,
III, Vb.

Syromastus rhombeus L. /. quadrata
F. Va.

Dermaptera

ft Forficula auricularia L. lb, II, III, IV, Vb, Via, VII, VIII.

Araneae

(det. W. Hackman)

Dysderidae Theridiidae

f Dysdera crocata C. L. Koch IV, Vb. * Ctenium (Robertus) lividiim Bl. lb.

1 90

CHAPTER II

t Theridiuni bimaculatum L. lb, Va.
f Th. ovatiim CI. lb (c).

Linyphiidae
Dicymbium nigrum BI. lb.
Lepthyphantes tenuis Bl. Vb, Via.
Linyphia clathrata Sund. lb, Va,
Via.

L. pusilla Sund. Vb.
Stenomyphantes bucctdentiis CI. {line-
atus L.) la, Va, b, VII.

Erigonidae (Micryphantidae)
Oedothorax fiisciis Bl. Vb.
Oe. retusiis Westr. Vb.
Silometopus interjectus Cambr. lb.
Styloctetor romanus Cambr. Va.

Araneidae
Araneiis redii Scop. Vb.
Meta merianae Scop. Vb.

Tetragnathidae
Pachygnatha degeeri Sund. lb, Vb,
Via, VII.
P. listeri Sund. lb.
** Tetragnatha extensa L. Vb.

Agelenidae
Agelena labyrinthica CI. II, Vb.

Pisauridae
Pisaura mirabilis CI. lb, Vb.

Lycosidae
Alopecosa pulverulenta CI. lb, Vb,
Via, VII.
Arctosa leopardus Sund. Vb (c).

A. perita Latr. Va.
Pardosa nigriceps Thor. Va.
* P. palustris L. la.
P.proxima C. L. Koch IV, Vb.
P.puUata CI. la, III, Va.
Trochosa riiricola DeG. II.
T. terricola Thor. Via.
Xerolycosa miniata C. L. Koch Va.

Gnaphosidae

Drassodes lapidosus macer Thor. II,

IV, Vb.

Zelotes pedestris C. L. Koch Vb.

Clubionidae
Cheiracanthium erraticum Walck.
Va.

Clubiona compta C. L. Koch Vb.
C. neglecta Cambr. lb, IV, Va, Via.
C. recliisa Cambr. lb, II, Va, VII.
C terrestris Westr. III.
ft Micaria puUcaria Sund. lb, IV, Vb,
Via.
Phriirolithus festivus C. L. Koch Vb.

Xysticidae
Xysticus cristatus CI. lb, II, IV,
Vb, Via, VII.
X. kochi Thor. Vb.

Philodromidae
** Tibellus maritimiis Menge Va.

Salticidae
Euophrys frontalis Walck. Vb.
Eu. molesta Cambr. Vb.
Heliophanes flavipes C. L. Koch Vb.

Opilionida

(det. W. Hackman)

Lacinius ephippiatus C. L. Koch lb.

ft Phalangiiim opilio L. lb, IV, Va, b,
VI, VII.

The human transport of animals across the Northern Atlantic

191

Chilopoda

(det. E. Palmen)

ft Cryptops hortensis Leach Vb, Via,

VII.
tt C. parisi Brol. Ill, IV.

ft Haplophilus subterraneus Leach la,

IV, Via, VII, VIII.
tt Lithobius forficatus L. la, b, II, III,

Vb, Via, VII.

Diplopoda

(det. E. Palmen)

tt Brachyiuliis littoralis Verh. Vb.

tt Cylindroiulus britannicus Verh. Vb.

tt C. frisius Verh. Vb.

tt C. silvarum Mein. Vb.

tt C. teutonicits Pocock lb.

Glomeris marginata Vill. VIII.
tt Ophyiulus fallax Mein. Vb, VIII.

t Polydesmus angustus Latz. lb.
tt P- inconstans Latz. {coriaceus Por.)
Vb, Via.

Schizophyllimi sabulosum L. Vb,
VII.

Tachypodoiulus albipes C. L. Koch
Vb, VII, VIII.

Oniscoidea (Isopoda Terrestria)

(det. E. Palmen)

tt Androniscus dentiger Verh. Vb.

Armadillidiiim depressum Brdt. Vb.
t A. vulgare Latr. la, b, IV, Vb, Via,

VII.

Ligia oceanica L. Ic.
tt Oniscus asellus L. lb, II, III, IV,

Vb, Via, VII, VIII.

t Philoscia muscorum Scop. lb, II, IV,

Vb, VIII.
tt Porcellio scaber Latr. la, b, II, III,

IV, Vb, Via, VII, VIII.
tt Trichoniscus provisorius Racov. II.

Mollusca (shell-bearing)

(det. H. W. Walden)

Ashfordia granulata Alder IV, Vb.
** Ciotiella (Cochlicopa) lubrica Miill.

lb, Vb, Via, VII, VIII.

C (C). minima Siem. VII.

Clausilia bidentata Stroem II, Vb,

Via, VIII.
tt Discus rotundatiis Miill. III.

Helicella (Candidula) caperata Mont.

la, b, Va, b. Via, VII.

H. (Cernuella) virgata Da Costa

II, IV, Vb, Via, VII.
■f Helix aspersa Mull. lb, IV,

VIII.

t Hygromia (Trichia) hispida L. lb,

III, Via, VII, VIII.
tti^- {T.) striolata Pfeiff. Via, VII.

Lauria cylindracea Da Costa IV, Vb.
t Monacha {Theba) cantiana Mont.

VIII.
t Oxy chillis alliarius Miill. Vb.
t O. cellarius Miill. la.
tt O. draparnaldi Beck II, IV, Vb,

Via, VII.
t O. helveticus Blum VIII.

Retinella nitidula Drap. Vb.

Vitrina pellucida Mull. Vb, Via.

192 CHAPTER II

TABLE 4. Animals found on ballast-places in S. W. England.

Coleoptera

Number of species

Tot

al

Eur.-.'\merican

Introduced into N. Amer.

242

72

68= 28%

Carabidac

83

22

21 = 25 %

Curculionidae

37

12

12= 32%

Others

122

38

35 = 29 %

Hem. Heteroptera

26

I

Dermaptera

I

I

I = 100 %

Araneae

45

8

4 = 9 %

Opilionida

2

I

I = 50 %

Chilopoda

4

4

4 = 100 %

Diplopoda

1 1

8

8= 73%

Isopoda Terr.

8

6

6= 75%

Mollusca

18

10

9= 50%

All animals

357

1 1 1

lOI = 28 %

The Carabid beetles

Special attention was paid to Carabid beetles, every specimen of this group being
collected in order to get an idea of the relative abundance of the different species.
The results are given in table 5.

From the survey given there, it seems appropriate to draw attention to the
following points:

1. Of 83 Carabid species collected on the ballast places, 21 are here regarded
as introduced into North America, that is 25 per cent.

2. Of 40 Carabid species regarded as introduced into North America from
Europe (table 2, p. 144), these 21 species make no less than 53 per cent. If 3 species
are removed from the list, because they are not permanent inhabitants of the
British Isles (Amara anthohia, Carabus auratus and cancellatus) and 2 species
{Clivina collaris and /owor), found by other collectors on the Avon banks at Bristol
(p. 186), are added to the "ballast species", it appears that out of 37 North Ameri-
can species of Carabidae which may have been introduced from the British Isles,
no less than 23, that is 62 per cent, have been observed on the stated or supposed
ballast-places of southwestern England.

3. Another point of importance is the frequency and abundance of the "emigrants"
among Carabids collected on the ballast-places (Diagr. 3 and 4); only the occur-
rence on "waste places" is here considerd. If frequency is determined by the num-

The human transport of animals across the Northern Atlantic

193

TABLE 5. Carabid beetles collected on the ballast-places. Symbols as in the list of

species, p. 186 a.f.

Abax ater Vill

Acupalpus dubius Schky.
t A. meridianus L. ...

Agomim atratum Dft.

A. dor sale Pont

A. micans Nic

ft A. miilleri Hbst

A. obscuriim Hbst. . . .
tf A. ruficorne Gze. . . .
ft Amara aenea DeG. . .
ft A. aulica Panz

A. communis Panz. . .

A. convexior Staph. . .

A. convexiuscula Mrsh. .

A. equestris Dft

W A. familiar is Y)h. . . .
ft A. lunicollis Schio. . .

A. montivaga Sturm . .

A. ovata F

A. plebeja Gyll

A. similata Gyll. . . .

A. tibialis Payk

t Anisodactylus binotatus F.
t Asaphidion flavipes L.

Badister bipiisttdatiis F. .
Bembidion assimile Gyll.

B. concinnum Steph. . .

B. guttula F

B. tricolor Bed

ft B. lampros Hbst. . . .

B. laterale Sam

B. hinulatum G.-F. . .

B. minimum F

B. normannum Dej. . .
t B. properans Steph. . .

13 — 565597 Lindroth

A. Waste places

la, b II III IV Vb

0+7
0+ I

0+ 7

+ 14

0+ I

0+2
0+ I

0+2
o + 10
0+2
+ 13
0+ 6

+ 12

17

Via

VII

42

32

Total:
Number

of
lo-

cal-
ities

spe-
ci-
mens

B. Saline places C. Sand

Ic

6 —

18 —

45 Ii-

5
3

I
76

I
2
2

4

I

19

3

I

16

40

12

3

15

7

3

38

13

Vc

15

VI b

24
3

VIII Va

17

I

26

6 —

194

CHAPTER II

la, b

III

IV

Vb

B. quadrimaculatntn L. . .
ft B. tetracolum Say (usttila-

titm auctt.)

B. variiim OI

t Bradycelliis harpalinus Serv.

B. verbasci Dft

Broscus cephalotes L. . . .
CaJathus erratus Sahib. . .

f C. fuscipes Gze

C. melanocephaliis L. . . .

C. mollis Mrsh

Carabiis violaceus L. . . .
Demetrias atricapilhis L.
Dichirotrichus pubescens

Payk

Dromius linearis Ol. . . .

D. melanocephalus Dej. . .
Dyschiriiis globosiis Hbst. .
Eiirynebria complanata L. .

ft Harpalus affinis Schrk.
(aeneus F.)

H. anxius Dft

H. attenuatus Steph. . . .

H. latus L

H. puncticeps Steph. {an-
gusticollis J. Mull.) . . .

H. rubripes Dft

\\ H. rufipes T>eG. (pubescens
Mull.)

//. seladon Schaub

H. tardus Panz

H. tenebrosus Dej

** Loricera pilicornis F. . . .

Metabletus foveatus G.-F. .

Microlestes maurus Sturm .
\\ Nebria brevicollis F. . . .

N. salina Frm. {degenerata

Schauf.)

ff Notiophilus biguttatus F.

A'', palustris Dft

A'^. substriatus Steph. . . .

Olisthopiis rotundatus Payk.

Pogonus chalceus Mrsh. . .

o + 2

o +
o +

5 +

I +

o+ 7

o + 4

13 + 17

I + o

+

8+ o

o + n

0+ I

1+ 3

26+3

9+0

28

13

19

Via

VII

Total:

loc. spec.

13

Ic

4

28

I
20

II

I

97

3

6

4

13

4

46

2

14

I

I

I

9

I

I

2

5

I

I

6

54

I

9

4

12

I

2

2

3

I

19

Vc

17

15

VIb

VIII

The human transport of animals across the Northern Atlantic

195

Pterostichns ciipreus L. . .

P. madidus F

P. niger Schall

P. jiigita F

ft P. stremiiis Panz

P. verualis Panz

P. versicolor Sturm {coe-
rulescens auctt.)

Stenolophus teutonus Schrk.
t Stomis pumicatus Panz. . .
f Trechus obtusus Er. . . .

T. quadristriatus Schrk.

la, b

II

III

IV

Vb

Via

VII

Total

la

Vc

VI b

loc.

spec.

Vll 1 V a

0+ 4
0+ I

0+ 2
0+ 2

I
I

3

—

I

4
I

I

7

I

2
7

I

I

I

3

I
I
2
2

I

I
I
2
2

4
6
I
I

2
8

I

2
I

4
10

—

2

3

I

1

i ~

Number of species
Number of specimens

34
207

22
107

3

3

10
63

32
163

15
43

25
170

64 5

756 I II

7
59

II

53

12 II
68 96

ber of investigated localities (out of 7) inhabited by a species (Diagr. 3), it appears
that the "emigrants" show a higher frequency (average: about 3 localities per
species) than the rest (average: about 2 localities per species). — Still more striking
is their higher abundance (Diagr. 4).

The 7 most abundant species in order of precedence, the seven "waste ground"
localities (table 5) taken together, are:

Harpalus affinis
Amara aenea
Nebria brevicollis
Harpalus riifipes

Acupalpus meridianus
Amara plebeja
Bembidion lampros

Number of localities
3 U

DIAGR. 3. Frequency groups of the 64 species of Carabid beetles found on waste
ground (A, table 5) on 7 ballast places in SW England.
Black = "emigrants". White = non-emigrants.

196

CHAPTER II

per cent
100t

Number of specimens
11—20 21—40 /11— 60 61—80

DIAGR. 4. Abundance groups of the 64 species of Carabid beetles found on waste
ground (A, table 5) on 7 ballast places in SW England.
Black = "emigrants". White = non-emigrants.

Of these all but one (Amara plebeja) occur in North America as "emigrants".

If both frequency and abundance are considered, both on a 6 degree scale (as
in diagr. 3 and 4), it appears that of 16 species topping the list 11 are "emigrants".

It certainly cannot be regarded as presumptuous to declare that the composition
of the Carabid fauna, especially of its "emigrant" element, on the ballast-places
investigated strongly supports the idea of a ship transport of the said element
across the Atlantic.

Still more pronounced is the geographical composition of some other of the col-
lected groups of animals:

Of 37 species of Curcidionids 12 are "emigrants" = 32 per cent

,, 18 ,, ,, Molluscs 9 ,, ,, — 50 ,, ,,

,, 23 ,, ,, "Iso-Myriapods" 19 ,, ,, = 83 ,, ,,

On the other hand, the ballast material contains only few species of Hemiptera
Heteroptera and Araneae in common with North America (for explanation, vide
below, p. 199).

The plants

Without any pretensions to completeness, the more abundant or otherwise
conspicuous vascular plants of the ballast-places were noted. The identification of
some critical species was made or controlled by Mr. K. A. Mattisson, Botanical
Institute, University of Lund. The complete list of observed species, divided into
the three types of investigated localities, follows here:

The human transport of animals across the Northern Atlantic

List of vascular plants observed on the ballast-places^

t = occurring in North America, regarded as introduced. ^
ff = ,, ,, Newfoundland, ,, ,, ,,

* = ,, ,, North America, ,, ,, indigenous.

197

A. On waste ground (loc. I a, b, II, III, IV, V b, VI a, VII).

ft Achillea millefolium L.

f Alliaria petiolata Bieb.
{officinalis Andrz.)

f Anisantha (Bromiis) sterilis L.
ft Artemisia vulgaris L.

t Ballota nigra L.
ft Bel I is perennis L.
ft Brassica tiigra L.
ft Calystegia {Convolvulus) sepium L.
ft CapseUa bursa-pastoris L. (coll.)

f Carduus crispus L.
ft Cerastium vulgatum L.

f Chaerophyllum temulum L.
ft Chrysanthemum leucanthemum L.
ft Cirsium arvense L.
ft C. vulgare Savi {lanceolatum L.)
ft Coronopus didymus L.
ft Dactylis glomerata L.

f Daucus carota carota L.

f Epilobium hirsutum L.
ft Equisetum arvense L.
ft Festuca ovina L.
tt Galium aparine L.

t Geranium dissectum L.

* G. robertianum L.
tt Holcus lanatus L.

t Hordeum murinum L.
tt Hypochaeris radicata L.

Kentranthus ruber L.
tt Lolium perenne L.
tt Lotus corniculatus L.
tt Matricaria maritima inodora L.

* M. matricarioides Less, {discoidea D.
C.)

t Melilotus altissima Thuill.
tt Myosotis arvensis L.

t Plantago coronopus L.
tt P- lanceolata L.
tt P- major L.

t -P. media L.
tt -Poo annua L.
tt -P- pratensis L.
tt -P- trivialis L.

t Potentilla reptans L.
tt Ranunculus repens L.

t Reseda luteola L.
tt Rumex crispus L.

t Sarothamnus {Cytisus) scoparius L.
tt Senecio jacobaea L.
tt »S'. vulgaris L.
tt Silene cucubalus Wibel {vulgaris

Moench)
tt Sisymbrium officinale L.
tt Solanum dulcamara L.
tt Sonchus oleraceus L.
tt Stellaria media L.

t Trifolium campestre Schreb. {procum-
bens auctt.)
tt ^- hybridum L.
tt ^- pratense L.
tt 7"- repens L.
tt Tussilago farfara L.

t tZ/ex europaeus L.
tt Urtica dioica L.
tt r^i^^« angustifolia L.
tt ^- cracca L.

t Vulpia bromoides L.

1 The nomenclature of species follows Clapham, Tutin, & Warburg, "Flora of the British
Isles" (1952). If different from Fernald, in "Gray's Manual" (1950), or other current
literature, the latter name is noted as a synonym in brackets.

2 According to Fernald (1950); Chaerophyllum, Hordemn, and Vulpia according to Clap-
ham, Tutin, & Warburg (1952).

jg8 CHAPTER II

B. On saline places (loc. I c, V c, VI b).

tt Artemisia vulgaris L. Phragmites communis Trin.

Beta vulgaris maritima L. * Piiccinellia maritima Huds.

* Glaux maritima L. * Spergularia (Spergula) salina Presl.

Halimione (Obione) pedunadata L. {marina auctt., p. p.)

t Hordeiim murimim L. tt Triglochin maritimum L.
tt Matricaria maritima maritima L.

C. On sand (loc. V a).

t Ammophila (Psamma) arenaria L. Euphorbia paralias L.

tt Anagallis arvensis L. E. portlandica L.

tt Cirsium arvense L. Ononis repens repens L.

t Cynoglossum officinale L. tt Senecio jacobaea L.

t Erodium ciciitarium L.

The plant list gives very clear evidence. Of the total number of 77 species,
72 (94 per cent) are known to occur in North America and no less than 65 (84 per
cent) are there regarded as European introductions, 44 species (57 per cent) also
in Newfoundland.

If only the flora of waste ground (63 species) is considered, all plants except
Kentranthus ruber (wich is originally introduced in the British Isles) are recorded
from North America and all the rest, except Geranium robertianum and Matricaria
matricarioides, are regarded as adventives from Europe, that is 95 per cent!
42 of these (67 per cent) occur in Newfoundland.

It may be concluded that ballast transport has been still more efficient in carrying
plants across the Atlantic than in carrying the lower terrestrial fauna. This is not
surprising, considering the generally higher resistance and longer diapause (period
of rest) in plant diaspores by which they no doubt as a rule are more suited to
endure the voyage.

The reduced number of species common to Europe and North America among
the plants of saline localities and of pure sand is in conformity with the results
obtained from the fauna (below, p. 199).

The selection of animal species which managed to cross the Atlantic

The successful transport of an animal species with ballast from one continent
to another involves a long series of lucky chances. One would be inclined, perhaps,
to tegard the selection of species which survived the adventure as a matter of

The human transport of animals across the Northern Atlantic 199

pure chance too, provided that the species in question regularly occurs on places
where ships took ballast. This is not so. From the account above it is evident that
certain groups of animals, for instance Carabid beetles and "Iso-Myriapods",
have crossed the Atlantic, or at least made successful settlements on the other
side, more often than plant bugs {Hemiptera Heteroptera), Spiders [Araneae) and
others. This, of course, is not due to their taxonomic position but to one or more
characteristic features of their natural history.

By grouping the animals found on the English ballast-places according to such
principles, instead of taxonomically, it would be possible to fix the foremost
qualifications favouring transport with ballast. The necessary analysis of the ecolog-
ical requirements and of certain other biological properties of the species was
carried out for all Coleopterous insects observed on the ballast-places (table 6).

The biological and ecological properties recorded in the 7 columns of table 6
are valid for the fullgrown insect (imago), the larva in several species having a
diflFerent biology. The method used is defensible from the point of view that the
imago (especially a fertilized female) is far more important for a successful coloni-
zation of an oversea area than is the larva.

Let us first consider the columns of table 6 separatedly.

1st column. The planticolous species (p) constitute 28 per cent (48 species) of
the 174 Coleoptera from the ballast-places not introduced into North America,
but only 19 per cent (13 of 68 species) of the "emigrants". Terricolous species are
favoured, as has already been pointed out by Brown (1940, 1950). Similarly,
Molluscs and Iso-Myriapods are strongly represented among the introduced
forms, whereas very few of the many planticolous Hemiptera and Araneae (true
spiders) were carried over. Planticolous forms were less likely to be taken on board
with the material generally used- as ballast (above, p. 164) and, if this happened,
they ran the risk of being blown overboard due to their habit of climbing.

2nd column. The hygrophilous species (h) constitute 25 per cent (44 species)
of the "non-emigrants", but only 6 per cent (4 species) of the "emigrants".
Xero- and mesophilous species are favoured, partly because dry ballast was more usual,
partly because there was considerable risk of desiccation during the long voyage
or, if the ballast became wet, the cause was usually salt water, which is injurious
to normal hygrophilous insects.

3rd column. The "emigrants" and the remaining species were distributed on
the three investigated kinds of soil as follows:

waste ground pure sand saline localities

"Emigrants" 61 species = 90 % 6 = 9% 1=1%

Others 1 30 ,, - 75 % 14 = 8 % 30 = 17 %

200 CHAPTER II

The essential difference depends on the fact that almost no species from saline
localities (in the collected material only the Chrysomelid beetle Phaedon cochleariae
F.), artd no halobiontic (salt-demanding) species, were introduced into North
America. This may sound absurd but must be explained by the fact that the ballast,
at least as a rule, was not taken from the tidal zone (cf. above, p. 165). The
soil there was wet, heavy to handle, and eventually lost part of its valuable
weight; it was often muddy and difficult to clean out of the ship afterwards.

4th column. There is no difference between phytophagous beetles (p) and species
requiring other kinds of food, the former group constituting 25 per cent
(17 species) of the "emigrants" and likewise 25 per cent (44 species) of the
remainder.

5th column. A slight difference is found between polyphagous and oligo- or
monophagous species, that is those specializing on restricted kinds of food. The
former group makes 85 per cent (58 species) of the "emigrants", 79 per cent (137
species) of the remainder. One would have expected a clearer demonstration of the
fact that a more or less pronounced monophagy disfavours introduction over long
distances. Apparently the figures are not quite representative, as is clear from a
somewhat different calculation on Curculionids (below, p. 210).

6th column. The constantly short-winged (brachypterous) species and those
occurring in both macro- and brachypterous individuals (dimorphic species)
(b and d of column 6) are better represented among the "emigrants", 32 per cent
(22 species), than in the rest, 25 per cent (44 species). Flightless insects no doubt
had more chance to stay onboard during the voyage, escaping the risk of being
blown into the sea. The Hemiptera, as a rule capable of flight in the adult stage,
and the true spiders (Araneae), most of which are able to ascend by "ballooning",
at least as young, are also very poorly represented among the introduced species.
It is characteristic that the only true spider both occurring in the English ballast-
material and regarded as a European introduction in the fauna of Newfoundland
(Hackman, 1954, p. 40), is Micaria pulicaria Sund., a pronounced ground-species
without the habit of ballooning, found on four of the eight investigated ballast-
places.

yth column. Five of the species observed on the ballast-places are constantly
parthenogenetic {Amischa analis, and four species of Otiorrhynchus), only females
being known, and all of them have been carried over to North America. This is
another proof of the extraordinary advantage of this form of reproduction for the
dispersal of a species: for a single specimen, in any stage of development, to be
deposited in a suitable spot, is sufficient to colonize a new area! The parthenogenesis
in Coleoptera, with very few exceptions (Suomalainen, 1953), is combined with
polyploidy (increase of chromosome number) and possibly this in itself effects a

The human transport of animals across the Northern Atlantic 201

higher viability in comparison with related bisexual forms and species (Lindroth,
1954 e). The aspect of parthenogenesis in the first place applies to Curculionids
(Weevils; below, p. 210). Concerning the Staphylinid Amischa analis Gr., vide
Strand (1951). In some Iso-Myriapods, too, a more or less constantly partheno-
genetic reproduction has largely contributed towards the wide passive dispersal
(mainly with man) of these species. In the Newfoundland fauna this applies to
the Chilopod Lamyctes fulvicornis Mein. (Palmen, 1954, p. 144), the Diplopod
Proteroiulus fuscus a. St. (Palmen, 1952, p. 17), and the Oniscoid Trichoniscus
pusillus pusillus Brandt (Palmen, 195 1, p. 2).

Another animal, the extremely rapid dispersal of which may be attributed at
least in part to parthenogenesis, is the euryhaline Mollusc Hydrobia (Potamopyrgus)
jenkinsi Smith {vide Bondesen & Kaiser, 1950). Its original home is unknown but
it may have been carried to NW. Europe by human transport from America. The
Lumbricid worms Allolobophora caliginosaSav., Bimastus tenuis Eis., Dendrobaena
subrubicunda Eis., Eisenia rosea Sav.,and Eisetiiella tetraedra Sav., almost cosmopoli-
tan in distribution and evidently introduced into North America from Europe, are
likewise parthenogenetic, at least in part (Gavrilov, 1935, 1940; Muldal, 1952;
Omodeo, 1952, p. 246), and this may also apply to the rest of peregrine
Oligochaeta.

Functionally similar are hermaphroditic species capable of self-fertilization.
Possible examples among terrestrial (and, in part, limnic) animals are to be found
among Mollusca and Oligochaeta. Of the species dealt with in this book, Arion
ater L. (Kiinkel, 1916, p. 409 a.f.; Perrot, 1939) and several species of Lymnaea
(Hubendick, 195 1, p. 32) are considered capable of self-fertilization^ and the same
has been supposed with respect to the form of Deroceras laeve Miill. with reduced
penis (Boettger, 1932, p. 402; Meeuse & Hubert, 1949, p. 25), as well as for
members of the following families (in part even the same species), which have
crossed the North Atlantic by human agencies: Arionidae, Helicidae, Limacidae,
Physidae, Planorbidae, and Zonitidae {vide bibliography in Larambergue, 1939,
p. 41-43). The occurrence of self-fertilization in Oligochaeta has been suggested in
those cases, in which reproduction was observed in isolated individuals, as in
Dendrobaena octaedra Sav. and Eisenia foetida Sav., both of world-wide distribu-
tion, but parthenogenesis, as in the species mentioned above, seems equally
probable.

^ It must be regarded as very difficult to separate, conclusively, selflFertilization from
parthenogenesis in hermaphroditic animals without cytological investigation.

202 CHAPTER II

TABLE 6. Coleoptera from the ballast places of southwestern England divided into
certain biological and ecological groups.

1st column. Selection of stratum.^

t = terricolous (ground-living); (t) = regularly climbing the vegetation
p = planticolous (plant-living).

True subterraneous species were not considered.

2nd column. Selection of moisture.^

X = xerophilous (preferring dry places); (x) = less pronounced

m = mesophilous (intermediate)

h = hygrophilous (preferring wet places); (h) = less pronounced.

3rd column. Selection of soil. 2

c = on waste ground; (c) = also on sand or (and) saline soil

s = on pure sand (loc. V a)

t = on saline soil (tidal zone, ± clayish).

4th column. Selection of food.i

p = phytophagous (feeding on living plants, incl. mosses and fungi)

z = zoophagous (feeding on other animals)

s = schizophagous (feeding on decomposing organic matters)

o = zoo- or schizophagous; m = mixed diet.

5th column. Specialisation on food.^

m = monophagous (restricted to plants or animals belonging to one single genus)
o = oligophagous (restricted to plants or animals belonging to one single family

or corresponding group)
p =polyphagous (taking more diversified food).

6th column. Means of dispersal. ^ ^

m = macropterous (stated or supposed to fly); (m) = brachypterous in other areas
b = brachypterous (fligthless)^; (b) = macropterous in other areas
d = dimorphic (occurring on the ballast places in both forms).

7th column. Means of reproduction. 1
s — bisexual; p = parthenogenetic.

^ According to general knowledge of the species.
* According to observations on the ballast places.

' The following species seem to be flightless in spite of comparatively well developed
hind-wings: the species of Quedius and Staphylimis, Chrysomela banksi.

The human transport of animals across the Northern Atlantic

203

Species regarded as European introductions into
North America in bold face types

-a •«
o u

o -^

Fam. Carabidae

Abax ater

Acupalpus dubius ....

A. meridianus

Agonum atratum ....

A. dorsale

A. micans

A. MuUeri

A. obscurum

A. ruf icorne

Amara aenea

A. aulica

A. communis

A. convexior

A. convexiuscula

A. equestris

A. familiaris

A. lunicoUis

A. montivaga

A. ovata

A. plebeja

A. similata

A. tibialis

Anisodactylus binotatus .
Asaphidion f lavipes . .
Badister bipustulatus , .
Bembidion assimile . . .

B. concinnum

B. guttula

B. iricolor

B. lampros

B. laterale

B. lunulatum

B. minimum

B. normannum

B. properans

B. quadrimaculatum . . .
B. tetracolum {ustulatum)
B. varium

h
m

h

h
m
(h)
h

m
m

(x)
(x)

m

(h)
m
h
h
h
h
m
h
h
h
h
m
m
m
h

c

t

c

t
(c)

c

(c)
(c)

c

(c)
c
c
c

c
c

c
t
t
t
t
c
t
t
t
t

(c)
c
c
t

m
m
m
m

m

m
m

m
z

P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P

m
m
(b)
b?
m
m
m

m
tn

m
m

m

d

m
(m)

m
(b)
(b)

(b)
m

204

Bradycellus harpalinus .

B. verbasci

Broscus cephalotes ....
Calathus erratus

C. fuscipes

C. melanocephalus . . . .

C. mollis

Carabus violaceus . . . .
Demetrias atricapillus . . .
Dichirotrichus pubescens .
Dromius linearis

D. melanocephalus . . . .
Dyschirius globosus . . .
Eurynebria complanata . .
Harpalus affinis (aeneus) .

H. anxius

H. attenuatus

H. latus

H. puncticeps (angusticollis)

H. rubripes

H. ruf ipes (pubescens) . . .

H. seladon

H. tardus

H. tenebrosus

Loricera pilicornis . . . .
Metabletus foveatus . . .
Microlestes maurus . . . .
Nebria brevicollis . . . .
N. salina {degenerata) . . .
Notiophilus biguttatus

N. palustris

N. substriatus

Olisthopus rotundatus . .

Pogonus chalceus

Pterostichus cupreus . . .

P. madidus

P. niger

P. nigrita

P. strenuus

P. vernalis

P. versicolor (coertilescens)
Stenolophus teutonus . .
Stomis pumicatus . . . .
Trechus obtusus . . . ,

CHAPTER II

I

2

3

4

5

6

7

(t)

m

c

m

P

(m)

s

(t)

m

c

m

P

m

s

X

s

z

P

m

s

X

s

m

P

(b)

s

m

c

m

P

b

s

m

c

m

P

(b)

s

X

s

m

P

(m)

s

m

c

z

P

b

s

X

(c)

m

P

m

s

h

t

m

P

m

s

X

(c)

m

P

(b)

s

X

(c)

m

P

m

s

m

t

z

P

b

s

(h)

s

z

P

m

s

X

(c)

m

P

m

s

X

s

m

P

m

s

X

c

m

P

m

s

m

c

m

P

m

s

(x)

c

m

P

m

s

X

c

m

P

m

s

m

c

m

P

m

s

m

c

m

P

m

s

X

c

m

P

m

s

X

c

m

P

m

s

h

(c)

z

P

m

s

X

c

z

P

b

s

X

c

z

P

(b)

s

m

(c)

z

P

m

s

m

c

z

P

m

s

m

c

z

P

d

s

m

c

z

P

(b)

s

m

c

z

P

m

s

X

c

m

P

d

s

h

t

z

P

(b)

s

ni

c

z

P

m

s

ni

(c)

z

P

b

s

m

c

z

P

m?

s

h

c

z

P

m

s

(h)

(c)

z

P

(b)

s

h

c

z

P

d

s

m

c

z

P

m

s

(h)

c

m

P

m

s

m

c

m

P

b

s

m

c

m

P

(b)

s

1

The human transport of animals across the Northern Atlantic

205

T. quadristriatus

Fam. Hydrophilidae
Helophorus brevipalpis . . . .
Megasternutn boletophagum
Ochthebius impressicollis . . .

O. marinus

Sphaeridium scarabaeoides .

Fam. Silphidae

Ablattaria laevigata

Silpha tristis

Fam. Staphylinidae
Aleochara bipustulata . . . ,

Amischa analis

A. decipiens

Astenus longelytratus

Astilbus canaliculatus . . . ,

Atheta angustula

A. atramentaria

A. exigua ...

A. fungi

A. oblita

A. orbata

A. triangulum

A. vestita

Bledius spectabilis

Conosoma lividum

C. testaceum (pubescens) . . .

Falagria obscura

Halobrecta flavipes

Hypocyptus longicornis . . .

Lathrobium geminum

Lesteva heeri

Mycetoporus splendidus . .

Myrmecopora uvida

Oxytelus tetracarinatus . . .

Paederus fuscipes

P. litoralis

Philonthus astutus

Ph. concinnus

Ph. cruentatus

Ph. fimetarius

Ph. fuscipennis

Ph. laminatus

Ph. marginatus

2

3

4

5

6

7

X

c

m

p

m

s

h

t

z

p

m

s

m

c

s

p

m

s

h

t

z

p

m

s

h

t

z

p

m

s

m

s

s

p

m

s

m

c

m

p

b

s

m

(c)

m

p

b

s

m

c

p

m

s

m

c

p

m

p

m

c

p

m

s

m

c

z

p

m

s

m

c

p

b

s

(h)

c

p

m

s

m

c

s

p

m

s

(h)

c

p

m

s

m

c

p

m

s

m

c

p

m

s

(X)

s

p

m

s

ni

c

p

m

s

h

t

p

m

s

h

t

p

m

s

m

c

p

b

s

m

c

p

m

s

m

c

p

m

s

h

t

p

m

s

m

c

s

p

m

s

(h)

c

z

p

(b)

s

h

t

p

b

s

m

c

z

p

tn

s

h

t

p

m

s

m

s

s

p

m

s

h

t

z

p

m

s

m

c

z

p

b

s

m

s

p

m

s

m

c

s

p

m

s

m

c

s

p

nti

s

m

c

s

p

m

s

m

(c)

s

p

m

s

m

c

p

m

s

m

t

s

p

m

s

206

CHAPTER II

Ph. nigritulus

Ph. varians

Ph. varius

Philorinum sordidum ....
Quedius curtipennis ....
Q. fuliginosus {subfuliginosus)

Q. maritimus

Q. picipennis (molochimis)

Q. picipes

Q. rufipes

Q. schatzmayri

Q. tristis

Sipalia circellaris

Staphylinus ater

S. caesareus

S. olens

Stenus brunnipes

S. canaliculatus

S. clavicornis

S. flavipes

S. formicetorum

S. fulvicornis

S. impressus

S. juno

S. "nanus"

S. picipes

S. similis

S. subaeneus

Stilicus orbiculatus

Tachinus rufipes

T. subterraneus

Tachyporus chrysomelinus

T. hypnorum

T. nitidulus

T. obtusus

T. pusillus

T. solutus

Xantholinus angustatus . . .

Zyras limbatus

Fam. Histeridae
Hister purpurascens . . .

I

2

3

4

5

6

7

m

c

o

p

m

s

m

c

s

p

m

s

m

(c)

s

p

m

s

p

m

c

p

p

ni

s

m

c

z

p

bi

s

m

c

z

p

fai

s

m

c

z

p

b

s

m

c

z

p

bi

s

m

c

z

p

bi

s

(x)

c

z

p

bi

s

(x)

c

z

p

bi

s

m

c

z

p

bi

s

m

c

o

p

b

s

(X)

c

z

p

bi

s

m

c

z

p

bi

s

(x)

c

z

p

bi

s

(h)

c

z

p

b

s

h

t

z

p

m

s

m

(c)

z

p

m

s

(h)

t

z

p

b

s

h

t

z

p

m

s

h

c

z

p

m

s

m

c

z

p

m

s

h

t

z

p

m

s

(h)

c

z

p

m

s

(h)

c

z

p

m

s

h

c

z

p

m

s

(h)

c

z

p

m

s

m

c

o

p

ni

s

m

c

s

p

m

s

m

t

s

p

m

s

m

c

o

p

m

s

m

(c)

o

p

ni

s

m

c

o

p

m

s

m

c

o

p

m

s

m

c

o

p

m

s

m

c

o

p

ni

s

m

c

z

p

m

s

m

c

o

o

m

s

t

m

c

o

p

m

s

^ Vide footnote 3, p. 202.

The human transport of animals across the Northern Atlantic

207

Fam. Lampyridae
Lampyris noctiluca . .

Fam. Cantharidae
Cantharis rufa ....

C. rustica

Malachius viridis ....
Rhagonycha limbata . .

Fam. Elateridae
Agriotes lineatus . . .

Fam. Byrrhidae
Cytilus sericeus ....
Simplocaria semistriata

Fam. Nitidulidae
Mellgethes aeneus . .

M. flavipes

M. picipes

Fam. Cryptophagidae
Atomaria fuscata ....
Cryptophagus vini . . .

Fam. Phalacridae
Olibrus aeneus ....

O. liquidus

Stilbus testaceus ....

Fam. Lathridiidae
Corticarina fuscula . . .

C. gibbosa

Enicmus transversus . .

Fam. Coccinellidae
Coccidula rufa ....
Coccinella bipunctata .
C. 22-punctata ....

C. 14-punctata

C. 7-punctata

Micraspis i6-punctata
Rhizobius litura ....
Subcoccinella 24-punctata

Fam. Oedemeridae
Oedemera lurida ....
Oe. nobilis

Fam. Anthicidae
Anthicus humilis . . .

Fam. Tenebrionidae
Melanimon tibiale . . .
Phylan gibbus

I

2

3

4

5

6

7

t

m

c

z

b

s

p

m

c

m

P

m

s

p

m

c

m

P

m

s

p

m

c

m

P

m

s

p

m

c

m

P

m

s

t

m

c

m

P

m

s

t

m

c

P

m

s

t

m

c

P

m

s

p

m

c

P

m

s

p

m

c

P

P

m

s

p

m

c

P

P

m

s

t

m

s

P

P

m

s

p

m

c

P

m

s

p

m

c

P

m

s

p

m

c

P

m

s

p

m

c

P

P

m

s

t

m

c

P

P

m

s

p

m

c

P

P

m

s

t

m

c

P

P

m

s

p

(h)

c

z

P

m

s

p

m

c

z

P

m

s

p

m

c

z

P

m

s

p

m

c

z

P

m

s

p

m

c

z

P

m

s

p

m

c

z

P

m

s

t

X

c

z

P

b

s

p

m

c

p

P

b

s

p

m

c

p

P

m

s

p

m

(c)

p

P

m

s

t

m

t

P

m

s

t

X

s

s

P

m

s

t

X

s

s

P

b

s

2o8

CHAPTER II

F"am. Scarabaeidae

Aegialia arenaria

Aphodius fimetarius . . .

A. granarius

A. rufipes

Fam. Ccrambycidae
Clytus arietis

Fam. Chrysomelidae
Chalcoides fulvicornis . . .

Chrysomela Banksi

Phaedon cochleariae . . .

Ph. tumidulus

Phyllotreta undulata . . .

Fam. Curculionidae

Apion aestivum

A. apricans

A. assimile

A. flavipes

A. hookeri

A. nigritarse

A. ononicola

A. ononis

A. sanguineum {miniatum) .

A. ulicis

A. violaceum

A. virens

Ceuthorrhynchus assimilis

C. hirtulus

C. troglodytes

Clonus alauda

C. scrophulariae

Cneorrhinus exaratus ....

Lisoma deflexum

Otiorrhynchus ligneus . .

O. ovatus

O. rugosostriatus

O. sulcatus

Philopedon plagiatus . . .
Phytonomus ononidis . . .
Ph. posticus {variabilis) . .
Polydrosus confluens ....

I

2

3

4

5

6

7

t

X

s

s

p

b

s

t

m

c

s

p

m

s

t

m

c

s

p

m

s

t

m

c

s

p

m

s

p

m

c

p

o

m

s

P

m

c

p

m

m

s

t

m

c

p

m

hi

s

p

h

t

p

o

m

s

p

h

c

p

o

m

s

p

m

s

p

o

m

s

p

m

c

p

m

m

s

p

m

c

p

m

m

s

p

m

c

p

m

m

s

p

m

c

p

m

m

s

p

m

c

p

m

m

s

p

m

c

p

m

m

s

p

X

s

p

m

m

s

p

X

s

p

m

m

s

p

m

c

p

m

m

s

p

(x)

c

p

m

m

s

p

m

c

p

m

m

s

p

m

c

p

m

b

s

p

m

c

p

o

m

s

p

m

s

p

o

m

s

(p)

(x)

c

p

o

m

s

p

m

c

p

m

m

s

p

m

c

p

o

m

s

t

(x)

c

p

o

b

s

(p)

m

c

p

o

b

s

(X)

c

p

P

b

p

m

c

p

P

b

p

(X)

c

p

P

b

p

m

c

p

P

b

p

X

s

p

P

b

s

p

X

s

p

m

b

s

p

m

c

p

o

(b)

s

p

(x)

c

p

o

b

s

Vide footnote 3, p. 202.

The human transport of animals across the Northern Atlantic

209

Sibinia potentillae . ,
Sitona griseus . . . ,
S. hispidulus . . . .
S. humeralis . . . .
S. lepidus iflavescens)

S. lineatus

S. puncticollis . . . .
S. regensteinensis . .
S. sulcifrons . . . .
Tychius picirostris .

I

2

3

4

5

6

7

p

(x)

c

P

m

s

p

X

s

P

m

s

(p)

(x)

c

P

d

s

p

m

c

P

(m)

s

p

m

c

P

m

s

p

m

c

P

(m)

s

p

m

c

P

(m)

s

p

(x)

c

P

(b)

s

p

m

c

P

m

(b)

s

p

m

c

P

m

s

Summarizing, then: to be judged especially fit to survive transport in ballast
across the Atlantic and to becoming settled in North America, a terrestrial animal,
e.g. an insect, should combine the following six properties: it lives on the ground
(terricolous), has no pronounced moisture requirements (non-hygrophilous), pre-
fers open ground of a waste-place character (at least it is not halobiontic, salt-
demanding), is not dependant on special kind of food (polyphagous), is flightless,
and has a parthenogenetic reproduction. The only species of Coleoptera from the
English ballast-places filling all these requirements are the four Otiorrhynchus
weevils and, sure enough, they all crossed the Atlantic.

It is of considerable interest to illustrate, in form of a diagram (diagram 5), to
what extent the beetles of the ballast places combine the six "favourable qualities"
just mentioned, keeping "emigrants" and the remaining species apart. The differ-
ence is striking: 73 per cent of the "emigrants", but only 46 per cent of the rest,
combine more than 3 of the 6 "favourable qualities".

A similar calculation has been made for Carabid beetles alone (diagram 6),
omitting the mode of reproduction (last column of table 6), because all species
are bisexual, but adding the stage of hibernation (according to Lindroth, 1945,
1949). This may be regarded as having some influence because the transport of
a fuUgrown beetle (above all of a fertilized female) gives the maximum chance for
a successful colonization. Most ships in the Newfoundland trade left England in
the spring (above p. 162) and constant imaginal hibernators were therefore fa-
voured. These make 71 per cent (15 species) of the 21 "emigrants" but 66 per cent
(41 species) of the remaining species; only among the latter there are some species
(6 in number: Amara convexiuscula, A. equestris, Calathus mollis, Dichirotrichus
pubescens, Eurynebria complanata, and Trechus quadristriatus), in which the larval
hibernation is practically without individual exceptions. — Of the Carabidae of

14 — 565597 Lindroth

2IO

CHAPTER II

per cent
SOt

3 4

Ballast fauna
Coleoptera, 242 species

DIAGR. 5. Combination of one to six ecological and biological characters (table 6,
columns 1-3, 5-7; and p. 209) favouring ballast transport, among 242 species of
Coleoptera observed on ballast places in SW England.

Black = "emigrants" (67 species). White = non-emigrants.

the ballast-places, all of the 21 "emigrants" but only 81 per cent (51) of the re-
maining 62 species combine more than 3 "favourable characters", as shown in
diagram 6.

The estimation of characters favouring ballast-transport was also carried out by
comparing the species introduced into Newfoundland with the "non-emigrants"
of the English ballast-places. The Carabid and Curculionid families were selected
for this purpose.

Carabidae planti- hygro- saline macro-

colous philous localities pterous

Introduced Newfoundland species (23) o 5 = 22% o 14 = 61%

"Non-emigrants" of ballast-places (63) 3 = 5% 20 = 32% 14 = 23% 46 = 74%

The difference in the other factors of table 6, as well as in the stage of hiberna-
tion, was insignificant in this case.

Curculionidae planti

colous
Introduced Newfoundland
species (29) 15 = 52 %

"Non-emigrants" of ballast-
places (25) 24 = 96 %

The advantage of a polyphagous choice of food is very striking in this case, and
also the predominance of flightless forms among introduced species.

oligo- or

macro-

monophagous

pterous

bisexual

15 = 52%

13=45%

16 = 55%

25 = 100 %

18 = 72%

25 = 100%

The human transport of animals across the Northern Atlantic

per cent
50 T

211

-nnflj

T SO

2 3 4

Combination of "favourable qualities"
Carabidae, 83 species

DiAGR. 6. Combination of one to five "favourable qualities" (table 6, columns 1-3,
5-6; and p. 209), among 83 species of Carabid beetles observed on ballast places in
SW England.

Black = "emigrants" (21 species). White — non-emigrants.

From the above it can be concluded that the selection of animal species which
managed to cross the Atlantic by ship, first and foremost in ballast, has not been
a random one. Forms associated with cultivated soil, possessing low moisture
requirements, little specialization of food and little power of active dispersal were
always favoured, especially if these properties were combined with parthenogenetic
reproduction.

Eastward transport

The entire preceding part of this chapter has been devoted to the European in-
troductions into North America. Turning to the reverse phenomenon, the Ameri-
can element in the fauna of Europe, we find quite a different picture: this element
plays an inferior role in the European fauna.

On page 125 a summary is given (table i) of the animal species common to
Europe and North America, belonging to certain selected taxonomic groups and
listed on page 17-124. The total number of species is 908. Of these no less than
309-344 species (33-38 per cent) are considered (at least in part) to be introduced
with man from Europe to North America. In comparison herewith, the American
introductions in Europe are quite unimportant, amounting to 27-34 species
(3-4 per cent).

The difference is especially striking in some groups of lower terrestrial animals.
In the case of Carabid beetles, 37-38 species were introduced from Europe to North
America, none in the opposite direction; in Chilopods (Centipedes) 12-13 (o)
species, in Diplopods (Millipedes) 15 (o) species, in Terrestrial Isopods (Woodlice)
21-22 (o) species, in Lumbricids (Earthworms) 15 (2) species. Schubart (1929,

212 CHAPTER II

p. 316) has already drawn attention to this remarkable fact, as far as Diplopods
are concerned.

Thd situation is well-known also to applied entomologists. They noticed that
the number of originally European insect pests in North America is higher than that
of originally American ones in Europe. A few instances of the latter type are
mentioned below (p. 222), the most famous being the Colorado Beetle {Leptino-
tarsa decemlineata Say), the Grape Phylloxera {Phylloxera vitifoliae Fitch), and
the Woolly Aphis [Eriosoma lanigerum Hausm.). "The Fall Webworm" {Hyphan-
tria cunea Drury) has become a potential pest in southeastern Europe. "The
Northern Army Worm" {Leucania unipuncta Haw.), with an almost cosmopolitan
distribution, is also said to be of North American origin.

Still more insignificant is the group of American animals not associated with
agricultural plants which have been unintentionally introduced into Europe and
become naturalized outdoors. For instance, only two cases of this kind have been
observed among the Coleoptera: the Longicorn beetle Neoclytus acuminatus F.
and the small Weevil Stenopelmus rufinasus Gyll. Both are now widely distributed,
the former in the regions surrounding the northern part of the Adriatic Sea, the
latter in western Europe. Both have been imported with their foodplants, the
Neoclytus (arrived before 1850; Miiller, 1949-53, p. 148-149) with American tim-
ber, the Stenopelmus (since 1898) with the water-fern Azolla caroliniana Willd.
which is of North American origin but generally cultivated in Europe as an orna-
ment in ponds and pools.

This is fully in accordance with the geographical character of the European
flora. It has sent hundreds of emigrants west across the Atlantic, which is illustra-
ted above on the Newfoundland flora (p. 146) and the vegetation of the English
ballastplaces (p. 196; vide also Gilmore, 1932), but weeds and other recent in-
troductions originating in North America, completely naturalized and widely
distributed in Europe, are surprisingly scarce. Well-known instances are^: —

Elodea canadensis Michx.^ — Transamerican, N to N. Labr. — Europe (widely,
since 1836), N to N Sweden.

Epilobium glandtdosum Lehm. (incl. adenocaulon Haussk.)^ — Transamerican. —
Europe (widely, since later part of 19th century), N to C Finland.

Erigeron canadense L. — Transamerican. — Europe (generally), N to N Sweden.
Almost cosmopolitan.

Jiinciis tenuis Will, {macer Gray).^ — N. America (generally), also C. & S. America.
— Europe (widely, since before 1850), N to C Sweden. N. Africa. Australia.

^ For further information, vide Jessen & Lind, 1922-23, Hulten, 1950, Fernald, 1950,
and Clapham, Warburg, & Tutin, 1952.

- Probably originally cultivated in Europe and escaped.

^ By some students, e.g. Heslop Harrison (1953), regarded as native in Ireland.

The human transport of animals across the Northern Atlantic 213

Lepidium densiflorum Schrad. — Transamerican (but probably introduced in the
east). — Europe (generally but often casual), N to N Norway. Also in other conti-
nents.

Matricaria matricarioides Less, {discoidea D.C., suaveolens Pursh., nee L.). — Trans-
american, native of NW America & NE Asia. — Europe (generally, since about 1850),
N to northernmost Scand. S. America, New Zealand.

Oenothera biennis L.^ — N. America E of Rocky Mts. — Europe (generally, since
1 8th century), N to C Finland.

Oxalis stricta L. — Transamerican, S to Mexico. — Europe (widely, since before
1700), N to C Finland. Also in other continents.

Veronica peregrina L.i — Transamerican, also C & S America. — W & C Europe
(since i8th century), N to Denmark.

To these should be added some 15-20 species of North American origin, lo-
cally and mostly only temporarily established, especially on the British Islands.
— On Ranunculus cymbalaria Pursh., vide below (p. 244).

This is a poor representation, indeed! It should also be observed that several
of the above plants are late arrivals, apparently introduced after modern steam
trade had commenced.

Turning back to animals, there is only one ecological group in which North
American species unintentionally introduced into Europe are far more numerous
than those transported in the opposite direction: the hothouse animals, more or
less unable to establish themselves out of doors.

European hothouse species of North^ American origin.

A. Terrestrial animals.

Formicidae Helicodiscus parallelus Say

/-, . , 7- 7 , c Opeas pumilum PfeiflF.

Lrematogaster lineoiata bay ^ .

r> ^ • ^ TVT 1 Subulina octona Brug.

ronera opaciceps Nyl. „ . . , ,

Zonitotdes arbor eus Say^

Araneae .

Lumbricidae

Eperigone maculata Banks

Btmastus beddardi Mich.

Mollusca Eisenia carolinensis Mich.

Hawaiia minuscula Binn.

^ Probably originally cultivated in Europe and escaped.

- Some of these may possibly come from parts of America south of the U.S. A. -Mexican
boundary.

* In part established in outdoor conditions.

214 CHAPTER II

B. Freshwater animals.
MoUusca

Gyraulus parvus Say Lymnaea columella Say

Heltsoma dtiryi Weth. L. cubensis Pfeiff.

H. tenuis Phil. Physa ancillaria Say

H. trivolve Say Ph. gyrina Say

Lymnaea catascopium Say Ph. heterostropha Sayi

The corresponding European element in North America is almost non-existent.

The reasons for its poor effect

Judging from the present character of trade and exchange of goods between
Europe and North America, one would perhaps feel inclined to expect that an
unintentional transport of animals and plants across the Atlantic would have been
more easily realized in an eastward direction. There is more material suitable for the
purpose, such as vegetable food, wool, timber, &c., shipped from North America to
Europe than in the opposite direction.

The present conditions, however, are but little pertinent to the matter. It
seems that the main bulk of introduced species, especially of the Europeans
naturalized in North America, arrived prior to modern steam and motor traffic, the
first transatlantic line of this kind being opened in 1840 (Cunard Line: Liverpool-
New York), Earlier records from North America are available for at least the
following European species: —

Sitotroga cerealella 01., the Angoumois Grain Moth, is reported to have been
observed in North America as early as 1743 (Essig, 193 1, p. 824).

The Hessian Fly, Mayetiola {Phytophaga) destructor, was first described by Say
in 1816 and observed on Long Island, N.Y., as early as 1779, but is usually regarded
as a European introduction.

Argyroploce pomonella L., the Codling Moth, was also introduced in the i8th
century.

The Carabid Beetle Harpalus affinis Schrk. {aeneus F.) was described from North
America in 1805 by Beauvois under the name of viridiaeneus, a clear synonym.

Similarly, the father of North American entomology, Thomas Say (i 787-1 834),
described some insects and other arthropods now regarded as of Old World
origin, for instance the Carabid beetle Bembidion tetracolum Say [ustulatum auctt.,
nee L.), the Dung-beetle Aphodius fasciatus 01. (putridus Hbst.; as tenellus Say),
the Milliped Nopoiulus venustus Mein. (as pusillus Say; vide Palmen, 1952, p. 26),

^ In part established in outdoor conditions.

The human transport of animals across the Northern Atlantic 215

the Wood-lice Porcellio spinicornis Say (pictus Brandt) and P. scaber Latr. (the
latter as niger Say; vide Palmen, 195 1, p. 15). Say also mentioned the occurrence
in North America of Armadillidium vulgare Latr. and Oniscus asellus L.

Dejean (1828, p. 466) reports the Carabid beetle Amara aenea DeG. from North
America and this is confirmed by a specimen in his collection (Lindroth, 1955 b,
p. III).

Randall, in 1838, described the Carabid Clivina collaris Hbst. (as elongata;
vide Lindroth, 1954 a, p. 122) and the dung-beetle Aphodius fimetarius L. (as
nodifrons).

One of the Otiorrhynchus (Brachyrhinus) Weevils, the genus as a whole being
introduced into North America (Brown, 1940, p. 67), was early recognized:
sulcatus F., about 1830 (Schwarz, 1890, p. 38; Essig, 1931, p. 189).

The Pear Psylla, Psylla pyricola Foerst., and the Elm Leaf Beetle, Galerucella
luteola Miill., were noticed in the U.S.A. in the 1830's (Essig, 193 1, p. 848-850).

In Newfoundland, between 1827 and 1835, Philip Henry Gosse {vide Bruton,
1930) collected, among other insects, "Forficula sp." (no doubt auricularia L.)i
and the beetles Coccinella bipunctata L., Aphodius fimetarius L., and two species
of Sitona, all introduced.

Finally, according to a collection of insects made by the Norwegian naturalist
Peter Stuwitz in Newfoundland in the period 1 839-1 842 and preserved at the
Zoological Museum, Oslo, at least the following European species reached the
island prior to the opening of steamer connections (with Halifax, in 1842):

Carabidae Scarabaeidae

Agonum millleri Hbst. Aphodius fimetarius L.
A. ruficorne Gze.

Bembidion tetracolum Say Curculionidae

Harpalus affinis Schrk. Barynotus obsciirus F.

Otiorrhynchus ovattis L.

Elateridae O. sulcatus F.

Agriotes lineatus L. Sitona lepidus Gyll. (fiavescens Mrsh.)

It is thus evident that the old trade conditions of sailing vessels days are largely
responsible for the fact that transatlantic introduction of foreign animals and
plants took place almost exclusively in a westward direction. In addition, as fully
discussed earlier in this chapter, the one-sided carrying of ballast westward only,
seems to strengthen the case in a sufficient way.

A slightly different picture is provided by animals imported with their host plants.

^ Generally, Forficula auricularia L. is regarded as an introduction of the 20th century
(Essig, 1931, p. 923).

2l6 CHAPTER II

This traffic is still going on on a large scale and the gradually more rigorous
quarantine will never be able to eliminate completely the undesirable landing of
unexpected and unwelcome passengers.

As far as animals feeding upon the common agri- and horticultural plants of
the temperate zone are concerned, there is a marked preponderance of species
carried west across the North Atlantic (cf. p. 212) which is easily explained
by the greater number of cultivated plants transferred from the Old to the New
World.

In importations connected with hothouse gardening we meet with the reverse
situation: the eastward transport has been strongly dominating. Above all, the
botanical gardens and aquariums of Europe, erected for scientific purposes, have
no doubt been more inclined to enlarge their collections with specimens of the
rich flora of North America than similar institutions of the latter region to obtain
European plants. It should also be observed that European animals, first arrived
as hothouse inquilines into North America, very often were able to spread and
become settled under outdoor conditions, whereas the corresponding element of
European hothouses often arrived from subtropical, or even tropical, parts of North
America and thus as a rule was prevented from leaving its artificial surroundings.
Only species which happened to be introduced from Europe into cold regions,
such as Newfoundland (certain Myriapods and Terrestrial Isopods; p. 108 a.f.)
or the Pacific Northwest (also some species of Carabid beetles; p. 170), have
also remained indoors in North America.

Another explanation

The main bulk of European species, animals as well as plants, which managed
to cross the Atlantic with man and become settled in North America, even on their
native continent demonstrate a high degree of dependence upon human culture.
Most of the plants in question are pronounced weeds and the animals, almost all
members of the lower soil fauna, belong to a corresponding synanthropic element.
Many of the insect species, for instance several Weevils {Curculiom'dae), are directly
bound to Crucifers, Leguminosists, and other weeds.

These plants constitute an old element in Europe, as do the corresponding ani-
mals. The first civilizations of the Occident founded upon agriculture, especially
those of Hither Asia (Assyrians, Babylonians, &c.), were pronounced river cultures,
immediately bordering upon deserts and steppes. Hence man got his most im-
portant culture-plants, the cereals, from the unwooded land but at the same time
many useless plants invaded the arable land from the surrounding steppe and

The human transport of animals across the Northern Atlantic 217

transformed into weeds. They were of course accompanied by insects and other
lower animals feeding upon or otherwise connected with them.

Eventually civilization based on agriculture moved westward, forested land
around the Mediterranean was developed into culture-steppe occupied by culti-
vated plants but also by weeds. Locally, primary steppe of this region contributed
a number of additional weeds, with adherent vermin.

During the Roman age, about the beginning of our chronology, the modern
colonization of Central Europe started, then slowly spread northward. The first
permanent glades were cut out of the continuous forest carpet and invaded, with
the aid of man, by cultivated plants— and weeds. Thus the culture-steppe con-
quered Europe, "step by step", the isolated patches of arable land increased and
fused together. Already at the time of Columbus the coastland of Western Europe
was almost devoid of forests and transformed into an artificial steppe inhabited
by man's constant followers among plants and animals, invited or selfinvited. They
were all ready to embark the ships and, if the voyage was favourable, to settle
wherever they landed in a suitable climate, provided man had prepared the soil.
Similar views, concerning the synanthropic bird fauna, have been express ed by
Lack (1954, p. 200-202).

How different were conditions on the American side when transatlantic trade
was starting! — The high Indian cultures, founded upon agriculture, were restricted
to Central America and parts of the Andes farther south. In northeastern North
America, at least, the tribes were more or less migratory and had hardly progressed
beyond the neolithic stage. A very primitive form of agriculture, with maize and
beans as foremost plants, was practiced by the Iroquois in the St. Lawrence
valley (Coleman, 1930, p. 11). Farther south within the limits of the present U.S.A.,
also along the Atlantic coast, agriculture seems to have been carried on more in-
tensively and sweet potatoes, tobacco, cotton, sunflower, &c., were cultivated.
But nowhere was the plow used. The general custom was to burn over the ground
and to plant some grains of maize &c. into holes dug at short distances with the
aid of primitive shovels, hoes or spades (Hodge, 1907, p. 24-27). Nothing like the
arable land of the contemporaneous Eastern Hemisphere was produced in this way
and though some tribes planted corn in the same spots from year to year, the culti-
vation of the soil seems generally to have been of a more or less casual character,
except among the Pueblos of the present New Mexico and Arizona, where agricul-
ture was the dominating trade, based on irrigation.

Therefore, in North America, the chance of native steppe plants invading per-
manent arable land and transforming into constant weeds was considerably less
than in the Old World. This holds true especially for northern coastal districts,

2i8 CHAPTER II

on the Atlantic as well as on the Pacific side, partly because the Indian agriculture
was very primitive there, partly because there was no direct connection with the
central" prairie enabling steppe plants to invade the artificial clearings. Conse-
quently, when the White Man first arrived and permanent settlement founded upon
agriculture started, there were few indigenous plants present able to intrude as
weeds. This gave free entrance to the corresponding floral element from Europe
which, almost free from competition, soon became dominating on cultivated and
waste ground, above all within the northern half of the Atlantic coast. Around the
ports of Newfoundland and Nova Scotia it is still easy to find acres of land in-
habited by a purely European flora. This region was perfectly prepared then, to
receive and establish the more or less synanthropic members of the European
fauna brought in by trade, but it was very little suited to provide ships destined
east with stowaways of North American origin.

The economic importance of introduced animals

It is a fact, only too well-known, that animals and plants purposely or unintention-
ally introduced into a foreign country often reproduce and spread very quickly
and soon reach an abundance far surpassing the normal conditions in their original
patria. As far as domesticated or otherwise useful animals are concerned this of
course is only to the advantage of man. More numerous, however, are the instances
of weeds, phytophagous insects, plant diseases, &c., which in the new continent
have developed to veritable pests, though many of them in their native country
do not cause much annoyance.

Theoretically, the explanation of such super-normal abundance of introduced
species, animals as well as plants, may be found among any of the following three
groups of facts: —

1. The environment, that is soil, climate, supply of food, &c., may be more
favourable in the new country.

2. There may be more "space" available, the introduced species being able
to colonize and "ecological niche" previously not, or only thinly, inhabited by
indigenous species.

3. The introduced species may have escaped from its native enemies which in
the original patria control its abundance.

The first explanation seems to hold true at least in cases where a new and more
abundant host (plant or animal) is available in the new area. Thus the main
reason for the mass reproduction of the Colorado beetle [Leptinotarsa decemlineata
Say) was that it invaded land (in eastern North America and in Europe) where a

The human transport of animals across the Northern Atlantic 219

new host, Solanum tuberosum, was enormously more abundant than the Solanum
rostratum of its original habitat in the Rocky Mountains.— Similarly, the European
vine (Vitis vinifera) was a more suitable host to the Phylloxera vitifoUae Fitch than
were the corresponding American species.— Species adapting themselves to in-
door conditions, especially in greenhouses, such as many Diplopods, Terrestrial
Isopods (Woodlice), &c., may gain thereby, not only an almost cosmopolitan
distribution, but also, due to an equal and favourable microclimate, a local abun-
dance widely surpassing that of their original outdoor habitat.

However, these are exceptions. It seems reasonable to assume that normally
an animal or plant species is perfectly adapted to climatic and other conditions
prevailing in its natural habitat, that it finds its optimum where it developed (from
a phylogenetical point of view), usually in the central part of its distribution area,
and that a successful introduction by the aid of man into a foreign country means
that it by lucky chance entered a new area with approximately the same, and not
better, environmental conditions. Then this alone cannot be responsible for
subsequent super-abundance.

The second explanation is based on the assumption that "competition" (excluding
the relations between predator or parasite and prey or host) is a major factor
determining the composition of fauna and flora of a given area, especially the assort-
ment of species within a micro-habitat. Without any doubt this is true as far as
sedentary organisms are concerned. Between species and individuals of higher,
autotrophic plants there is a continuous competition, not only for space, but also
for light and nutriment. It seems therefore defensible to explain the colonization
of waste ground in towns and ports of the Canadian East by an almost unmixed
European flora {vide above, p. 218) as at least partly due to the fact that practically
no corresponding ecological element of the indigenous flora was present when the
White Man arrived.

In the second place, mammals and birds, with their higher-developed psychical
functions, especially their instinct for ownership of a wide area ("territory";
Germ.: Revier) around their breeding places, certainly in their normal state live
under permanent (intra- as well as inter-specific) competition which is essentially
difi'erent from the blind struggle for life in most non-sedentary evertebrates. The
effect is that the population of for instance a certain bird species is many times
sparser than if it had been regulated by food supply alone. The rapid increase of
area and abundance of the English Sparrow (Passer domesticus L.) and the Starling
(Sturnus vulgaris L., fig. 4), both purposely introduced into North America, may
very well have been determined by the lack of indigenous competitors among birds
living in the immediate neighbourhood of man.

Turning to the lower terrestrial fauna, the literature contains a good many

220 CHAPTER II

purely theoretical assumptions but very few direct observations of interspecific
competition as a major factor governing distribution. Personally, I feel especially
suspicious of the slogan: "competition between near related species", because,
in this connection, the ecological and not the taxonomical relationship must be
decisive.

Some of the most reliable observations on competition were made on earthworms
(fam. Lumbricidae), implying that "peregrine", i.e. the anthropochorous, more or
less cosmopolitan species, after introduction into a new country, intrude upon the
endemic species of the original fauna or even completely exterminate them over
large areas (Michaelsen, 1903, p. 24; Smith, 1928, p. 347 a.f.). It seems also natural
that the rather permanent burrow system of an earthworm should represent a
sort of individual "territory", avoided by other individuals of the same or ecolog-
ically related species and that a competition about space takes place, similar to
that between sedentary organisms.

Klots (1951, p. 201), Ferguson (1955, p. 188), and others remarked that the
indigenous Pieris napi L. (Green-veined White) of North America has noticeably
decreased in places where the introduced Pieris rapae L. (Small or Cabbage
White) had become established (since i860). It is difficult to imagine a com-
petition for food in this case as napi appears to prefer wild, thinleaved Cruciferous
plants, but rapae the thickleaved, usually cultivated forms (Brassica, &c.) (Petersen,
1954a, p. 195). It seems more reasonable to suggest that the normal courtship
and other sexual activity of the "weaker" species (napi) has been disturbed by
the presence of the very similar rapae; the upper side of the male is almost identical.
This explanation may have relevance for other diurnal insects of great vagility
and guided by optical sense (the scent of the male in the two Pieris species seems
to be quite different; Ford, 1946, p. 97). For nocturnal Lepidoptera the olfactory
sense governing sexual behaviour may be similarly deceived by the intrusion of
a related species with similar smell of the female. Dr. Petersen {in lit.) suggests
that, in the case of the two Pieris mentioned, a higher resistance against parasites
in rapae may be an important factor for its higher viability as compared with
napi. It would perhaps be worth while to investigate whether the purposely in-
troduced Braconid wasp Apanteles glomeratus L. (p. 222) has played a role there.

From my own experience on Carabid beetles, I have already (Lindroth, 1949,
p. 554-560) given at some length the reasons for my deeply felt doubts as to the
validity of "competition" (in the limited sense, i.e. exclusive of predacy and
parasitism) as a major factor determining the abundance or even the existence of
a species in a certain habitat. In the lawns of Halifax, Nova Scotia, the introduced
Amara aenea DeG. was so exceedingly abundant that the population density widely
surpassed that of all Amara species together in any similar spot studied by me

The human transport of antmals across the Northern Atlantic 221

in Europe, and this in spite of the fact that, in abundant species, intra-spGcific
competition (between individuals of the same species) must be considered stronger
than interspecific competition (between the individuals of different though pos-
sibly near related species). In my opinion, terrestrial habitats very seldom and
then for only short periods possess a "saturated" fauna and therefore competition,
in the limited sense here used, hardly constitutes a governing factor for lower
animals without instincts of "ownership", except in cases where they are highly
specialized, for instance in choice of food.

The third explanation (above) contains the most obvious facts contradicting
the opinion that as a rule reduced competition is responsible for the often extra-
ordinarily rapid dispersal of an introduced animal species, above all embodied in
the method of ''biological control'' against this special form of insect pests. This is
based on the assumption that the abundance of a species in its native country is
balanced by natural enemies (predators, parasites and diseases) from which it partly
or entirely escapes in a new area. By careful observations in the original patria,
the most important of these natural enemies may be discovered, artificially repro-
duced and finally released in the infested country.

The best examples of success in biological control of this kind emanate from
the United States. In that country, due to the extraordinarily high number of
introduced culture-plants, the demands for proceedings against imported pests
have been greater than anywhere else, and likewise the economical powers to
carry them out.

The following examples refer to insects unintentionally or purposely exchanged
between Europe and North America and are selected from Howard (1930), Essig
(1931), "Handbuch der Pflanzenkrankheiten" (1949-54), and other current eco-
nomic-entomological literature.

A. Species of European origin

Galerucella luteola O. F. Miill. (xanthomelaena Schrk.) (Elm Leaf Beetle). Intro-
duced as early as the i83o's, now over the main part of U.S.A. and very noxious.
A species of Tetrastichus (fam. Chalcididae), parasiting the eggs in Europe, and a
few Tachinid flies, have been tentatively introduced.

Phytonomus punctatus F. (Clover Leaf Weevil) and Ph. posticus Gyll. {variabilis
Hbst.) (Alfalfa Weevil) {vide map, fig. 6).^ The latter has been successfully con-

^ Flight of Phytonomus posticus has been frequently observed in U.S.A. It is therefore
possible that macropterous homozygotes have been introduced by which the dispersal
would be considerably favoured. In Europe, the species is dimorphic with respect to the
hind-wings.

CHAPTER II

trolled by the Ichneumonid wasp Bathyplectes curculioms Thorns., introduced
from Europe, the former by the fungus Entomophthora sphaerosperma Fres.

Pieris rapae L. (Cabbage or Small White). First recorded in N. America about
1856 (in Quebec), now widely distributed. The Braconid wasp Apanteles glomeratus
L., imported from Europe in 1883, is now well established and useful.

Lymantria {Ocneria, Porthetria) dispar L. (Gypsy Moth). Introduced for purposes
of study and unintentionally escaped about 1869, a severe pest from 1889. A
large number of different enemies were imported from Europe and Japan, among
others the Carabid beetle Calosoma sycophanta L., which has proved useful.

Euproctis phaeorrhoea Don. {chrysorrhoea auctt., nee L.) (Brown-tail Moth).
First recorded in the 1890's (Mass.), injurious since 1897. Controlled by means of
several European parasitic wasps and flies.

Argyroploce {Carpocapsa, Cydia) pomonella L. (Codling Moth). This was in-
troduced into N. America already in the i8th century and is now generally distrib-
uted. Attempts at biological control have met with only moderate success but
recently the Ichneumonid wasp Ephialtes caudatus Ratz., from Europe, has become
established in British Columbia.

Pyrausta nubilalis Hbn. (European Corn Borer). First recorded in 1917 (Boston),
now widely distributed and very disastrous. A special organisation, "The Inter-
national Corn Borer Investigation", has carried out large scale rearing and trans-
port of parasites (Ichneumonids, Braconids) from Europe.

Mayetiola (Phytophaga) destructor Say (Hessian Fly). This Cecidiomyid fly,
although first described from America and known there since the i8th century, is
usually regarded as a European introduction. Attempts at control have been per-
formed with several European parasitic wasps.

Nematus (Pristiphora) erichsoni Htg.^ (Larch Sawfly), injurious in Canada, has
been effectively controlled by the aid of the European Ichneumonid wasp Meso-
letus tenthredinidis Marl.

B. Species of North American origin

Leptinotarsa decemlineata Say (Colorado Beetle). Native of the Rocky Mountains.
First introduced specimens observed in Europe as early as 1876 but not until
about 1920 did it become established, in the Bordeaux region, whence it spread
with terrible rapidity {vide map, fig. 3, p. 136). Attempts at biological control
have had little success but the Carabid beetle Lehia grandis Hentz and a couple

^ On the possibility of Pristiphora erichsoni being indigenous in North America, vide
Coppel & Leius (Can. Entomol. 87. 1955, p. 107-108).

The human transport of animals across the Northern Atlantic 223

of Tachinid flies, all of them imported from N. America, have established them-
selves in France.

Phthorimaea operculella Zell. (Potato Tuber Moth). Probably of American
origin, now almost cosmopolitan. Control by means of several parasitic wasps
was attempted, i.a. a Habrobracon imported from California to France, but without
noticeable success.

Phylloxera vitifoliae Fitch (vastatrix Planch.) (Vine Aphis or Grape Phylloxera).
Indigenous in E N. America, it was introduced into California about 1858, in
Europe not earlier than 1863, but now a terrible pest. No biological control has
succeeded in this case; the insect is kept down by grafting European vine upon
more resistant American roots.

Eriosoma lanigeriim Hausm. (Woolly Aphis). Introduced from N. America before
1800, now generally distributed in Europe (except in the north). Its severe damage
in Europe is accentuated by the fact that it has cancelled the yearly migration be-
tween the fruit-trees and a second host (Ulmus). The Chalcid wasp Aphelinus mali
Hald. was introduced from N. America into France in 1920, has subsequently
become well established in most parts of Europe and elsewhere, and proved very
useful.

The danger of noxious insects being introduced with the continuously increasing
air traffic seems to have been exaggerated. The appearance of a European Grasshop-
per, Roeseliana roeselii Hag., near the airports of Montreal was, however, inter-
preted in this way (Urquhart & Beaudry, 1953).

On the minimum size of a viable population

Students of animal populations often postulate that there exists a minimum
number (a minimum density) of individuals constituting the population, below
which the population (if isolated), or even the entire species, is unable to recover.
They apparently assume that a sufficiently low number of individuals in itself,
regardless of other factors, may be the initial stage of complete extinction.

The picture of an introduced animal (or plant) species colonizing a new continent
is fundamentally opposite (already pointed out by Mayr, 1954, p. 174). There
is no doubt that these rapidly increasing populations in many cases are descend-
ents of one single female, fertilized before, during or after the intercontinental
transport.

We are entitled to maintain that in such cases no "minimum size" of population

224

CHAPTER II

exists, provided that the genetic constitution of the organism is sufficiently in
accordance with the new environment. Eurytopic ("plastic", non-demanding)
species, like the ecological ubiquists and the geographical cosmopolitans, are in
little need of genetic variability, caused either by re-combinations or by mutations
of genes. As has been shown by experiments, the pronouncedly diffuse thermal
preferendum of a eurytopic species (the Carabid beetle Pterostichus nigrita F.;
Lindroth, 1949, p. 50-53) may be explained by the relative insensibility of each
individual, and not by the presence of a genetically heterogenous population.

And how about the parthenogenetic species, such as most Otiorrhynchus weevils
and some related genera, now making their triumphal progress throughout all
temperate parts of the world? Their prevalence over bisexual forms and species
among immigrants is due to the fact that the introduction of one single individual,
regardless of stage, is sufficient for a permanent colonization. Their genome is
rigid, shut out from new combinations, and there is nothing indicating that their
mutation frequency is higher than normal. The genetic constitution of these
successful parthenogenetic forms is apparently suitable and has no need of im-
provement.

Extinction is not first and foremost a question of population density but of
genetic degeneration, accentuated by unfavourable changes of environment {vide
Mayr, 1944, p. 224).

Bibliography of Chapter II

Alfaro, a., 1 941. El escarabejo de la patata (Leptinotarsa decemlineata Say). —
Bol. Patol. Veg. y Ent. Agric. 10. Madrid, p. 39-80.

1943- La invasion del escarabejo de la patata al iniciarse la campana de

1943. — Ibidem. 12. p. 1-8.

Blair, K. G., 193 i. The beetles of the Scilly Islands. — Proc. Zool. Soc. London, p.

1211-1258.
BoCHER, T. W., 1948. Contributions to the flora and plant geography of West

Greenland. I. Selaginella rupestris and Sisyrinchium montanum. — Medd. om

Gronl. 147: 3. Copenhagen, p. 1-26.
BoETTGER, C. R., 1932. Die Besiedelung neu angelegter Warmhauser durch Tiere.

— Zeitschr. Morph. Okol. Tiere. 24. Berlin, p. 394-407.

BoNDESEN, P. & Kaiser, E. W., 1950. Hydrobia (Potamopyrgus) jenkinsi Smith in

Denmark, ilustrated by its ecology. — Oikos. I: 2 (1949). Lund, p. 252-281.
Braddick, L. E., 1953. The Port of Topsham — its ships and shipbuilding. — De-

vonsh. Assoc. Adv. Sci., Lit. & Art. 85. Torquay, p. 18-34.
Brooks, S. T., 1936. The land and freshwater MoUusca of Newfoundland. — Ann.

Carn. Mus. 25. Pittsburgh, p. 83-108.
Brooks, S. T. & B. W., 1940. Geographical distribution of the recent Mollusca of

Newfoundland. — Ibidem. 38. p. 53-75.
Brown, W. J., 1940. Notes on the American distribution of some species of Coleo-

ptera common to the European and North American continents. — Can. Ent. 72.

Guelph, Ont., p. 65-78.

1950. The extralimital distribution of some species of coleoptera. — Ibidem.

82. p. 197-205.

Bruton, F. a., 1930. Philip Henry Gosse's Entomology of Newfoundland. — Ent.

News. 41. Philadelphia, p. 34-38.
Carpenter, G. H., 1895. Notes on a new British beetle. Otiorrhynchus auropuncta-

tus Gyll. — Irish Nat. 4. Dublin, p. 213.
Clapham, a. R., Tutin, T. G. & Warburg, E. F., 1952. Flora of the British Isles.

— Cambridge, pp. I-LI, 1-1591.

Cochrane, J. A., 1938. The story of Newfoundland. — Boston, p. 1-257.

Coleman, A. P., 1930. The geographical and ethnical background. — Cambr. Hist,
Brit. Emp. 6. Cambridge, p. 1-19.

Cosens a., 1923. Reports on insects of the year. — 53rd Ann. Rep. Ent. Soc. Ont.
(Ont. Dept. Agric). Toronto, p. 10.

Darlington, P. J., Jr., 1927. Aegialia arenaria Muls. in New England, with local
records for other species. — Psyche. 34. St. Albans, Vt., p. 98-99.

Dejean, p. F. M. a., 1828. Species General des Coleopteres. III. — Paris (p. 466).

EssiG, E. O., 1931. A history of entomology. — New York, p. 1-1029.

1934- The historical background of entomology in relation to the early develop-
ment of agriculture in California. — Pan-Pac. Ent. 10. San Francisco, p. i-ii.

15 "565597 Lindroth

226 CHAPTER II

Evans, J. W., 1952. The injurious insects of the British Commonwealth. — London,

pp. I-VII, 1-242.
Fall, H. C, 1934. A new name and other miscellaneous notes. — Pan-Pac. Ent. 10.

San Francisco, p. 171-174.
Ferguson, D. C, 1955. The Lepidoptera of Nova Scotia. — Bull. N. S. Mus. Sci.

2. Halifax, p. 161-375.
Fernald, M. L., 1950. Gray's Manual of Botany. 8th ed. — • Cambridge, Mass., pp.

I-LXIV, 1-1632.
Ford, E. B., 1946. Butterflies. — The New Naturalist. London, pp. I-XIV, 1-368.
Fowler, J. J., 1901. A visit to St. Andrews, N. B., with a catalogue of plants

collected in its vicinity. — Proc. Nat. Hist. Ass. Miramichi. 2. Chatham, N. B.,

p. 21-28.
Fuente, J. M., de la, 1918. Catalogo sistematico-geografico de los Coleopteros en

la peninsula iberica, &c. — Bol. Soc. Ent. Espana. i. Zaragoza.
Gathorne-Hardy, G. M., 1921. The Norse discoverers of America. — Oxford,

p. 1-304.
Gavrilov, K., 1935. Contributions a I'etude de I'autofecondation chez les Meta-

zoaires hermaphrodites. — Bull. Assoc. Russ. Rech. Sci. 2. Sect. Sci. Nat. & Math.

Praha, p. 163-216.

1940. Sur la reproduction de Eiseniella tetraedra (Sav.) forma typica. — Acta

Zool. 29 (1939). Stockholm, p. 439-464.

GiLMORE, M. R., 1932. Plant vagrants in America. — Papers Mich. Ac. Sci., Arts

& Lett. 15. Ann Arbor, Mich., p. 65-79.
Gray's Manual of Botany, vide Fernald, 1950.

Gribble, J. B., 1830. Memorial of Barnstaple. — Barnstaple, p. 1-640.
Hackman, W., 1954. The spiders of Newfoundland. — Acta Zool. Fenn. 79. Hel-

singfors, p. 1-99.
Handbuch der Pflanzenkrankheiten (P. Sorauer), 1949-54. 5- d. IV. V. Berlin &

Hamburg.
Harrington, W. H., 1891. Notes on a few Canadian Rhyncophora. — Can. Ent.

23. London, Ont., p. 21-27.
Harris, C. A., 1930. Newfoundland, 1783 to 1867, and 1867-1921. — The Cambr.

Hist. Brit. Emp. VI. Cambridge, pp. 422-437, 673-685.
Hatch, M. H., 1946a. Notes on European Coleoptera in Washington, including a

new species of Megasternum. — Pan-Pac. Ent. 22. San Francisco, p. 77-80.

1946b. Note on introduced species of Carabus in North America. — Bull.

Brookl. Ent. Soc. 41. Brooklyn, N. Y., p. 71.

■ 1949. Studies on the fauna of Pacific Northwest greenhouses, &c. — Journ.

N. Y. Ent. Soc. 57. New York, p. 141-165.

1953- The beetles of the Pacific Northwest. I. Introduction and Adephaga. —

Univ, Wash., Publ. Biol. 16. Seattle, Wash., p. 1-340.
Hodge, F. W. (editor), 1907-10. Handbook of American Indians north of Mexico.

1-2. — Washington, D. C, pp. 1-972, 1-1221.
HoLDHAUS, K., 1927-28. Die geographische Verbreitung der Insekten. — Handb.

der Ent. (C. Schroder). 2. Jena, p. 592-1058.
1954. Die Spuren der Eiszeit in der Tierwelt Europas. — Abh. Zool. Bot.

Gesellsch. 18. Wien, p. 1-493.

Bibliography 227

HoLDHAUS, K. & LiNDROTH, C. H., 1939- Die europaischen Koleopteren mit bo-

reoalpiner Verbreitung. — Ann. Nat. Mus. 50. Wien, p. 123-293.
Howard, L. O., 1930. A history of applied entomology. — Smiths. Misc. Coll. 84.

Washington, D. C, p. 1-566.
HuBENDiCK, B., 1951. Recent Lymnaeidae. Their variation, morphology, taxonomy,

nomenclature, and distribution. — K. Vet. Ak. Handl. (4) 3. Stockholm, p. 1-223.
HuLTEN, E., 1950. Atlas of the distribution of vascular plants in NW. Europe. —

Stockholm, pp. 1-120, 1-512.
IvERSEN, J., 1938. Et botanisk Vidne om Nordboernes Vinlandrejser. — Naturhist.

Tidende. 2: 8. Copenhagen, p. 113-116.
Jeannel, R., 1941-42. Coleopteres Carabiques. 1-2. — Faune de France. 39. 40.

Paris, p. 1-1173.
Jessen, K. & LiND, J., 1922-23. Det Danske Markukrudts Historic. — K. Dansk.

Vid.Selsk. Skr., Nat. Math. Afd. (8) 8. Copenhagen, p. 1-496.
Johnson, W. F. & Halbert, J. N., 1902. A list of the beetles of Ireland. — Proc. R.

Irish Ac. (3) 6. Dubhn, p. 535-827.
Klots, a. B., 195 1. A field guide to the butterflies of North America, east of the

Great Plains. — Cambridge, Mass., pp. I-XVI, 1-349.
Krogerus, H., 1954. Investigations on the Lepidoptera of Newfoundland. I. Macro-

lepidoptera. — Acta Zool. Fenn. 82. Helsingfors, p. 1-80.
Kruger, L., 1899. Insektenwanderungen zwischen Deutschland und den Vereinigten

Staaten von Nordamerika und ihre wirtschaftliche Bedeutung. — (Ent. Verein)

Stettin, pp. I-VIII, 1-174.
KiJNKEL, K., 1916. Zur Biologic der Lungenschnecken. — Heidelberg, p. 1-440.
Lack, D., 1954. The natural regulation of animal numbers. — Oxford, p. I-VIII,

1-343-
Lapouge, V. de, 1908. Tableaux de determination des formes du genre "Carabus".

— L'Echange. (Revue Linneenne.) 24. Moulins (p. 19).
Larambergue, M. de, 1939. Etude de I'autofecondation chez les Gasteropodes

Pulmones, &c. — Bull. Biol. France et Belg. 73. Paris, p. 19-231.
Leng, C. W., 1920, 1927-48. Catalogue of the Coleoptera of America, north of

Mexico. With 5 suppl. — Mount Vernon, N. Y.
LiNDROTH, C. H., 1945, 1949. Die fennoskandischen Carabidae. Eine tiergeogra-

phische Studie. I. III. — Gbgs Vet. Vitt. Samh. Handl. (6) B. 4. Goteborg, pp.

1-709, 1-911.

1954a. Random notes on North American Carabidae. (Coleopt.). — Bull. Mus.

Comp. Zool. III. Cambridge, Mass., p. 117-161.

1954b. Carabidae common to Europe and North America. — Coleopt. Bull. 8.

Washington, D. C, p. 35-52.

1954c. Carabid beetles from Nova Scotia. — Can. Ent. 86. Ottawa, p. 299-310.

i954d. Carabid beetles from eastern and southern Labrador. — Ibidem, p.

364-370.

19546. Experimentelle Beobachtungen an parthenogenetischem und bisexuel-

lem Otiorrhynchus dubius Stroem (Col., Curculionidae). — Ent. Tidskr. 75.
Stockholm, p. 111-116.

1955a. The Carabid beetles of Newfoundland. — Opusc. Ent., Suppl. XII.

Lund, p. 1-168.

228 CHAPTER II

LiNDROTH, C. H., 1955 b. Dejean's types of North American Carabidae (Col.). —
Opusc. Ent. 20. Lund, p. 10-34.

1956. The Irish form of Carabus granulatus L. — Ent. M. Mag. 92. London,

p. 7-8.

Lowe, A., 1951. The plants of Vineland the Good. — The Icelandic Canadian.

10: 2. Winnipeg, p. 15-22.
Malkin, B. & Hatch, M. H., 1953. Colpodes buchanani in Oregon. — Pan-Pac.

Ent. 29. San Francisco, p. 134.
Mayr, E., 1944. Systematica and the origin of species. — (Columb. Univ. Press.).

New York, pp. I-XIV, 1-334-

1954- Change of genetic environment and evolution. — In Huxley, Hardy, &

Ford, Evolution as a process. London, p. 157-180.

Meeuse, a. D. J. & Hubert, B., 1949. The Mollusc fauna of glasshouses in the

Netherlands. — Basteria. 13. Liege, p. 1-30.
MiCHAELSEN, W., 1903. Die geographische Verbreitung der Oligochaeten. — Berlin,

pp. I-VI, 1-186.
MuLDAL, S., 1952. The chromosomes of the earthworms, i. The evolution of poly-
ploidy. — Heredity. 6. Edinburgh, p. 55-76.
Naess, a., 1954. Hvor la Vinland? (The location of Vinland.) — Oslo, p. 1-246.
Needham, J. G. & Westfall, M. J., Jr., 1955. A manual of the Dragonflies of North

America (Anisoptera). — Berkeley & Los Angeles, p. 1-615.
Newton, A. P., 1930. Newfoundland, to 1783. — Cambr. Hist. Brit. Emp. VI.

Cambridge, p. 121-145.
Oekland, F., 1927. Einige Argumente aus der Verbreitung der nordeuropaischen

Fauna mit Bezug auf Wegeners Verschiebungstheorie. — Nyt Mag. f. Naturv.

65. Oslo, p. 339-367-
Omodeo, p., 1952. Cariologia dei Lumbricidae. — Caryologia. 4. Pisa, p. 173-

275-
Palmen, E., 1 95 1, a survey of the Oniscoidea (Isopoda Terr.) of Newfoundland. — ■
Ann. Zool. Soc. Zool. Bot. Fenn. Vanamo. 14. Helsingfors, p. 1-27.

1952. Survey of the Diplopoda of Newfoundland. — Ibidem. 15. p. 1-31.

1954- Survey of the Chilopoda of Newfoundland. — Arch. Soc. Zool. Bot.

Fenn. Vanamo. 8. Helsingfors, p. 131-149.

Palmer, M. G., 1946. The fauna and flora of the Ilfracombe district of North

Devon. — Exeter, p. 1-266.
Parkinson, C. N. (editor), 1948. The trade winds. A study of the British overseas

trade during the French Wars 1793-1815. — London, p. 1-336.
Pearce, E. J., 1926, 1927. A list of the Coleoptera of Dorset. — Dors. Nat. Hist. &

Antiq. Field Club. Dorchester, pp. 1-78, 1-7.
Perrot, J.-L., 1939. Contribution a I'etude de la fecondation chez les Pulmones

Stylommatophores. Le cas de I'Arion empiricorum Fer. — Bull. Biol. France et

Belg. 73. Paris, p. 408-432.
Petersen, B., 1954a. Egg-laying and habitat selection in some Pieris species. — Ent.

Tidskr. 75. Stockholm, p. 194-203.

1954b. Some trends of speciation in the cold-adapted Holarctic fauna. — Zool.

Bidr. 30. Uppsala, p. 233-314.

Prowse, D. W., 1895. A history of Newfoundland. — London, p. 1-742.

Bibliography 229

Reeves, G. I., 1927. The control of the Alfalfa Weevil. — Farmers' Bull., U. S.. Dept.

Agric. 1528. Washington, D. C, p. 1-22.
Rogers, J. D., 191 1. Newfoundland. In: C. P. Lucas, A historical geography of the

British colonies. V: 4. — Oxford, p. 1-274.
Rouleau, E., 1949. Enumeratio plantarum vascularum Terrae-Novae. — Contr.

Inst. Bot. Univ. 64. Montreal, p. 61-83.
ScHUBART, O., 1929. Zur Diplopodenfauna einer Weltstadt (Berlin). — Zool. Anz. 85.

Leipzig, p. 303-316.
ScHWARZ, E. A., 1890. On Otiorhynchidae. — Insect Life (U. S. Dept. Agric). 3.

Washington, D. C, p. 37-38.
Smith, F. 1928. An account of changes in the earthworm fauna of Illinois and a de-
scription of one new species. — Bull. 111. State Nat. Hist. Surv. 17. Urbana, 111.,

P- 347-362.
Strand, A., 1951. The Norwegian species of Amischa Thoms. (Col. Staph.). —

Norsk Ent. Tidsskr. 8. Oslo, p. 219-224.
SuoMALAiNEN, E., 1953. Die Polyploidie bei den parthenogenetischen Riisselkafern.

— Verb. Deutsch. Zool. Gesellsch. (1952). Zool. Anz., Suppl. 17. Leipzig, p.

280-289.
Ulbrich, J., 1930. Die Bisamratte. Lebensweise, Gang ihrer Ausbreitung in Europa,

wirtschaftliche Bedeutung und Bekampfung. — Dresden, p. 1-137.
Urquhart, F. a. & Beaudry, J. R., 1953. A recently introduced species of European

grasshopper. — Can. Ent. 85. Ottawa, p. 78-79.
Valle, K. J., 1955. Odonata from Newfoundland. — Ann. Ent. Fenn. 21. Helsing-

fors, p. 57-60.
Van Dyke, E. C, 1945. A review of the North American species of the genus Carabus

Linnaeus. — Ent. Amer. 24. Brooklyn, N. Y., p. 87-137.
Walker, E. M., 1953. The Odonata of Canada and Alaska, i. — Toronto, p. 1-292.
Wing, L., 1943. Spread of the Starling and English Sparrow. — The Auk. 60.

Lancaster, Pa., p. 74-87.
0KLAND, vide Oekland.

Chapter III

THE TRUE, PRE-HUMAN RELATIONSHIP BETWEEN THE
PALAEARCTIC AND NEARCTIC FAUNAS

The so-called Amphiatlantic species

The word "Amphiatlantic" (German: amphiatlantisch) was proposed by 0kland
(1927, p. 352) in order to cover animal (and of course, also plant) species common
to Europe and North America but absent from Asia and thus believed to form
instances of direct faunal exchange across the North Atlantic. The word would
be equally suitable for species common to Africa and Central or South America
(if any) but is here used in the same sense as by 0kland, for the northern continents
only.

A true Amphiatlantic species occurs only in Europe (possibly also in the African
and Asiatic parts of the Mediterranean region) and the eastern parts of North
America. If the word "Amphiatlantic" is used as a historical, not merely as a
geographical concept, implying that a species so termed took part in a direct
faunal exchange across the North Atlantic, then even animals and plants with
a wider distribution, towards the east in the Old World, to the west in North
America, may be included, provided a pronounced gap can be demonstrated
somewhere in the continents surrounding the North Pacific.

However, it is safer, at least for the moment, to keep such historical aspects
out of the picture. The main question would then be to judge whether a seem-
ingly Amphiatlantic distribution, in the restricted sense of the word, is an un-
questionable reality or only due to incomplete knowledge of the species' area.
The latter possibility must always be earnestly considered since the fauna, not
only of northern Asia but also of Western North America, is far from sufficiently
investigated in many groups of animals.

This reservation cannot, however, conceal the fact that a long series of animal
species, belonging to different taxonomical and biological groups, show a perfectly
Amphiatlantic distribution, as is amply exemplified in the list of Eur-American
species (p. 17-124). Many of these have already been treated in the preceding
chapter of this book and their crossing of the Atlantic was proved (or at least
considered) as a quite late event, caused by introduction with man, mainly in the
direction from Europe to North America.

The relationship between the palaearctic and nearctic faunas

231

This is, however, only part of the explanation of the Amphiatlantic type of
distribution though it apparently holds true in the majority of cases, as far as
terrestrial Evertebrates are concerned. Putting these aside, there still remains
to be considered a good number of animal species which are— or may be— old,
prehuman inhabitants of the countries bordering the North Atlantic but are absent
from northern and eastern Asia as well as the Pacific side of the American continent.
Let us call them the primary Amphiatlantic species.

The following are good illustrations of this geographical group: —

A. Primary Amphiatlantic species or subspecies lacking in Asia (except possibly
in Novaja Zemlja) as well as in Pacific North America.

Aves

Alca tarda L.

Catharacta skua Briinn.^

Cepphus grylle L. (except sbsp. mandti

Mandt)
Charadrius hiaticida L.,/. typ. (incl.

sbsp. psammodroma Sal.)
Fratercula arctica L.
Fulmarus glacialis L.,/. typ. (fig. 27)
Larus mar inns L.
Morns (Sida) bassanus L. (fig. 28)
Phalacrocorax carbo L.,/. typ.
Sterna dougalU Mont.,/, typ.
Thalasseus sandvicensis Lath.
Uria aalge Pont.,/, typ.

Lepidoptera

Anarta lapponica Thunb. W^ (fig. 30)
Crymodes exidis Lef. (fig. 38)
Plusia (Autographa) parilis Hbn. W
(fig- 31)

Hymenoptera, Formicidae

Ponera coarctata Latr. (fig. 35).

Araneae

Agyneta cauta Cambr.
Crustulina sticta Cambr.
Leptyphantes minutus Blackw.
Monocephalus parasiticus Westr.
Rhaebothorax borealis Jacks. W
Walckenaera vigilax Blackw.

Coleoptera, Staphylinidae

Micralymtna marinum Strom (fig. 32)

B. Amphiatlantic species or subspecies differing from group A only in that
they extend into Western Asia.

Aves

Erolia maritima Briinn.
Rissa tridactyla L.,/. typ.
Sterna hirundo L.,/. typ.
Uria lomvia L.,/. typ.

Araneae

Bathyphantes nigrinus Westr.
Phlegra fasciata Hahn

^ Perhaps no permanent resident of North America.
* W = "Westarctic" species (vide p. 237).

232

CHAPTER III

FIG. 27. Breeding area (about 1950) of the Fulmar, Fiihnariis
glacialis L. — Solid dots =^ forma typica, the Atlantic Fulmar.
Pointed dots = sbsp. rodgersii Cass., the Pacific Fulmar. — The
bird is steadily spreading in western Europe.

(After Fisher, 1952.)

(From Rosenberg, Faglar i Sverige.)

The relationship between the palaearctic and nearctic faunas

233

(From Rosenberg, Faglari Sverige.)

FIG. 28. Breeding area of the Gannet, Mortis (Sula) bassaniis L., about 1940. Size
of dots in four categories, indicating colonies of less than 100 pairs, 100-1,000 pairs,
1,000 to 10,000 pairs, and more than 10,000 pairs, respectively.
(After Fisher & Vevers, 1943.)

C. Less pronounced Amphiatlantic species, lacking in Asia (as far as known) but
Transamerican.

Lepidoptera

Anarta leucocycla Stdgr. W
A. melanopa Thunb.

Araneae

Cbibiona norvegica E. Str. W
C. trivialis C. L. Koch
Dictyna annultpes Blackw. W

Diplocentria bidentata Em.
Gnaphosa or lies Chamb. W
Islandiana alata Em. W
Leptyphantes complicatiis Em.
Pardosa saltuaria L. Koch
Pocadicnemis piimila Blackw.
Thanatus striatus C. L. Koch

Other Arthropods may belong here but the absence of records from Asiatic
Russia is possibly due to insufficient knowledge. Instances zrt:—Cidaria ruberata
Frr. and Eupithecia gelidata Moeschl. (Lepidoptera), Aeschna subarctica Walk.
(Odonata).

Doubtful cases

In the above lists (A-C) of more or less pronounced Amphiatlantic animals a few
species have been omitted which were regarded as primary Amphiatlantic by

234

CHAPTER III

several students, whereas others supposed that human transport has been respons-
ible for their occurrence in North America.

This applies to two ants, Tetramorium caespitum L. and its parasite, Anergates
atratulus Schenck. Wheeler, and others, assumed that the Tetramorium, and con-
sequently also its parasite, had been introduced from Europe with the early colo-
nists and both species are still regarded as adventive by Smith (195 1, p. 823).
This opinion was strongly opposed by Creighton (1950, p. 289): —"The discovery
of Anergates in America rules out the possibility that caespitum might have been
imported to this country"; and (I.e., p. 244):— "—I cannot agree that the first
advent of caespitum on this continent is a result of importation. To do so implies
that Anergates has also been imported. I believe that it can be demonstrated that
the probability for this having occurred is too remote to be credible."

It seems to me that Creighton has not sufficiently considered the nature of
ballast transport. If ballast was taken in great quantities from heaps of rubbish
in a port {vide pp. 165, 200) it is not at all unlikely that a whole Tetramorium nest,
with inquilines and parasites, might have been brought on board. Though scat-
tered and disturbed during the process, it might very well have reorganized itself
during the long journey (p. 162) and become unloaded with the ballast on some
North American shore after arrival. It should also be remembered that the myr-
mecophilous Isopod (Woodlouse) Platyarthrus hoffmanseggi Brandt apparently
reached North America in the same way. Yet, Anergates atratulus is so rare in
Britain that it was probably carried across from the European mainland.

A much more intricate case is that of the snail Cepaea {Helix) hortensis O. F. Miill.
(fig. 29). This was likewise long regarded as a European introduction into North
America {vide Johnson, 1906, p. 79-80) though, after it had been found in "pre-
Columbian kitchen-midding deposits", Pilsbry (1894, p. 321) suggested that the
importation might have taken place with the Vikings of the eleventh century.
Later, subfossils of proposed greater age were recorded:— "in the glacial Pleisto-
cene of Maine" (Dall, 1910, p. 20); "in a shell heap— associated with bones of the
large extinct mink— Mustela macrodon Prentiss.", on an island in Penobscot
Bay, Maine (Johnson, 1915, p. 131); "in a prehistoric shell-heap on Mahone Bay,
about 75 miles west of Halifax, N. S." (Wurtemberg, 1919, p. 71). Since this in-
formation was published, conchologists and zoogeographers have almost unani-
mously accepted Cepaea hortensis as indigenous in North America (Scharff, 1909,
p. 18-20; 191 1, p. 39; Taylor, 191 1, p. 361-362; 0kland, 1927, p. 342 a.f.). Pilsbry
(1939, p. 8) also changed his view but left open the question of whether the present
separated occurrences of the species should be regarded as remnants of a wide
Holarctic area, or if they should be explained as being the result of migration
along a former transatlantic land-bridge, as advocated by Scharff. Wegener (1929,

The relationship between the palaearctic and nearctic faunas

235

FIG. 29. Distribution of Cepaea {Helix)
hortensis O. F. Miill. — Open circles =
doubtful or accidental occurrence.

(According to Johnson, 1906, Taylor, 191 i,

0KLAND, 1925, 1927, PiLSBRY, 1939, &C.)

(From PiLSBRY, 1939.)

p. 105) regarded the distribution of Cepaea hortensis as one of the strongest zoo-
geographical facts in favour of his theory of continental drift {vide below, p. 284).

It seems to me that the solution of the enigma "introduced contra native" in the
case of Cepaea hortensis in North America stands and falls with the dating of the
subfossil remains in Maine and Nova Scotia quoted above. The only published
statement possibly allowing this, as far as I know, is the observation that in Penob-
scot Bay the shells were found associated with bones of an extinct species of mink.
This has apparently impressed subsequent students of the question with the assumed
high age of the Cepaea. The known history of Mustela macrodon Prentiss., however,
is very disappointing in this respect (Anderson, 1946, p. 192-193):— "According to
Hardy the animal became extinct in Maine about the year i860. Traditionally
said to have been commonly trapped along the coast of the Bay of Fundy in southern
New Brunswick, — ". Until further, reliably dated subfossils of Cepaea hortensis
are discovered, nothing seems to prevent those hitherto found being of "post-
Columbian" age.

236 CHAPTER Iir

The following facts may be regarded as indicators of late arrival with man of
Cepaea hortensis into North America: —

(a) The restriction of area in North America to the sea-shore of the Northeast.
There is only one old, unlocalized inland record, from northern Vermont, by A.
Binney {vide Johnson, 1906, p. 74), which requires confirmation. In Europe (map,
fig. 29), Cepaea hortensis is not at all restricted to coastal regions, nor to districts
showing an oceanic climate. Attempts to explain its absence from inland localities
as due to unsuitable edaphic and other conditions (Johnson, 1906, p. 80; Pilsbry,
1939, p. 8) are not in the least convincing since the species in Europe shows no
tendency to fastidiousness in choice of soil; it cannot be termed "calciphile",
more than all shell-bearing snails are, though Pilsbry (I.e.) does so. The restricted
distribution in North America has no doubt historical reasons.

(b) The frequent occurrence of Cepaea hortensis in North America on small
islands off the coast has been used as an argument against the theory of introduc-
tion. Actually, this argues in favour of the theory. Brooks & Brooks (1940, pp.
59, 61) found a couple of European snails (Limax marginatus Miill. and Vertigo
alpestris Aid.) on small, uninhabited islands off the coast of SE Newfoundland and
likewise regarded this as a sign of indigenous occurrence, but it is given a far more
natural explanation by the assumption that sailing vessels discharged their ballast
on these islands, especially at low tide, before entering the port of destination. As
mentioned above (p. 157), tipping of ballast into the sea was prohibited in old days.

(c) The oldest record of living Cepaea hortensis from North America seems to
be from St. Pierre-Miquelon, in 1822 (Taylor, 191 1, p. 366); from Quebec in
1829, New England (Mass.) in 1837 (Johnson, 1906). Though this gives no evi-
dence as to time and place of a possible first introduction, it suggests the New-
foundland region as the primary immigration area. The species was also unmen-
tioned in two lists of New England shells from 1833 (according to Johnson, 1906,

P- 73)-

(d) A. Binney (according to Johnson, I.e., and Taylor, I.e.) observed that on the

small Salt Island off Gloucester, Mass., Cepaea hortensis in 1837 occurred exclusively
in the unhanded variety, but in 1851 he reported that the banded variety was
"not uncommon" on the same island. This indicates an instability of the popula-
tion contradicting the idea of old age on the spot in question. The shell char-
acter mentioned has a hereditary base and the factor "unhanded" behaves as
a dominant (according to the well-known breeding experiments by A. Lang;
vide also Boettger, 1950). Apparently Salt island was originally colonized by
snails with unhanded shells, in part heterozygotes.

(e) Cepaea hortensis, if native in North America, would display a quite unique
type of distribution. It would constitute the single Amphiatlantic animal species.

The relationship between the palaeartic and nearctic faunas 237

the distribution of which could not be explained either by active or passive oversea
dispersal or as a relict of a former circumpolar area.

Besides Cepaea hortensis, 0kland (1927, p. 353-355) regards three species of
terrestrial Arthropods as primary Amphiatlantic: the Wood-louse Oniscus asellus
L. and two Ground-beetles of the genus Carabus, nemoralis Miill. and problema-
ticus Hbst. {catenulatus auctt.). The two first mentioned, without any doubt, are
European introductions into North America; the proposed endemic North Ameri-
can form of Carabus nemoralis is a fiction {vide Lindroth, 1955 a, p. 27), and the
records of Carabus problematicus, as "sbsp. californicus Mtsch.", from western
North America are based on specimens with wrong locality labels {vide Lindroth,
1954a, p. 46).

There are a few more Eur- American animals, belonging to different taxonomical
groups, which have a more or less pronouncedly Amphiatlantic distribution but
which may be explained either by insufficient knowledge of the actual area and/or
by early introduction with man into North America. Instances are the Click-
beetle {Elateridae) Corymbites sjaelandicus Miill., the Moth {Noctuidae) Amphipyra
tragopogonis L., the Spiders Gonatium rubens Blackw. and Haplodrassus signifer
C. L. Koch, and the Snail Vallonia excentrica Sterki. These are to be found among
species preceded by a cross in brackets (f) in the list given in Chapter I.

"Westarctic" species

The concepts "Westarctic" and "Amphiatlantic" overlap, and certain members
of the first-mentioned group were enumerated above (and marked with a "W")
among the Amphiatlantic species.' The name "Westarctic" was, however, given
from a Scandinavian point of view^ in order to cover species of a northern or
alpine character with affinities towards the west, to North America. The word was
created by Th. Fries (1913) on a botanical basis and was soon applied to insects
{Lepidoptera) by Wahlgren (1919). For more recent views on Westarctic plants,
the reader is referred to Hansen (1929, p. 224 a.f.), Nordhagen (1935, p. 143 a.f.),
and Nannfeldt (1940, p. 39).

The definition of a Westarctic plant or animal is that its occurrence in north-
western Europe (Fennoscandia) is cut off towards the east but more or less con-
nected with an area in North America by means of occurrences in Greenland, pos-
sibly also in Iceland and/or Spitzbergen. The concept i not merely a descriptive
one, it contains implicitly the idea that these organisms have reached Fennoscandia

^ In North America this group would be termed "Eastarctic"!

238

CHAPTER III

FIG. 30. Distribution of Anarta (Sympistis)
lapponica Thunb., a "Westarctic" moth. The
figured specimen is from Scandinavia.

(Photo P. Ardo.)

from a western direction across the North Atlantic and it is usually thought that the
dispersal was facilitated by one or two now submerged land-bridges. This hypo-
thesis was strongly opposed by Hulten (1937, e.g. pp. 126-127, 137; 1950, p. 52).

The reasons why a Westarctic species is not always an Amphiatlantic species,
and vice versa, is that the former may occur in Pacific North America and northeast-
ern Asia (E. of River Lena) but not in the mountains of Central Europe; further-
more it is supposed to show clear cold-adaption. However, the essentials of the two
groups, above all the historical interpretation, are common to both. So it would
seem more appropriate to deal with them in the future under one heading. If so,
the term "Amphiatlantic" is much to be preferred because it is universally under-
stood. It would be easy to make subdivisions: into more or less pronounced Amphi-

The relationship between the palaearctic and nearctic faunas

239

FIG. 31. Distribution of Plusia (Autographa)
parilis Hbn., a "Westarctic" moth. The fig-
ured specimen is from Scandinavia.

(Photo P. Ardo.)

atlantic species (according to the size of gap on both sides of the Pacific; groups
A-C above) on the one hand, into northern, arctic, boreo-alpine, &c., Amphiatlantic
species on the other.

Westarctic animals are much fewer in number than plants are. Some of the more
pronounced were enumerated above (p. 231) (and marked with a "W"). According
to the definition, that is with allowance for an occurrence in eastern Siberia, the
following are also instances of Westarctic distribution: —

Lepidoptera

Anarta fiinebris , Hbn.
A. melaleuca Thunb.
A. zetterstedti Stgr.

Anomogyna laetabilis Zett.
Arctia quenseli Payk.
Cidaria sabini Kby.

240 CHAPTER III

Colias nastes Bsd. S copula frigidaria Moeschl.

Hillia iris Zett. Spaelotis clandestina Harr.

Plusia, (Autographa) diasema Bsd.

Araneae

Bathyphantes puUatus Cambr. Hilaira frigida Thor.

Estrandia grandaeva Keys.

Apart from the inappropriateness of using the name "Amphiatlantic" for species
at the same time "Amphipacific", there are three reasons for excluding species
of this type from the group. — In xhe first place, it is usually impossible to declare
for certain that a considered species is actually lacking in Siberia west of the Lena
River; for instance this gap exists, according to our present knowledge, in the
otherwise probably circumpolar Carabid beetles Diachila arctica Gyll. and Elaphrus
lapponicus Gyll.— In the second place, a gap of distribution (also described, for
Lepidoptera, by Kusnezov, 1935) within the large West-Siberian plain, so different
from the mountainous region east of Yenisei, may be due to lack of conditions of
life; the limit between these regions corresponds to a rather sharp fauna! limit
{vide for instance Johansen, 1955). — In the third place, the western parts of
Siberia during Pleistocene time, and probably more than once, have been sub-
merged under water (Piroznikov, 1937) and an originally equal area of distribution
through northern Eurasia may have become interrupted thereby.

These objections do not prevent particularly well investigated representatives
of this somewhat dubious type of Westarctic distribution being used as arguments
in the discussion of transatlantic faunal connections.

The American element of the British flora

The existence of a truly American element in the flora of Ireland, to a lesser extent
in western Scotland and in Wales, has long attracted the attention of botanists.
Though no quite corresponding faunal element peculiar to the British Islands is
known, the phenomenon has such a clear bearing upon our problem of possible
transatlantic connections that it should be briefly touched upon here.
The plant species involved are:

Eriocaulon septangulare With. W. Irel. (widely); Inner Hebrides (Skye

& Coll).
Limosella siihidata Ives S. & N. Wales.

Myriophyllum alterniflorum americanum N. E. Ireland.
Pugsl.

The relationship between the palaearctic and nearctic faunas 241

Potamogeton epihydrus Raf. Outer Hebrides (South Uist).

Sisyrinchium angustifolium Mill. W. Ireland (widely).

Spiranthes romanzoffiana Cham. N. E. & S. W. Irel.; Inner Hebr. (Coll

& Colonsay), W. Argyllshire (Scotl.).

In North America these species have a wide distribution, all reaching New-
foundland in the east, the Spiranthes even central Labrador; the Sisyrinchium
goes to Anticosti Island and SE. Labrador.

The "American Group" has been carefully studied by Praeger (1932, 1934)
and, most recently, by Heslop-Harrison (1953). For some unknown reason, he
does not mention the LimoseUa but his list contains Juncus tenuis Willd. and J.
Dudleyi Wieg., as well as Najas flexilis Willd. Juncus Dudleyi was included tenta-
tively only, as probably introduced, and the same, according to other students
(Clapham, Tutin & Warburg, 1952, p. 1244), applies to J. tenuis (cf. p. 212).
Concerning Najas flexilis, known from several scattered stations on the European
mainland and widely distributed there during inter- and early postglacial times,
Heslop-Harrison himself admits that it is no more entitled to be listed in the
"American" group than for instance Lobelia dortmanna L.

Five of the six species above (all except Sisyrinchium) are aquatic or grow on
very wet places. Water-plants have often been brought by man, purposely (as
the water-fern Azolla caroliniana Willd.) or unintentionally (as Elodea canadensis
Michx.), from North America to Europe, but there seems to be no reason for
a similar explanation of the six American plants in the British Isles. By means
of pollen-analysis, it has even been demonstrated by Jessen (1949, p. 202)
that Eriocaulon septangulare was growing in Irish pools already in early Atlantic
time.

One animal organism, the fre&h-water Sponge Heteromeyenia ryderi Potts, has
a distribution quite similar to that of the above plants (Arndt, 1928). Its main area
is in eastern North America, from Florida to Nova Scotia and Newfoundland,
it is widely distributed in Ireland (Stephens, 1920) and also found on the isle of
Mull in the Inner Hebrides, as well as on the Faeroes (Sparck, 1934). Any idea of
human transport of Heteromeyenia ryderi from North America to its outposts in
western Europe seems as out of touch with reality as for the corresponding plant
species.

The possible explanation of the presence of an American floral (and faunal)
element on the British Isles (and the Faeroes) will be discussed in the next para-
graph.

16 — 565597 Lindroth

242 CHAPTER III

The chance of spontaneous transatlantic dispersal at the present time

We have now a selection of species, mostly animals but also some plants, which
more than any others are expected to show evidence of a direct exchange between
Europe and North America, without the interference of man.

In order to estimate their evidence it is necessary to scrutinize the animals
more concretely, so to speak more "individually", than is possible on the basis of
the taxonomic and purely geographical arrangement of the preceding paragraphs.
They should be grouped according to biological properties which may affect
their powers of dispersal.

In this respect, the following types are represented: —

I. Actively flying animals. All the birds (16 species or subspecies) and all the
Lepidoptera (16 species), referred to the Amphiatlantic or in widest sense "Westarc-
tic" groups. Yet the different taxonomical and biological groups of birds and
Lepidoptera are unequally represented.

The Amphiatlantic birds are all excellent flyers and closely connected with the
sea, 14 are Swimmers and both of the remaining (Charadrius hiaticula and Erolia
mariiima) are Waders breeding on the seashore. For most of them the Atlantic
breeding area is almost continuous, with stations on the Faeroes, in Iceland, Green-
land, &c. (fig. 27). Exceptions are, in the first place the three species of Terns
{Sterna dougalli, S. hirundo, Thalasseus sandvicensis) with a more southerly breeding
area, but the Terns belong to the veritable gipsies of the Atlantic Ocean {vide, for
instance, KuUenberg, 1946); in the second place Morus bassanus (fig. 28) which
is lacking in Greenland, and Catharacta skua, only doubtfully recorded as breeding
there.

The small breeding area of Charadrius hiaticula in arctic North America (inch
Greenland) is historically connected with the eastern side of the Atlantic, as dem-
onstrated by its route of migration along the European coast. The main popula-
tion of Erolia maritima even in Greenland stays over the winter.

It cannot be denied that all Amphiatlantic birds are easily able to cross
the Atlantic or that individual birds actually do so. The Swimmers especially
are able to rest and take food on the open sea. Continents more than oceans
constitute obstacles to their distribution. No inland birds and no weak flyers
are represented among Amphiatlantic birds. The Passeriformes are completely
lacking.

The 16 species of Lepidoptera include 13 Noctuid moths (fam. Noctuidae), that
is a good 80 per cent, 2 Geometers (fam. Geometridae; Cidaria sabini and Scopula
frigidaria), and one Butterfly {Colias nastes). These figures are not at all in accord-

The relationship between the palaearctic and nearctic faunas 243

ance with the proportions displayed by the faunas of countries bordering the North
Atlantic;!

Macrolepido- Butterflies Noctuid Geometrid

ptera, total (Diurna) moths moths

Newfoundland (Krogerus, 1954) 280 38 = 14% 126 = 45% 89=32%
Greenland (Henriksen, 1939) 29 4=14% 17 = 59% 6 = 21%
Iceland^ (Lindroth, 1931) 28 — 14 = 50% 12 = 43%
Norwayi (Haanshus, 1933) 720 96=13% 257 = 36% 237 = 33%
British Islesi (Kloet & Hincks, 1945) 850 66= 8% 336 = 40% 285 = 34%
Amphiatlantic and "Westarctic" spe-
cies 16 1= 6% 13 = 81% 2=13%

Evidently, Noctuid moths are strongly over-represented among Amphiatlantic
(and geographically related) Lepidoptera. The explanation is no doubt that the
members of this group are superior to other Lepidoptera in the art of flying, v^^ith
the exception of the Hawk-moths (fam. Sphingidae) which, however, are no resi-
dents of arctic and subarctic regions. The Anartas numbering no less than 6
of the 13 species, are extremely rapid flyers, on the wing both night and day,
though, it must be admitted, their perseverance of flight has not been investigated.
Also, among the Butterflies, the genus Colias contains some of the most capable
flyers; especially nastes and hecla, almost confined to the arctic region, are able
to withstand even rather strong winds.— No records were available to me of the
flight powers of the two Geometers concerned.

Crymodes exulis (fig. 38) is the member reaching farthest east of a series of
three Noctuids of American origin which via Greenland have proceeded at least
as far as Iceland on their air-borne advance (p. 254).

An instance of at least partly active dispersal across the North Atlantic is formed
by the famous Monarch Butterfly {Danaus plexippus), a North American species
regularly appearing in Britain in spite of the fact that its food plant (Asclepias)
is not indigenous there {vide Ford, 1945, p. 157-159).

2. Animals especially suited for passive aerial transport. This applies to all of
the Araneae listed above, 21 in number, which, at least as young, disperse them-
selves by "ballooning". Even if it is admitted that our present knowledge of the
distribution of spiders is far from complete, implying that certain of the listed
species actually belong to a more or less circumpolar type (but also that others, not

^ Some of the figures are not quite up to date but this hardly affects the records of per-
centage.

2 I have later received a revised list of Icelandic Lepidoptera by Mr. Niels L. Wolff,
Hellerup (Denmark), which, however, does not essentially alter the percental figures given
above.

244 CHAPTER III

considered, should belong here!), the rich representation of the Araneae among ani-
mals with an Amphiatlantic (or similar) distribution is quite striking. It is difficult
to get away from the idea that this is a consequence of their special mode of
dispersal. And it is well worth mentioning that one of the species in question,
Walckenaera vigilax, was among the spiders included in the famous collection of
"aerial plankton", made by Glick (1939) from air-planes in the U.S.A., 3 speci-
mens captured at elevations of from 1,000 to 3,000 feet above the ground
(Crosby & Bishop, 1936).

3. Flightless animals living in close connection zvith seawater and therefore liable
to become dispersed by active swimming or passive transport by currents and
waves. The single instance among the animals above is the Staphylinid beetle
Micralymma marinum (fig. 32), inhabiting the tidal zone, on rocky places with
a rich supply of Fucus and other seaweeds. Its distribution in the North Atlantic
region has already been associated with the Gulfstream (Lindroth, 1931, p. 497-
499) and it is difficult to understand it otherwise than as the result of direct
transport.

Two other insects, confined to the seaweed drifts of the shore, the Hydrophilid
beetle Cercyon litoralis Gyll. and the Coelopid fly Orygma luctuosum Meig. (Nielsen,
Ringdahl & Tuxen, 1954, p. 84), have a similar Amphiatlantic distribution but
probably do not endure submersion during any stage of development. The most
probable explanation seems to be that they were introduced from Europe into
North America.

It is a well-known fact that pieces of seaweed, seeds of terrestrial plants, &c.,
are regularly carried with the Gulfstream from North America to the European
west-coast and the exclusive seashore plant Mertensia maritima L., unknown
in Eurasia E. of the White Sea, is generally regarded as a Gulfstream immigrant,
even by Hulten (1937, p. 64) who is otherwise extremely sceptical of all kinds
of transatlantic floral exchange.

The quite isolated area of Ranunculus cymbalaria Pursh on the coast of SW
Scandinavia, on both sides of the Swedish-Norwegian border limit, was likewise
originally (Nordhagen, 1916; still by Sterner, 1945, p. 40) interpreted as the result
of transport with the Gulfstream. Hulten (1937, p. loi) regarded it as an old
relict-occurrence, a strained idea, but later (1950, p. 81) changed his mind,
apparently in accordance with Tambs-Lyche (1937), who thought of introduction
with ballast from North America. This explanation seems to be the most accept-
able one.

The famous flightless Great Auk {Penguinus or Alca impennis L.), now extinct,
once had a pronouncedly Amphiatlantic distribution (fig. 33). The limited means
of dispersal of this bird refer it to the present group.

The relationship between the palaearctic and nearctic faunas

FIG. 32. Distribution of the Staphylinid beetle Micralymma
marinum Strom {stimpsoni Lee), inhabiting the tidal zone, in
relation to the Gulfstream. The figured specimen is from
Newfoundland.

(From LiNDROTH, 1931, revised.)

(Photo P. Ardo.)

4. A fourth group would consist of warm-blooded animals, active in winter
and therefore possibly able to traverse the sea between arctic islands and continents
on the ice. This group is well represented among Circumpolar (p. 291), but not
among Amphiatlantic animals, except perhaps once, temporarily, by the Muskox
(Ovibos moschatus L.) (vide map by Ekman, 1922, p. 396).

5. Freshwater species with stage of development adapted to passive transport
with birds. The Sponge Heteromeyenia ryderi. Like others of the fresh-water
Spongillids, this species forms "gemmulae", less than i mm. in diameter, which

246

CHAPTER III

^^^^ii

^\^

;:-

^Hr

t\ "^

^^^

w>

\^

)>^

^\ \ ^^ — "{

mli^

i^syjy^''^ \

i

%-.. I

^^/i

'T

"x y^A

^

%^

^

y5i

\

VUL '■• juO^-

^

'r-^/

_^_^

\ ' ( --■■' \

'-^^

FIG. 33. The extinct Great Auk, Penguinus (Alca) im-
pennis L. Late- and postglacial distribution. — Solid
dots = on historical evidence. Open dots = subfossil
remains. The figured bird belongs to the Zool. Inst.,
Lund.

(According to Blasius, 1903, Ekman, 1922, &c.)

(Photo Hanna Grans.)

are extremely resistant and constitute the normal stage of dispersal of the organism.
The gemmulae may be carried by water but also, after drying up, by the wind; for
greater distances, transport with birds is usually accepted as the normal means of
dispersal {vide, for instance, Thienemann, 1950, p. 156).

It was therefore suggested by some students that bird-transport from North
America might have been responsible for the European occurrence of Heteromeyenia

The relationship between the palaearctic and nearctic faunas

247

FIG. 34. The situation of air pressure and winds over the North Atlantic at midday
during three days, September 30th to October 2nd, 1953, with considerable influx
of North American birds into the British Islands,

(From Williamson, 1954.)

ryderi. Others, for instance Scharff (1907, p. 34) and Stephens (1920, p. 247-248),
regard this sponge as belonging to a very old faunal element, evidence of a former
transatlantic land-connection. Arndt (1928, p. 159-163) tried to select a middle
course: he stressed the probability of bird-transport from Ireland to the Faeroes
but admits that the Irish population may be old. His view has been misinterpreted
by Sparck (1934) who himself ascribed even to the population of the Faeroes a
persistence in situ since the last interglacial period.

It seems advisable to consider the history of the Heteromeyenia in connection
with the related American element of the British flora. Phytogeographers have
almost unanimously accepted this as an old element too, a "relict" from a period
of transatlantic land-connection. Quite recently, however, Heslop-Harrison (1953)
has given voice to an opposing view: he is inclined to interprete the occurrence of
these American plants in the British Isles as due to transport with migrating birds
and especially draws attention to the routes of the Greenland White-fronted
Goose (Anser albifrons flavirostris Dalg. & Sc.) which winters partly in Ireland.
The weak point of Heslop-Harrison's argument is that, according to him, only
one of the species in question, Sisyrinchium angustifolium, grows in Greenland
(Bocher, 1948) an this species, it should be added, is the one least likely to be
dispersed by birds. ^ He therefore feels compelled to assume that other birds have
brought the plant diaspores from NE North America to Greenland, where they
"changed birds", without being able, for climatic reasons, to germinate on the spot.
This is a most far-fetched hypothesis.

If we are at all willing to accept bird-transport as a possible agency in the present
case, it seems necessary to evaluate the chance of birds arriving in the British

^ Dr. T. W. Bocher and Dr. J. Iversen have informed me, however, that the Greenlandic
and the Irish Sisyrinchium are different. Dr. Iversen has grown both forms together. (Cf.
p. 250).

The relationship between the palaearctic and nearetic faunas

249

FIG. 35. World distribution
of Ponera coarctata Latr.,
no doubt an old species,
belonging to a very primi-
tive group of ants.

1 = forma typica

2 = sbsp. pennsylvanica

Buckl.

3 — sbsp. boerorum Forel

4 = sbsp. mackayensis Forel

The figured worker (from
Smith, 1947) is of sbsp.
pennsylvanica.

Isles directly from parts of North America inhabited by the plants considered,
that is, not north of Newfoundland. No bird has a normal route of migration like
this, it is a question of abnormal movements or pure wind-drift.

Now, Williamson (1954) has given a very interesting report of occasional
appearances of American birds in the British Isles in the fall of 1953 and, what is
more important, has been able to correlate the observations with the meteorological
situations preceding the influx of these birds (fig. 34). Their route appeared as a
clear wind-drift and the offshore drift on the American side, according to the
weather, at least in one instance could have taken place as far south as Cape Hat-
teras. The American birds noted in Great Britain and Ireland, on this and other oc-
casions in late years, include geese, ducks, waders, passerines, cuckoos &c. (Alex-
ander & Fitter, 1955) and Williamson (I.e., p. 26) speaks of "the now regular

annual occurrence of nearetic waders at English reservoirs and sewage-farms". Is
the sudden appearance of the American freshwater snail Lymnaea catoscopium Say,
in "a warm engine-pond in a timber yard", in Leith, Scotland (Kevan, 1943),
possibly to be explained in this way, and not through human transport?

The direct transatlantic bird-connection, direction east, is accidental but not
too scarce. How great is the chance of these birds coming into contact with dia-
spores of the discussed plants? Heslop-Harrison (I.e., p. 114) stresses the point
"that they are all aquatic, marsh or lake-margin plants". Though this statement
does not apply very well to the Sisyrinchium, which also takes a different position
in other respects, Heslop-Harrison is probably right in regarding this ecological
property of the group (equally relevant for Heteromeyen a ryderi) as an indication
in favour of bird-transport. The reasons seem to be: —

250

CHAPTER III

(a) Nowhere is there such a congregation of migrating birds, particularly of
swimmers and waders, as on the shores of lakes and of the sea.

(b) Small objects of any kind attach themselves most easily to the feet and feather
of a bird in water or on very wet places, especially together with mud. After the
bird has taken wing, the mud dries up quickly and remains well fixed.

(c) Ubiquistic freshwater organisms, by means of their usually much wider
geographical distribution in comparison with terrestrial ones, and at the same time
their confinement to a more disjunct mosaic of suitable habitats, often of a tem-
porary character, apparently are much better adapted to passive dispersal than
terrestrial organisms are. Among the methods exploited, transport with birds
should not be underestimated {vide the extract by Thienemann, 1950, p. 156-
158).

(d) A special case is provided by Potamogeton epihydrus. The fruits of Pota-
mogeton are a normal part of the food of swimming birds, especially of ducks,
and Lohammar (1954) has shown that the passage through a bird's intestines
highly favours the germination.

To my mind, the theory of bird-transport should be earnestly considered in the
case of four of the "American" plants: Eriocaulon septangulare, Limosella subulata,
Myriophyllum alterniflorum americanum, and Potamogeton epihydrus; in addition,
for the single animal of this geographical group, Heteromeyenia ryderi.

It may sound absurd that causal appearances of wind-driven birds should have
proved more efficient as an agency enlarging the area of transported organisms
than birds on their normal route of migration. This is, however, a seeming dis-
crepancy only. Dispersal of seeds, fresh- water animals, &c., along the great
highways of migrating birds has gone on regularly for thousands of years and re-
sulted in continuous species' areas, regarded by us as "natural". Yet the erratic
east-west dispersal is predestined to give a better effect proportionally, because it
usually means the removal into a place with a climate similar to that of the starting-
point.

The two remaining plants must be treated separately. They are less hygrophilous,
neither of them is a w-ater or even a shore plant, and their diaspores are different.
Spiranthes romatizoffiana is an Orchid with very numerous and very minute seeds
which are wind-spread and must be assumed to constitute a regular component
of aerial plankton. The resistance of the seeds to cold and exsiccation should be
investigated experimentally. Probably the distribution of the species is more
restricted by the limited possibility of the seedling finding the right fungus for
mycorrhiza than by powers of dispersal.

It is difficult to understand that the globular seeds of Sisyrinchium could be suited
for passive transport of any kind or that any vegetative part of the plant could

The relationship between the palaearctic and nearctic faunas 251

form a substitute. A further important peculiarity of S. angustifolium is its occur-
rence on the European mainland, most often regarded as escape from cultivation,
with one exception: in the Carpathians. According to repeated judgment (Lauter-
born, 1927, p. 84; Bocher, 1948, p. 15) the plant here gives the impression of
being indigenous. Serious consideration must therefore be given to the possibility
that the small European areas of Sisyrinchium angustifolium are actually pre- or
interglacial relicts, the remnants of a circumpolar area. At least, this hypothesis
seems more plausible for the Sisyrinchium than for Spiranthes romanzoffiana, as
emphasized by Hulten (1937, p. 132).

There seems to exist on the American side a small European element of boreal
(non-arctic) plants corresponding to the "Americans" in Europe just treated. At
least Fernald (1925, p. 272 a.f.) regards a comprehensive group of such plants,
particularly in Newfoundland, as old relicts. The majority of these, however, are
no doubt late introductions (vide above, p. 146). Heslop-Harrison (1953, p. m)
maps two species, Carex hostiana D.C. (fulva Host.) and Potamogeton polygonifolius
Pourr. (oblongus Viv.), which he seems to regard as native in North America. Both
are restricted to Newfoundland (incl. St. Pierre-Miquelon) and either Anticosti
(C. host.) or Sable Island, Nova Scotia (P. polyg.). By Clapham, Tutin & Warburg
(1952, p. 1370) Carex hostiana is indicated as "probably introduced" in North
America. It may be appropriate to apply to the Potamogeton the suggestion of bird-
transport put forth concerning P. epihydrus above. A glance through the American
"Check-List" (1931) shows convincingly that stray birds on migration from Europe
are not uncommon on the Atlantic coast of North America.^

On the American side also, like Heteromeyenia ryderi in Europe, there seems to
exist at least one equivalent among animals, the fresh-water snail Lymnaea (Radix)
peregra Miill., known from only two localities on the Avalon Peninsula of SE
Newfoundland (Brooks & Brooks, 1940, p. 62). It is worth mentioning that the
appearance of the same species ("ovata Drap.") on small, new-formed islands
off the German Northsea coast was most likely due to bird-transport (Thienemann,
1950, p. 157). Also in other parts of Europe the distribution of this species gives
evidence for an extremely high power of passive dispersal and this property of
ubiquistic and more or less cosmopolitan fresh-water Molluscs is generally ex-
plained as the effect of transport with birds (Boycott, 1936, p. 123-126; Hubendick,
1947, p. 508-512, "limosa L.").

Whether or not the isolated occurrence of the American Amphipod Gammarus

^ The European birds most regularly observed on the Atlantic coast of North America are:
Branta leticopsis Bechst., Crex crex L., Lams minuttis Pall., Anas crecca L. /. typ., Erolia
alpina L. /. typ., Philomachiis pitgnax L., Scolopax rusticola L. /. typ., Vanellus vanellus L.
All are Swimmers or Waders.

252

CHAPTER III

fasciatus Say {tigrinus Sext.; vide Hynes, 1954) may be due to bird-transport,
I am unable to judge. Segerstrale (1954, p. 68 a.f.) accepts this as the normal mode
of dispersal for G. lacustrts G. O. Sars.

5. Animals without special properties of active or passive dispersal. The ant
Ponera coarctata Latr. (fig. 35). The sexuals, males as well as females, are winged in
this species but do not make ordinary nuptial flights, the mating as a rule taking
place on the ground near the nest, or even within the nest. At any rate the fertilized
queens never fly (Wheeler, 1900; Michener & Michener, 195 1, p. 141-142). The
dispersal of this ant is therefore functionally the same as for a constantly wingless
species.

Ponera coarctata inhabits a curiously broken-up area. Besides the forma typica
of Europe and the Mediterranean region and the sbsp. pennsylvanica Buckl. of
eastern North America, there is one subspecies endemic to South Africa (Natal)
and one to Australia (Queensland) (Creighton, 1950, p. 47). The subfamily Po-
nerinae includes the most primitive of living ants (Wheeler, 1900) and must be
very old, geologically speaking, not as a group alone, but also as actual species.
This is confirmed by the fact that the extinct Ponera atavia Mayr, known by all
casts from the Baltic amber (about the age of more than 40 million years), comes
very close to coarctata (Wheeler, 191 5, p. 39-40).

The present distribution of Ponera coarctata must be interpreted as the remnant
of an almost world-wide area in Tertiary time and the occurrences on both sides of
the Atlantic do not constitute evidence in favour of the theory of a direct land-con-
nection (cf. the distribution of the beetle family Cupedidae, fig. 60).

The conclusions arising from the study of animals showing an Amphiatlantic,
or similar, distribution are: —

Their present area seems understandable without accepting a late (Pleistocene)
land-connection between the two continents. The existence of such (in one of the
interglacial periods) would undoubtedly have resulted in a much richer faunal
exchange in both directions, and a less "asymmetrical" range of distribution in
the individual, actually existing cases.

On the other hand, it is not correct to regard almost all Amphiatlantic species
as peripheral relicts of an earlier circumpolar distribution, as do Hulten (1937) and
Deevey (1949), nor to emphasize that "der Terminus 'amphiatlantische Arten'
in faunagenetischer Hinsicht gegenstandslos wird" (Reinig, 1937, p. 22). Dispersal
across the sea is no negligeable part of faunal and floral history.

The relationship between the palaearctic and nearctic faunas 253

The theories of earlier transatlantic land-connections

The Iceland — Greenland bridge

It may seem inconsistent, after the existence of a biologically efficient land-
connection between Europe and North America has been denied or at least
considered highly improbable, to raise again the problem of a "land-bridge"
within the North Atlantic region. Land-bridges, however, are not bound to join
continents, they may be, or have been, "blinds" connecting islands to a main-
land.

Perhaps the situation is best made clear to the reader in the form of a brief
account, brought up to date, of the zoogeographical conclusions to which the
present writer arrived (193 1) from a study of the insect fauna of Iceland, all the
more as later students have agreed, at least in the essentials.

The fauna of Iceland is practically purely European, the American element being
restricted to some few species: the Noctuid Moths Crino sommeri Lef., Crymodes
exulis Lef., Rhyacia (Caradrina) quadrangula Zett. (figs. 36-38), and the Gnat
("Black Fly") Simulium vittatum Zett. (fig. 39), all of them no doubt able to arrive
by air under present conditions. Possibly the Empidid Fly Rhamphomyia hirtula
Zett. (Greenl., Icel., Scotl.) belongs to the same group. The Icelandic form of the
Water-beetle Colymhetes dolabratus Payk., thomsoni Sharp, is decidedly more re-
lated to the sbsp. groenlandicus Aube, of Greenland and North America (Brinck,
1940, p. 37-40) and must likewise have invaded Iceland from a western direction,
by air or with drifting ice. The same is true for a genuine fresh-water animal, the
small Copepod Diaptomiis minutus Lilljeb., a Nearctic species, known also from
Greenland (Poulsen, 1939, p. 29).

Even among breeding birds, only three Icelandic species are of clearly Nearctic
origin: Biicephala islandica Gm., Gavia (Colymbus) immer Briinn., and Histrionicus
histrionicus L. (Timmermann, 1938-49, p. 123).

As far as vascular plants are concerned, Epilobiutn {Chamaenerium) latifolium L.
and two Orchids, Habenaria (Leucorchis) hyperborea L. and H. (L.) straminea
Fern., belong to an American element (the last-named also reaching the Faeroes).
All of them have seed excellently suited for wind-transport. Lowe (1950) has added
some forms of doubtful taxonomic position, including polyploids, the historical
interpretation of which is most delicate since a polyphyletic origin cannot be dis-
missed.

In all, this is a poor share of American inhabitants indeed for an island situated
900 km. or more from the British Islands and Scandinavia, but only 330 km. from
Greenland which, in its turn, is separated from southern Baffin Island by the

254

CHAPTER III

ii4

^^

w^

i^fp

■^•^J^?^!^

\ \V\^

3%:^L_

\^''\ \ y^

a^

\J\^\-

FIG. 36-38. The North Atlantic distri-
bution of three Noctuid Moths.
36. Rhyacia (Caradrina) qiiadrangula
Zett. — 37. Crino sommeri Laf. (In
E North America only an old record
from Labrador.) — 38. Crymodes exults
Laf.

All species are distributed west of
the limit of the map.

The figured specimens are from
Greenland.

36

37

38

v.y

(Photo P. Ardo.)

The relationship between the palaearctic and nearctic faunas

255

^

(Photo P. Ardo.)

FIG. 39. Approximate distribution of Simulium vittatuni Zett., a species of gnat or
"black fly", a regular nuisance to man and cattle.

Mainly according to A. Stone (in litt.) who thinks the species may be a complex one.
The figured specimen is from Greenland.

only slightly wider Davis Strait (350 km.). The Nearctic element of the Icelandic
insect fauna is less than one per cent!

On the Faeroes, as a matter of course, it is still more insignificant, consisting
only of two of the Noctuid Moths just mentioned, Crino sommeri and Crymodes
exulis, and, among animals other than insects, of the freshwater Sponge Hetero-
meyenia ryderi Potts, treated above (p. 245).

The fauna of Greenland, however, provides the great surprise within the chain
of islands across the North Atlantic. This large island, almost a continent of
more than 2 million square kilometers, though to more than four fifths covered
with ice, traditionally belongs to America. Actually, the shortest distance to
the North American mainland, in northern Labrador, is little more than half
(750 km.) of the distance to the European continent, in northern Norway
(1,400 km.).

The American influence on the Greenlandic fauna, as on its flora, is quite
obvious but it is far from dominating. Above all it is striking how unequally large
is this element among diff'erent groups of animals. In those sufficiently known and
large enough to allow a comparative analysis, the share of the different geographical
groups is as follows: —

256 CHAPTER III

Representative groups of the terrestrial Greenlandic fauna

(species and subspecies; obviously introduced forms omitted)

Total Holarctic Palaearctic Nearctic Endemic

Birds (Salomonsen &

Gitz, 1950) 68 30 = 44 % 9 = 13% 18 = 27 % 11 = 16 %

Macro- Lepidoptera

(Henriksen, 1939) 26 14 = 53% 3 = 12% 9=35%^ —

Coleoptera (ditto;

Vibe, m litt.) 24 7 = 29 % 11= 4^ % 6 = 25 %* —

Collembola (Hammer,

1953) 46 27 = 59 % 13 = 28 %3 5 = 11% 1=2%

Araneae (Braendegaard,

1946, & Holm, in litt.) 50 2 1 = 42 % 1 1 = 22 %^ 1 1 = 22 %* 7 = H %

Together 214 99 = 46 % 47 = 22 % 49 = 23 % 1 9 = 9 %

If the 16 endemic subspecies of birds are distributed among the three other geo-
graphical groups, according to their relation to other subspecies (Salomonsen &
Gitz, 1950), the figures are:

37 Holarctic, 12 Palaearctic, 19 Nearctic forms
= 54% =18% =28%

If, furthermore, the 7 Spiders and the single Collembol, unknown outside Green-
land, are counted as Nearctic, which they probably are, the 214 species and sub-
species of the list include:

106 Holarctic, 50 Palaearctic, 58 Nearctic forms

= 50 % = 23 % = 27 %

This means that, provided the selected groups are accepted as representative,
the terrestrial fauna of Greenland consists of one half Holarctic, and one fourth each
of Nearctic and Palaearctic forms (species and subspecies).

Before considering the fascinating question of the reasons for the peculiar zoo-
geographical character of Greenland, and the North Atlantic islands on the whole,

^ Included are Byrdia (Dasychira) groenlandica Wocke, found in the Canadian Arctic
according to Dr. Munroe {in litt.), and Rhyacia (Euxoa) drezvseni Stdgr., the occurrence of
which in North America proper is not quite settled (McDunnough, 1950, p. 396); further-
more, Spaelotis (Amphitrota) clandestina Harr. {unicolor Wlk.), recorded also from E Siberia
but represented in N Europe by sbsp. siiecica Aur.

* \nc\ud\i\f^Micralymmabrevilingiie Schio. and Coccinella transversoguttata Fald., occur-
ring in Siberia but not in Europe.

^ Including two species (Sminthurus concolor Mein. and Collinsia thulensis Jacks.) outside
Greenland known from Spitzbergen only.

* Including Islandiana princeps Braend., outside Greenland known from Iceland only.

The relationship between the palaearctic and nearctic faunas 257

it seems appropriate to make a closer inspection of their beetle faunas (tab. 7,
diagr. 7). We are now in a far more favourable position than in 1931, since at least
a preliminary survey has been made of the Coleoptera of Baffin Island (Brown,
1937). This insect order, even from other points of view, is well suited as starting-
point. It is tolerably numerous in species, even in northern regions, and, above all,
it contains many flightless, not easily dispersed forms which may, in their present
distribution, reflect faunal history with unusual clarity.

The detailed facts concerning the coleopterous fauna included in table 7 and
diagram 7, give very clear evidence as to the zoogeographical character of Green-
land. It appears that the purely Palaearctic element is not only quantitatively
dominant (11 species), but that it furthermore contains the following flightless
species with consequently reduced powers of dispersal:

Nebria gyllenhali f. typ. Otiorrhynchus arcticus (fig. 41)

Lathrobiiim fidvipenne O. dubius

Quedius boops Phytonomus elongattis.

TABLE 7. Beetlca (Coleoptera) of the Faeroes, Iceland, Greenland, and Baffin Is-
land, regarded as indigenous, with a summing-up of their distribution in other parts

of the Holarctic area.

( +) = introduced. — a, b, and c = different subspecies; ab or be — both subspecies,
or intermediates.

In the first column, m = macropterous (wings full), b = brachypterous (wings
reduced), (b) = wings full, or almost so, but probably non-functionary (incl. di-
morphic forms occurring in the brachypterous form only), d == dimorphic (both forms
represented, also within the area' considered).

The limit between E and W North America is supposed to run through Hudson
Bay and Mississippi. Ditto between W and E Siberia is River Yenisei.

All species stated or believed to be introduced into the four island areas considered
have been excluded. A division of this kind is in part a matter of personal judgment,
yet the rather rigorous discharge here practised at any rate has not over-dimensioned
the European element. Students interested in the species thus omitted are referred,
for the Faeroes to West (i939)\ for Iceland to Lindroth (1931)^, for Greenland to
Henriksen (1939)', for Baffin Island to Brown (1937).

^ For the Faeroes, the only additional species is Otiorrhynchus rugosostriatiis Gze. (S. G.
Larsson, in lift.), probably introduced from the British Islands.

^ Many imported beetles, but apparently only one native species, Gymnusa brevicollis
Payk., have been collected since 1931 (S. G. Larsson, in litt.). The North American P/zywa-
phora pulchella Newm. (Lindroth, 1931, p. 219) was no doubt reported by a pure mistake.

^ For the recent discovery in Greenland of Caenoscelis ? cryptophaga Reitt. and Doryio-
mus sp. we are indebted to Mr. C. Vibe who kindly made the specimens available to me.

I 7 ~ 565597 Lindroth

258

CHAPTER III

Carabidae

Amara alpina F. (fig. 40) .
A. aulica Panz

A. quenseli Schnh

Bembidion bipiinctatum L. .

B. grapei Gyll

B. redtenbacheri K. Dan. . ,
Calathus fuscipes Gze

C. melanocephalus L

Carabus problematicus Hbst. .
Cymindis unicolor Kby. . . .
Dichirotrichus pubescens Payk. .
Harpalus quadripiinctatus Dej

Loricera pilicornis F

Nebria gyllenhali Schnh. . .

N. nivalis Payk

A^. salina Fairm

Notiophilus aquaticus L. . . .

A^. biguttatus F

Olisthopus rotundatus Payk.
Patrobus assimilis Chd. . . .

P. atrornfus Strom

P. septentrionis Dej.^ . . . .
Pterostichus adstrictus Eschz. .

P. arcticola Chd

P. diligens Sturm

P. haematopus Dej

P. nigrita F

Trechus fulvus Dej

T. obtusus Er

Trichocellus cognatus Gyll. . .

J

1

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3

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7

8

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(b)

ab

a

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a

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—

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b

b

b

m

—

+

+

+

+

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( + )

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+

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+

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+

+

m

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+

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+

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+

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—

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d

+

+

+

+

+

+
+

+

+

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+

b

+

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(+)

(b)

+

+

+

+

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—

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(b)

ab

a

a

a

a

a

a

—

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(b)

ab

a

a

a

—

—

—

b

b

b

(b)

—

—

+

+

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(b)

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+

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+

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+

+

+

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—

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+

+

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+

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+

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+

+

+

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m

+

+

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4-

+

Taxonomical remarks, 1-8.

I. The North American Patrobus septentrionis Dej. is not consistently so different from
the Eurasian nominal form as previously assumed (Lindroth, 1955a, p. 84). Only the New-
foundland population seems to deserve a subspecific name. Ample series from Labrador,
Greenland, Iceland, and Scandinavia, have convinced me that the punctuation of head and

The relationship between the palaearctic and nearctic faunas

Haliplidae

Halipliis fulvus F. (excl. lap-
ponum Thorns.) ....

Dytiscidae

Agabus congener Payk. (coll.) .

A. moestus Curt

A. solieri Aube

Colymbetes dolabratus Payk. .
Deronectes miiltilineatus

Falkstr

Hydroporus erythrocephalus L.

H. lapponum Gyll

H. melanocephalus Mrsh. . .

H. memnonius Nic

H. nigrita F

H. palustris L

H. polaris Fall

H. pubescens Gyll

Gyrinidae

Gyrinus opacus C. R. Sahib.

Hydrophil tdae

Anacaena globulus Payk. . .
Cercyon litoralis Gyll. . . .
Helophorus aquaticus L. . .
H. arcticus Brown . . . .

H. brevipalpis Bed

H. flavipes F

Megasternum boletophagum
Mrsh

Liodidae

Hydnobius punctatiis Sturm

Scy dtna enidae

Stenichmis collaris M. & K.

1234567

9 10 II

m ! +

+
+

+

+

+ +

+

+ 1

+ !

— + + + +

+ +

+ i +

+ 1 +

— b

- (+)

- (+)

+ —

259

be

( + )

prothorax, more pronounced in the North American material, is a variable character even
within geographically limited populations.

2. For the taxonomy of Atheta hyperborea Brd. and islandica Kr., vide Brundin (1943,

P- 15-19)-

3. Boreaphilus nearcticus Blair. Two ? ? from Baffin Island (Mus. Ottawa) were cornpared
with the original specimens oi nordenskioldi Maklin (1878, p. 25; 1881, p. 42) from Siberia,

26o

CHAPTER III

Staphylinidae

I

2

3

4

5

6

7

8

9 10

n

1

Acidota crenata F

m

+

+

+

+

—

+

—

— +

+

Aleochara obscurella Gr. . . .

m

—

—

+

+

+

—

—

— —

—

Amischa analis Gr

m

—

+

+

+

+

+

—

- ( + )

( + )

Arpedium brochypterum Gr. .

(b)

+

+

+

+

+

—

—

— i ?

—

Atheta atramentaria Gyll. . .

m

+

+

+

+

+

+

—

— —

—

A. excellens Kr

m

+

+

+

+

+

+

—

— —

—

A. fungi Gr

m

+

+

+

+

+

+

+

— +

+

A. graminicola Gr

m

+

+

+

+

+

+

—

— +

+

A. hyperborea Brd.^

m

—

—

+

—

—

—

+

—

—

—

A. islandica Kr.^

m

—

?

+

p

+

+

+

—

—

—

A. melanocera Thorns. . . .

m

+

+

+

+

+

+

—

—

—

—

A. vestita Gr

m

—

—

+

+

+

+

—

—

—

—

Autalia puncticollis Sharp . .

m

—

—

+

+

+

—

—

— —

( + )

Boreaphilus nearcticus Blair'' .

b

—

—

—

—

—

—

+ . +

—

Bryoporiis sp.*

m

—

—

—

—

—

—

—

+

+

—

Cafiiis xantholoma Gr

m

—

—

+

+

—

+

—

—

—

—

Geodrotnicus globulicollis Zett. .

m

—

—

+

+

—

+

—

—

—

—

Gnypeta cavicollis J. Sahib. .

m

—

+

—

—

—

—

+

—

—

—

Gymmisa brevicollis Payk. . .

m

—

+

+

+

+

—

—

I +

+

Lathrobitim fidvipenne Gr. . .

b

+

+

+

+

+

+

+

—

—

—

Lesteva longelytrata Gze. . .

m

—

—

+

+

+

+

—

—

—

—

Micralymma brevdhigue Schio.^

b

1 +

+

—

—

—

—

+

+

—

+

M. marinum Strom (fiig. 32) .

b

—

—

+

+

+

+

+

—

+

—

Ocalea picata Steph

m

■ —

—

+

+

+

+

—

—

—

—

Omcdhim laeviusculum Gyll. .

m

—

—

+

+

+

+

—

—

—

—

0. ripariiim Thorns

m

—

—

+

+

+

+

—

—

—

—

Othius melanocephalus Gr. . .

b

—

—

+

+

+

+

—

—

—

—

0. viyrmecophilus Kies. . . .

b

—

—

+

+

+

—

—

—

—

—

0. punctulatus Gze

m

—

+

+

+

+

—

—

—

—

—

Oxypoda islandica Kr. . . .

m

—

+

+

+

—

+

—

—

—

—

0. soror Thorns

b

—

—

+

+

—

+

—

—

—

—

Philonthtis trossuhis Nordm. .

m

—

—

+

+

—

+

—

—

—

—

Quedius boops Gr

(b)

i —

+

+

+

+

+

+

—

—

—

Mesenkin and Sopotshnoj Island, and with a series from Tolstoinos (J. Sahlberg, 1880,
p. 106); all localities on the eastern side of the Yenisei estuary. Though I was unable to
study the S genitalia, it seems clear from external characters that the species, though closely
related, as already supposed by Blair (1933, p. 95), are not identical. The vertex is exca-
vated behind the eyes in nearcticus and this concavity has a strong, striated sculpture. In
nordenskioldi, the vertex is truncate towards the contracted neck.

4. The "Mycetoporus sp." of Brown (1937, p. 1 10), according to the single specimen (Mus.
Ottawa), is a Bryoporus of a species not occurring in Europe but related to rugipennis Pand.
It is possibly rufescens Lee. and I have ventured to mark the distribution as "E. North
America".

5. Micralymma brevilingue Schio. A comparison between ample series of this species,

The relationship between the palaearctic and nearctic faunas

261

Q. fuliginosus Gr

Q. fulvicoUis Steph

Q. nitipennis Steph

Q. umbrinus Er

Sipalia circellaris Gr

Stenus brunnipes Steph. . . .

S. canaliciilatus Gyll

S. carbonarius Gyll

S. clavicornis Scop

S. coarcticollis Epp

S. impressus Germ

S. nanus Steph. (coll.). . . .

S. nitidiusculus Steph

Tachinus corticinus Gr. . . .

T. riifipes DeG

Tachyporus atriceps Steph.

Pselaphidae

Bythinus puncticoUis Denny
{validus Aube)

Cantharidae

Malthodes mysticus Kies. . .

M. pumilus Breb

{atomus Thorns.)

Elateridae

Hypnoidus riparius F

Helodidae

Helodes mimita L

Byrrhidae

Byrrhus fasciatus Forst. . . .

Cytilus sericeus Forst

Simplocaria tessellata Lee*. .
5*. semistriata F

Crytophagidae

Caenoscelis ? cryptophaga Reitt.'

(b)
(b)
b
b
b
b
m
b
m
b
m
m
b
b

( + )

+

+

+

( + )

( + )

+

( + )

including "types" from Greenland (Mus. Copenh.) and Brown's specimens from Baffin
Island (Mus. Ottawa), and of dicksoni Makl. from arctic Siberia, 1 1 localities, between the
mouth of River Yenisei and St. Lawrence Bay in the Bering Strait (Swed. Riksmus.), has
convinced me that they all belong to one species. The differences from brevilingue mentioned

262

CHAPTER III

< Coccinellidae

Coccinella transversoguttata

Fald

C. ii-piwctata L

Scymnus limonii Donis. . . .

Scarabaeidae

Aphodius lappotmm Gyll. . .

Chrysotnelidae

Chrysomela staphylea L. . .
Phyllodecta polaris Sp.-Schn. .

Curculionidae

Apion cruentatum Walt. . . .
Barynotus squamosus Germ. .
Dorytomus taeniatiis F. . . .
Lepyrus labradorensis Blair

Notaris aethiops F

Otiorrhynchus arcticus O. Fbr.

(fig- 41)

O. atroapterus DeG

O. dubius Strom

O. rugifrons Gyll

Phytonomus elongatus Payk.

Ph. pedestris Payk

Sciaphilus asperatus Bonsd.
Strophosomus melanogrammns

Forst

Tropiphorus obtusus Bonsd. . .

b —

b —

m |- +

b —

m +

+ +

+ :, +

+ 1' +

- ( + )

- (a)

(+)

( + )

(+)

by Maklin (1881, p. 42) for dicksoni: the broader 5th and the narrower 9-ioth antennal
segments, shorter elytra, and more shining abdomen, are by no means constant. The S
genitalia seems to be almost identical, though the penis (median lobe) is slightly shorter, more
parallelsided and with blunter apex in Greenland specimens. The species (dicksoni) has also
been reported from the Pribilof Islands off Alaska (vide Leng, Suppl. I, 1927) and thus
apparently has a high-arctic circumpolar distribution. Mr. W. O. Steel, Maidenhead,
Berks., who is revising the genus, informs me that probably two or three subspecies are
involved, the population of the Pribilof Islands likewise deviating in some minor points.
6. Simplocaria. The Greenland species is apparently the North American tessellata Lee,
according to a series from Churchill, Man. (det W. J. Brown, Mus. Ottawa), and not identical
with the European metallica Sturm. The species are very closely related but in tessellata the
pilosity of elytra is a little denser, forming more contrasting dark and light spots. The
penis (median lobe) is slightly slenderer and a trifle more enlarged towards apex, the denticles
along the sides in front of the flattened apex are more pronounced; but these differences are
so slight that it is easily understood why Szekessy (1936, p. loi) could declare the genitalia
of Greenlandic and European c?o as identical. — Apparently two species (other than remota

The relationship between the palaearctic and nearctic faunas 263

Brown) were confused under "tessellata" in North America. At least specimens from New-
foundland belong to a third species.

7. Caenoscelis. Through the courtesy of Mr. Vibe I have been able to study 3 specimens
collected by him in Greenland. They agree with the descriptions of cryptophaga Reitt.
(187s, p. 87; 191 1, p. 66). A comparison with a typical specimen (from "North America")
would however be desirable. In Casey's key (1900, p. 104-107) they run to macilenta Csy.
(New York, Penns.) which is probably a synonym. At least I have ventured to give the distribu-
tion as "E. North America".

8. After the above was set up for print the Greenland Dorytomus has proved to be
distinct from taeniatus and probably a Nearctic species.

The Nearctic element (including two Holarctic species lacking in Europe)
amounts to 6 species only:^

Colymbetes dolabratus groenlandiciis Caenoscelis ? cryptophaga

Gyriniis opacus blairi Simplocaria tessellata

Micralymma brevilingue Coccinella transversoguttata

Of these only the Micralymma is flightless. It inhabits the tidal zone and is no
doubt easily dispersed by sea-water drift (cf. M. marinum, p. 244).

This point of view provides a satisfactory explanation for the unequal share
of Nearctic species in Greenland within the diff"erent groups of animals to which
attention was drawn above. They are most numerous in Macro-Lepidoptera (35
per cent) and Birds (28 per cent), with more or less developed powers of active
flight, in the second place among Spiders (22 per cent, or 36, if "endemic" species
are included), with their common habit of "ballooning" into the air. But in
Collembola (Spring-tails) the Nearctic species amount to 11 per cent only, con-
siderably less than in Coleoptera. Among comparatively able flyers, as the Tricho-
ptera (Caddies-flies), 6 of 9 (or 10) species are Nearctic (Henriksen, 1939, p. 23-24;
Vibe, 1950) but the Neuroptera (Lace- wings) , with poorly developed powers of
flight, are represented by one Holartic and two Palaearctic species.

The composition of the coleopterous fauna of Baffin Island is in complete
accordance herewith. Of the 16 species^ known so far from this island (Brown,
1937), only three species are common also to Greenland: the Water-beetles
Colymbetes dolabratus groenlandicus and Hydroporus melanocephalus, both good
flyers, and the Staphylinid Micralymma brevilingue, just mentioned as easily

^ After the above was written, the Dorytomus from Greenland has proved to be a prob-
ably Nearctic species, different from taeniatus F. {vide table 7). It is fully winged.

^ From Ellesmere Island in the high north (between about 76° and 83° north) only 5
species of Coleoptera are reported (Brown, 1937, and in litt.). Two, of the genera Crypto-
phagus and Enicmus, are accidental importations and the Quedius "fulgidus", according to
earlier practice of nomenclature within the genus, is no doubt mesomelinus Mrsh., a true
synanthrop, common in the dwellings of the West Greenland esquimos. The two remaining
species are Micralymma brevilingue Schio. and one Atheta, which has been investigated
by Dr. Brundin, who informs me that it represents a species unknown to him, possibly
undescribed (unfortunately a female), and thus almost certainly a Nearctic element.

264 CHAPTER III

dispersed by sea-drift. The remaining 13 species, 7 of them lacking the power of
flight {vide table 7), have been unable to get across to Greenland.

The. character of the interjacent sea as a faunistic obstacle is best illustrated by
maps of two functionally brachypterous species, the Ground-beetle Amara alpina
brunnipennis Dej. (fig. 40) and the Weevil Otiorrhynchus arcticus O. Fbr. (fig. 41),
both widespread on either side. It can be stated, without any exaggeration, that,
zcithin the Arctic of the entire northern circumpolar area, the comparatively narrow
strait between Baffin Island and Greenland has constituted the most effective barrier
to the dispersal of soil-bound animals. The Bering Strait has been far more sur-
mountable (below, p. 293). Though it should of course be admitted that Green-
land is a zoogeographical transition between the Palaearctic and Nearctic regions,
this island— if a choice is necessary— must be regarded as Palaearctic (a good reason
for continued Danish government!).

How is this surprising extension of the European fauna over the islands of the
North Atlantic to be explained?

In the first place it must be stated that the possibilities of passive dispersal (and
at least for the soil-bound, flightless fauna this is the effective agency) would seem
to favour an eastward transport across the Atlantic. Not only the Gulfstream
follows this course (fig. 32), the predominating winds are also western^ and the
barometric minima developed off the northeastern coast of North America, not-
withstanding all irregularity too well-known to meteorologists, move eastward.
Williamson (1954) has given good evidence for how stray birds on migration may
be sluiced across from North America by the aid of subsequent minimum pressures
(fig. 34). In exceptional cases, provided a minimum becomes immobilized to the
south of Iceland (Lindroth, 193 1, p. 524, fig. 42), favourable, mainly southeastern
winds may last long enough to make air-borne transport at least theoretically
possible, for instance from the British Isles to the Faeroes or Iceland, Salomonsen
(1951) has shown quite convincingly how a wind-driven flock of Field-fares (Tj/r^/w^

^ The air movements at high altitude may be different from the winds immediately
above the earths surface. Meteorological balloons, usually travelling above 20 kilometers,
have crossed the Atlantic in both directions. In Europe, the summer winds at this altitude are
usually eastern (Scherhag, 1948, p. 77 a. f.) but rather slow. The biogeographical consequences
of transported air-plankton is a question of survival, which should be tested experimentally.

DiAGR. 7. Species of Colcoptera common to the Faeroes, Iceland, Greenland and
Baffin Island, and selected regions of the Holarctis.

White = flying; black = flightless species.

Dimorphic species counted as flightless where (entirely or partially) represented
by the short-winged form. — Different subspecies treated as species.

Summary of table 7.

The relationship between the palaearctic and nearctic faunas

265

percent g j^j^ W. Sib. Scand Brit. Isl. Faroes Icel. Greenl. Baff. Isl. E.N.Am. W.N. An

100t

100 T

100t

IOOt

266

CHAPTER III

(Photo P. Ardo.)

FIG. 40. The distribution oi Amara alpina brunnipennis Dej. in eastern North America
and the Canadian Arctic. This Carabid beetle occurs also in Alaska and in the moun-
tains south to Colorado. It is probably constantly flightless. The figured specimen
is from Labrador.

pilaris L.) reached Greenland, probably from southwestern Norway, in 1937, and
gave rise to a permanent colonization (fig. 42). Similarly, the Palaearctic Gamma
Moth, Plusia {Phytometra) gamma L., captured in both eastern and western
Greenland, is surely a casual immigrant since it is normally not able to survive
winter even in Scandinavia.

From zoo- and phytogeographical facts it is easy to demonstrate, however, that
eastzvard dispersal, passive or semi-active, occurs more regularly in the North
Atlantic area. The American element in the British flora has little equivalence in
North America (p. 251), this applies also to the Noctuid Moths and a few other
flying insects of American origin which invaded Iceland and the Faeroes (p. 253).
Though several Nearctic species of winged insects and "ballooning" spiders have

The relationship between the palaearctic and near die faunas

267

(From HoLDHAUS, 1924.)

FIG. 41. Distribution of the Weevil Otiorrhynchus arcticus O. Fbr., a flightless,
sluggish, soil-bound insect with bisexual reproduction, and therefore not easily-
spread. The larva is subterraneous and feeds, like the imago, on various plants.
Crosses = subfossils, earlier than Postglacial.

reached Greenland (p. 263) no Palaearctic species, as far as is known, have arrived
in Baffin Island from the opposite direction.

To my mind, only two possibilities remain which, separately or by joint action,
may account for the dominance of Palaearctic, in part strictly European, species in
the faunas of Iceland and Greenland.

In the first place, transport with man may be responsible. Actually, voices have
been raised to the effect that practically the entire lower terrestrial fauna of Iceland
{vide Lindroth, 193 1, p. 507) and the European element of the Greenlandic fauna
and flora (Stephensen, 1921; Ostenfeld, 1926; Jensen, 1928) were introduced with
the early Norse colonists and it cannot be doubted that human transport from
Europe has played an important role in the composition of the fauna of these
islands.

As far as Coleoptera are concerned, I have tried to exclude from the list above
(table 7) all species for which this may be true, according to their choice of habitat
&c., but it is of course by no means impossible that other species also, not at all
connected with human culture, may have been brought over, by pure chance, for
instance, in the Greenlandic fauna: Atheta fungi Gv . (6o°59'-6i°i3'), Lathrobium
fulvipenne Gr. (only one specimen known), Quedius boops Gr. (6o''-62°), and
Scymnus limonii Donis. (3 specimens in the extreme south), which are not found
north of the old Norse colonization area (for this, vide "Atlas", 1921). *

268 CHAPTER III

A removal of these species, however, does not appreciably change the zoogeo-
graphical character of the island and I am completely unable to accept the idea
that, 'for instance, the culture-avoiding, wingless, sluggish, and, contrary to the
majority of species within the genus, obligatorily bisexual Weevil Otiorrhynchus
arcticus O. Fbr. (fig. 41) should have been introduced into Greenland or Iceland
(Lindroth, 193 1, p. 514-515). Also, Porsild (1932) has published weighty argu-
ments against the opinion of Ostenfeld, and others, that man is responsible for the
European element of vascular plants in Greenland.

The trade from Europe to Iceland and Greenland from oldest times had a
character essentially different from that with Newfoundland and the present Mari-
time Provinces of Canada. From the very first time after discovery, the firstnamed
island possessed a permanent population badly in need of all kinds of necessities
(provisions, timber, metals, &c.) and there was certainly no use of ballast on the
outward voyage. Consequently, the experiences gained from the study of the New-
foundland trade (p. 154 a.f.) have no application at all to the naval connections
of the northern islands. This explains why several species, for instance Notiophilus
biguttatus F., Amischa analis Gr., Coccinella ii-punctata L., Barynotus squamosus
Germ., Otiorrhynchus rugifrons Gyll., Sciaphilus asperatus Bonsd., Strophosomus
melanogrammus Forst., and Tropiphorus obtusus Bonsd., in table 7 are regarded as
introduced in eastern North America, but as indigenous in Iceland, where they
occur, in part or entirely, in the most inaccessible parts of the coast.

The most weighty objection to the opinion that the European element of the
faunas of Iceland and Greenland was imported with man is the large amount
of species in common among the species here regarded as indigenous.

From table 7 and diagr. 7, regarding Coleoptera, the following figures can be
extracted: —

Of 1 1 native Palaearctic species in Greenland, 8 (73 %) are common also to Iceland.
Of II native Palaearctic species in Greenland, 6 (55%) are common also to the

Faeroes.
Of 56 native Palaearctic species in Iceland, 38 (68%) are common also to the

Faeroes.

The introduced species^ of the same islands show a different picture: —

Of 23 introduced Palaearctic species in Greenland, 10 (43 %) are common also to

Iceland.
Of 23 introduced Palaearctic species in Greenland, 9 (39%) are common also to

the Faeroes.
Of 99 introduced Palaearctic species in Iceland, 32 (32%) are common also to

the Faeroes.

^ The records of introduced species among the Coleoptera of Greenland are from Hen-
riksen (4939, p. 43-48), of Iceland from Lindroth (1931, p. 171-236) and S. G. Larsson
{in litt.).

The relationship between the palaearctic and nearctic faunas

269

90' 80' W to" SO' tO'iO'^O'lO' 0" 10' JO" 30' fO' 50' 60' 70' ^

FIG. 42. Wind-drift of Fieldfares {Turdiis pilaris L.) to
Greenland in January, 1937.

Black circles = observed specimens. Broken line = prob-
able course of flight.

Unbroken line = isobars of January 19 1937 (at 7 p.m.).
Arrows = wind direction.
(From Salomonsen, 195 i). (From Rosenberg, Faglar i Sverige.)

A similar calculation, and with similar results, was carried out on the Collemboles
of Iceland and the Faeroes (Lindroth, 193 1, p. 516).

The accidental nature of introduction is thus clearly demonstrated. If this
holds true for the unquestionably imported species with a more or less pronounced
synanthropic habitat, how much more due to chance would not human transport
be of species indifferent to or rather avoiding the neighbourhood of man! Yet
within this latter faunal element the species common to the North Atlantic islands
are more, not less numerous. This is the clearest possible proof that their presence
is not due to human agencies.

We have thus rejected the idea that the pronouncedly European character of

270 CHAPTER III

the Icelandic, but also of the Greenlandic fauna could be the result of introduction
alone. At the same time we have denied that the said native element could have been
able to immigrate during present conditions, with the extension of land and sea
now existing in the North Atlantic region. One alternative only remains: the main
part of the fauna, especially of its soil-bound members, reached Iceland and Greenland
over a land-connection now submerged.

This is the old hypothesis of a transatlantic bridge, eagerly advocated above all
by Sharff (1907, 1909, 191 1), but with the fundamental divergence that it did
not extend to the mainland of North America or at least, for some reason, it was
not faunistically effective as far west.

The bottom configuration of the sea is rather favourable for the rise of the land-
connection required (fig. 43). A positive displacement of the shore-line amounting
to less than 600 meters would transform the Wyville-Thomson ridge into an
isthmus connecting Iceland and Greenland with the enlarged European continent
north of the present Scotland but also, it should be pointed out, via EUesmere Is-
land with Baffin Island. It would require very little more uplift to shape another
direct connection between Greenland and southern Baffin Island.

An important question is whether an un-interrupted connection would be necess-
ary in order to allow an immigration of the said faunal element from Europe to
Iceland and Greenland. Considering its large share of flightless forms, in Iceland
26 species of 56 (46 per cent), in Greenland 6 of 11 (55 per cent), constituting the
Palaearctic element, the answer seems to be in the affirmative. Ocean water of
more than 30 pro mille salinity causes physiological desiccation of a terrestrial
animal (Palmen, 1944, p. 155 a.f.; Lindroth, 1949, pp. 600, 613). Furthermore, the
heavy Weevils, such as Otiorrhynchus arcticus O. Fbr., perhaps the zoogeographic-
ally most important member of the Iceland-Greenland fauna (fig. 41), soon are sunk
to the bottom by the lapping waves (Lindroth, 193 1, p. 485). Even the Channel,
only 31 km. at its narrowest, constitutes a marked faunistic barrier (Lindroth,
1949, p. 613), though the British Isles were connected with the mainland in early
postglacial times. The unbridgeable obstacle to flightless insects formed by the
Davis Strait has just been demonstrated.

When it comes to dating the existence of the assumed Scotland-Greenland bridge
a biologist is largely up in the air. This much is certain, however, from geological
evidences: it cannot have persisted in late glacial or postglacial times. On the
other hand, the faunas of Iceland and Greenland bear no sign of long isolation.
Proposed endemic species all belong to insufficiently investigated animal groups,
such as Diptera, Hymenoptera and Hemiptera Homoptera among the insects. The
Coleoptera and Macrolepidoptera of these islands do not even contain clearly
defined endemic iwfepecies. Such, it is true, have been established among Mammals

The relationship between the palaearctic and nearctic faunas

271

*«?-

FIG. 43. The bottom configuration of the North Atlantic. The detailed figures give
greatest depth of the ridge or plateau between the islands.

(From LiNDROTH, 1931, supplemented by Dr Borje Kullenberg, Gothenburg, in lift.)

and Birds, especially of the Greenland fauna, but there is clear evidence in favour
of the view that subspecies formation is often a rapid procedure among Verte-
brates. The development into new "subspecies" of the Varying Hare, Lepus
timidus L., and the House-Mouse, Mus musculus L., on the Faeroes, both introduced
with man (Degerbol, 1940), is a good illustration.

By a process of elimination we thus arrive at the opinion that the land connect-
ing Greenland and Iceland with Europe must have existed in Pleistocene time
and before the last glaciation (Wiirm, Wisconsin). This is also supported by the
present distribution of the Icelandic fauna which clearly indicates survival of
most of its indigenous members within two isolated glacial refuges in the south
and southeast (fig. 46; Lindroth, 193 1, pp. 489 a.f., 557-563).

Another remarkable fact is the discovery, in a fossiliferous interglacial layer in
SW Iceland, of insect rests belonging to 6 species of Coleoptera {Nebria gyllenhali
Schnh., Patrobus septentrionis Dej., Pterostichus diligens Sturm, Agabus solieri

272 CHAPTER III

Aube, Tachinus corticinus Gr., Byrrhus fasciatus Forst.),^ all of them members of
the present Icelandic fauna (Thorkelsson, 1935).

The powers which may have affected displacement of the shore-line in favour
of the extension of land in the North Atlantic area are eustatic, isostatic, and tectonic
movements.

(a) Eustatic movements of the sea, implying a world-wide change of the water
volume of the oceans, in Pleistocene time were caused first and foremost by the
glaciations. Part of the precipitation was stored on land in the form of glaciers and
thus withdrawn from the oceans, the level of which sank correspondingly. Cal-
culations of the vertical amount to which this happened are, as a matter of course,
highly diverging. They depend entirely on the estimated simultaneous volume
of all glaciers of the world during a certain phase of a glaciation. For the maximum
of the last glaciation (Wiirm, Wisconsin) the figures are usually given as something
between 90 and 100 meters (Zeuner, 1950, p. 129; Woldstedt, 1954, p. 289).
During the last but one, the '' great glaciation" (Riss, Illinoian), the regression of the
sea must have been more extensive but it cannot have amounted to more than one
fourth of the vertical distance necessary for realizing the postulated Scotland-
Greenland bridge. It may have contributed to its rise but was not alone responsible
for it.

Eustatic regression of the sea is perfectly and positively connected with glacia-
tion (seen from a global point of view). Though this did not necessarily have its
maximum extension contemporaneously everywhere in the Northern Hemisphere,
countries situated as far north as Iceland and Greenland can never have remained
climatically unaffected by the ice during a glacial period. This means that, if eusta-
tic regression has played a part in the existence of the Scotland-Greenland bridge,
this must have been realized during a period of arctic or at least subarctic climate on
the islands considered. This assumption may seem tempting because, if so, the
faunistic barrier between Greenland and Baffin Island could be explained as a
barrier of ice. Facts, however, argue against this. The coleopterous fauna of
Iceland contains one single arctic species, the Water-beetle Colymbetes dolabratus
Payk., immigrated from a western direction. The rest of the indigenous fauna is
boreal, not even subarctic; many of its members do not ascend to timber limit
anywhere on the earth. Among the Coleoptera of Greenland there are only two
Palaearctic species, the Staphylinids Atheta hyperborea Brund. and Gnypeta cavi-
collis J. Sahib., which may be termed arctic; both are winged.

A satisfactory explanation of these conditions is found only in the assumption
that the Scotland-Greenland land-bridge existed during a time of boreal climate and

^ In addition unidentified fragments of a Hydroporus, not living in Iceland of today.

The relationship between the palaearctic and nearctic faunas 273

eustatic movements of the sea have therefore played at most a quite inferior role in
its formation.

(b) Isostatic movements are local and due to ice-pressure during periods of
glaciation. In the central part of an inland ice they may amount to more than 200
meters, for instance in Scandinavia. If Greenland were cleared of ice, it would
undoubtedly rise to a considerable degree but, as we have learned from investiga-
tions in Scandinavia, the "isobase o", the line of equilibrium, with no postglacial
upheaval and, consequently, no previous glacial down-pressure, runs close to the
former ice-margin, or even inside it. The rise would affect only the immediately
surrounding sea-bottom.

An increase of the Greenlandic inland-ice, on the other hand, and a contempor-
aneous ice-cap over Iceland, would possibly cause a slight upheaval of the interjac-
ent sea-bottom but involve very little isostatic effect on the almost three times as
long sea-bed between Iceland and Scotland. The proposed land-bridge could not
arise, or even become considerably favoured, by isostatic movements.

(c) The possible influence of tectonic movements and volcanic activity remains
to be considered.

Iceland is a country with still vividly living volcanic powers. In comparatively
recent time, geologically speaking, these have been active within a much wider
area. Basalt of the same kind as that which built up the frame of the Icelandic
plateau occurs in parts of East Greenland, on the Faeroes, on some of the Inner
Hebrides, and in northeastern Ireland. Certain fossiliferous layers of this system
in Iceland, the Faeroes and Ireland have been regarded as homologous and con-
temperaneous. It is generally assumed that these now widely separated basalt
occurrences are only remnants of a former continuous land, formed through vol-
canic eruptions in Tertiary time, from Eocene and onwards. Soon after its rise,
this mighty basalt plateau became subject to violent tectonic movements and
eventually was broken down to present conditions.

Geologists apparently are not able to answer the question most important to a
biogeographer: at what time was the connection broken and the present isolation
of these islands definite? According to Thoroddsen (1914), Iceland was already a
separate island at the beginning of the Pliocene. Later geological students have
thought that Thoroddsen generally overestimated the age of Icelandic geological
formations but the current view seems to presume that the Tertiary transatlantic
basalt isthmus was broken down almost to a definite extent before Pleistocene
time.^

^ Dr. Sigurdur Thorarinsson, the wellknown Icelandic geologist, informs me, however,
{in litt.) that to his mind no definite facts seem to argue against the possibility that the Ter-
tiary land-bridge persisted into part of the Pleistocene period.

18 — 565597 Lindroth

274

CHAPTER III

Since the eustatic and isostatic movements of the shore-line in Pleistocene
time apparently were insufficient to create a bridge connecting Greenland and
Iceland with Europe, as described above, it is tempting to think of a prolonged per-
sistence of the Tertiary basalt plateau into part of the Pleistocene. The absence of
undoubtedly endemic species, and even true subspecies, in the insect fauna of
Iceland and Greenland prevents the acceptance of survival in situ since Tertiary
(Pliocene) time. Considering this, the present author (Lindroth, 193 1, p. 550-557)
arrived at the conclusion that the proposed land-connection most probably should
be dated to the last (Riss-Wiirm) interglacial period.

Facts contradicting this view have since appeared in the form of the discovery
of fossiliferous deposits in Central Sweden from the said period. The included
insects (Lindroth, 1948) give evidence of a climate at least as warm as the present
one during part of the Riss-Wiirm Interglacial in Scandinavia and this cannot be
considered in accordance with a situation implying that the Gulfstream was
prevented by a land-bridge from entering the Norwegian Sea^.

It should also be considered in this connection that the fauna of Iceland is
originally boreal, not arctic, the few exceptions {Colymbetes dolabratus, &c.) being
easily dispersed animals independent of any land-connection. The invasion of this
boreal fauna therefore took place in the warm middle part of some interglacial
period. Such a violent oscillation, from one stage: "land-bridge, blockade of
Gulfstream",— to another: "land-bridge sunk, 200 meters at least, Gulfstream
free passage",— cannot be accepted to have taken place in a time so short as the
temperate phase of an interglacial period.

It seems that a "removal" of the land-bridge one step further back in time, to
the last but one Interglacial (Mindel-Riss), is the most likely solution. From a
biological point of view, the difficulty in accepting this idea seems to me to lie
not so much in the lack of endemics in Iceland-Greenland, as in the necessity
of accepting survival in these islands through two subsequent glaciations, the
first of them being the severe Riss.

Yet if the land-bridge existed in the Mindel-Riss Interglacial, the climatic in-
fluence of the dislocated Gulfstream, at least on Iceland, ought to have been
highly increased and these favourable conditions may have lasted during part of
the following Riss glaciation.

There is one special reason to think that in Iceland, to a lesser degree also in
Greenland, local conditions may have been favourable enough, even during Riss,
to make a survival possible: the presence of volcanic hot springs. Tuxen (1944, p.
189-199) has discussed this question and rightly concludes that a field of hot

^ Vide also the table of climate and vegetation during the last interglacial period in Den-
mark by Jessen & Milthers (1928, p. 336).

The relationship between the palaearctic and nearctic faunas

275

c d

(Photo P. Ardo.)

FIG. 44. Bembidion grapei Gyll., a Carabid beetle showing polymorphism of the hind-
wings. Types a-b are counted as macropterous, c-d as brachypterous. — Ice-
landic specimens.

springs, whatever its effect otherwise, will never be able to withstand an approach-
ing land-ice. But thermal activity, in Iceland often giving the springs a temperature
above boiling-point, may considerably ameliorate the loco- and microclimatical
conditions of a possible ice-free refuge.

This opinion is confirmed by the occurrence of a Carabid beetle, Bembidion
grapei Gyll., in Iceland. It shows dimorphism or, rather, polymorphism with
respect to the hind-wings: in some specimens these are full and no doubt function-
ing (2 specimens were found in wind-drift material in Finland; Palmen, 1944, p.
37), in others they are more or less reduced and cannot be used (fig. 44). Breeding
experiments with another Carabid beetle, Pterostichus anthr acinus 111. (Lindroth,
1946) have shown that the phenomenon has a heriditary base, "short wing"
being a dominant. The full-winged individuals are therefore homozygotes and
owing to their superior powers of dispersal usually form a genetically pure zone
at the periphery of the species' area. At any rate, predominance of short-winged
specimens is characteristic of old populations.

Possibly more than one gene mutation is involved in the case of Bembidion
grapei (fig. 44), since different stages of wing-reduction occur, but the macro-
pterous individuals also in this species are homozygotes, as is quite evident from the
distribution in Fennoscandia (Lindroth, 1949, p. 401-405). The brachypterous
form is predominant along the Norwegian coast, within the area of ice-free refuges

276

CHAPTER III

FIG. 45. Distribution of long-winged (white) and short-winged (black) specimens
of the Carabid beetle Bembidion grapei Gyll. (fig. 44) in Fennoscandia. Cf. legend of
fig. 14 (p. 153).

The occurrence of the short-winged form indicates survival on ice-free refuges
of the western and northern coast during the last glaciation.

The relationship between the palaearctic and nearctic faunas

277

FIG. 46. Distribution of long-
winged (white) and short-
winged (black) Bembidion
grapei Gyll. (fig. 44) in Ice-
land. The two main refuge
areas during the last glaciation
are indicated. Cf. fig. 45.

during last glaciation (Wiirm), whereas a purely macropterous population inhabits
the eastern parts of Sweden and almost all of Finland (fig. 45).

Iceland is too small an area to have retained, in any part of the island, a purely
macropterous population. The postglacial period has been sufficiently long to
allow a universal colonization of the brachypterous form as well, and no obstacle
comparable to the mountain range of Scandinavia has prevented its dispersal.
The geographical distribution of the two forms (fig. 46) therefore, at first glance,
seems rather confused and due to chance. That is not the case, however. The
brachypterous form is most marked within two areas, around the EyjafjallajokuU
in the extreme South and along the coastal margin of the Vatnajokull in the
Southeast, which for other reasons are regarded as the main refuges during the
Wiirm-glaciation (Lindroth, 193 1, p. 489 a. f.). The index macropterous: brachyp-
terous is here 0.2 and o.i, respectively. For the rest of the island this figure is
0.7, slightly less in the Southwest, 0.6; in the North 0.9, in the isolated north-
western peninsula 1.5.

The interesting contribution to our present discussion is that in Iceland, outside
the two main refuge areas, there is a marked contrast in the occurrence of the
brachypterous form of Bembidion grapei in the immediate vicinity of hot springs,
as compared with other localities. Apparently the species is favoured by the warm
conditions. One would expect, then, that the hot springs would absorb in the
first place the most easily dispersed individuals from the mixed populations of
the surroundings and that consequently the macropterous form would be in the
majority around them.

The exact opposite is the case. The index macropt.: brachypt. for individuals
from the vicinity of hot springs is 0.3, for all others (outside the two refuge areas
of the South), 1.3 (diagr. 8).

278

CHAPTER III

per cent S E refuge

ICELAND
S refuge Hot springs

GREENLAND

Other places

Specimens

DiAGR. 8. Bembidion grapei Gyll. Distribution of four types of hind-wings (a-d,
fig. 44) among four types of habitats in Iceland and in Greenland.
White = full-winged; black = strongest wing-reduction.

In accordance with the current interpretation of the distribution of wing-
dimorphic forms, the most likely explanation is that the surroundings of hot springs
are inhabited by old populations. The index in favour of brachypterous forms is
almost the same as for the two large refuge areas in southern Iceland. To my mind,
this argues in favour of an assumption that, locally, small populations of terricolous
animals were able to survive the last glaciation also in other parts of the island in
the immediate neighbourhood of hot springs. For one of these localities, Krisuvik
in the Southwest, where Bembidion grapei (25 specimens) shows an index of 0.5, the
assumption of a glacial "micro-refugium" is confirmed by the isolated occurrence
of the Staphylinid beetle Othius melanocephalus Or., one of the most undoubted
"survivors" of the Icelandic fauna (Lindroth, 193 1, p. 484 a.f.).

In Greenland, Bembidion grapei occurs almost exclusively in the brachypterous
form (fig. 47; diagr. 8). Apparently selection has been strongly in favour of flightless
individuals in small, isolated populations, particularly during times when glacia-
tion was more total than at present. A location of refuges based on the distribution
of Bembidion grapei seems impossible but it should be mentioned that a large
series of 65 exclusively short-winged individuals from Julianehaab in the South-
west is from the vicinity of the most famous hot springs of Greenland.

The relationship between the palaearctic and nearctic faunas

279

R«pn>di*c*<«t tM OaodaiNt InsMui. f

FIG. 47. Distribution of long-winged (white) and short-winged (black) Bembidion
grapei Gyll. (fig. 44) in Greenland. Most of the few long-winged specimens are
found near the periphery of the species' area, in the north and the south.

The best criterion for the existence of the Scotland-Greenland bridge during
any of the interglacial periods would be the discovery of contemporaneous de-
posits in Scandinavia indicating an arctic climate contrasting with temperate, or
warmer, conditions in central and western Europe. Until then the dating of
this land-connection, which from a biogeographical point of view seems inevitable,
must probably be postponed.

Meanwhile it is highly desirable that botanists reconsider the problems of
"Westarctic" and other Amphiatlantic plants, and also of the Greenland flora,
from a more realistic point of view. Above all these ought to be treated by careful
investigation, including experiments, of seeds and other diaspores, in order to
get a clear idea of their different modes of dispersal. Speculations founded exclu-
sively on maps of distribution, as now often happens, do not give satisfactory re-
sults. In this way, botanists would also be able to approach the real nature of a

28o CHAPTER III

possible high-arctic route of migration, Greenland-Spitzbergen, assumed by some
authors (for instance Hansen, 1930, p. 226; Nordhagen, 1935, p. 157), to which
zoologists seem to be unable to make a contribution.

Affinities in the south

Animal distribution shows a few spectacular instances of subtropic or "low boreal"
forms indicating faunistic relationship between the southern parts of North
America or the West Indies and Europe, including adjacent regions of North
Africa and western Asia.

Usually the forms in question are not identical. Possible exceptions are two
Carabid beetles, Nomius pygmaeus Dej. and Blethisa eschscholtzi Zoubk., and one
Wood-louse (Terrestrial Isopod), Tylos latreilli Ad. & Sav. The latter inhabits
the epilittoral zone of the sea-shore of the Mediterranean region and the Atlantic
coast, north to central France, south to Dakar; on the American side, Florida, At-
lantic Central America, the Bermudas and West Indies. Contrary to Vandel (1945,
p. 229), I find the most probable explanation of this disjunct area to be introduc-
tion with ships from Europe into America. The habitat of the species quite invites
transport.

The Centiped Theatops erythrocephalus C. L. Koch, from southern Europe,
mentioned by Schmidt (1946, p. 145) as indigenous in western U.S.A., is no
doubt introduced in the latter area.

One would perhaps feel inclined to apply the same view to Nomius pygmaeus,
the more so since, in North America, it has repeatedly been observed in villages
and towns, even indoors, as a famous "stink beetle". In the Old World it is a grand
rarity and the assumption just mentioned would then necessarily imply introduc-
tion /rom North America. This, however, has become less probable since Basilewsky
(1954) has described a very closely related species, N. schoutedeni Basil., from
Ruanda Urundi in the interior of Belgian Congo. Now Nomius belongs to a
quite isolated group (usually termed Tribus) among the Carabidae, the few genera
of which have a pronouncedly disjunct, obviously relict, distribution, with several
representatives in Australia (Emden, 1936, p. 50-51; Jeannel, 1941, p. 290-292). It
may be suggested that not only subtribes and genera of this group, but also the
actually existing species, are very old and the present distribution of Nomius
pygmaeus may be regarded as a disjunct relict area which in itself gives no clue
to the history of the species.

The same may be true for Blethisa eschscholtzi (fig. 56). One single specimen is
known from North America, vicinity of Sanderson, SW Texas, but there seems
to be no reason to doubt the correctness of the locality label or the indigenous

The relationship between the palaearctic and nearctic faunas

281

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FIG. 48. World distribution of the Salamander genus Hydromantes, in California
H. platycephaliis Camp., in Europe H. genei Schleg. {Spelerpes fiiscus Schreib.).
The figured specimen is of the Californian species (from Stebbins, 1951).

occurrence on this far inland spot. Otherwise it is restricted to the steppe region
north of the Caspian Sea and of western Siberia. B. eschscholtzi is the most primi-
tive member of the genus Blethisa (Lindroth, 1945b, p. 13) and Semenov (1935,
p. 275) regarded it as a "relictum faunae tertiariae elementum". It is an old
species with disjunct relict area.

Among Vertebrates, the affinities in the south between Europe and North
America are restricted to common genera, or higher taxonomic units. The most
conspicuous cases are the following: —

Fam. Umbridae, fresh-water fishes, named Mud-minnows, with genus Umbra,
has two species in North America and one in Europe, U. krameri Wahlb. {lacustris
Gross), restricted to the basin of the Danube.

Among Salamanders (Urodela) there are two links suggesting a connection. —
(i) The famous Olm, Proteus anguineus Laur., a true inhabitant of the karst-caves
of SE Central Europe, possesses its closest relatives in the Mud Puppies, genus
Necturus, with 7 free-living species in eastern North America.— (2) The Pletho-
dontidae, abundant in North America, but lacking in Asia, are represented in
Europe (SE France, N Italy, Sardinia) only by a species of Hydromantes, genei
Schleg. (Spelerpes fuscus Schreib.), with two subspecies (Mertens & Miiller, 1940,

282 CHAPTER III

p. 13). The genus occurs nowhere else on the earth, except in California (fig. 48),
where the Mount Lyell Salamander, H. platycephalus Camp., is restricted to the
BoreaJ zone of Sierra Nevada (Stebbins, 195 1, p. 146).

The well-known genus Emys, among fresh-water Turtles, contains only two
species, orbicularis L. of Europe, North Africa and Hither Asia, and blandingi
Holbr., of central and eastern North America.

During present conditions a dispersal across the Atlantic of these Vertebrates,
unable to endure salt water, is out of the question, except with the aid of man,
and this again would be an absurd idea since in no case the same, only related
species occur on both sides.

Similar instances of related, but not identical, animals of a southerly Amphiat-
lantic distribution are more numerous among Invertebrates but it is sufficient
to mention a few examples.

Darlington (1934, p. 86-88) discovered and described two species of the Carabid
genus Perileptus, from Cuba and Jamaica, a most surprising event since this was
known as an entirely Old World genus and, moreover, the two species belonged
to the areolatus group, confined to Europe, North Africa and the Macaronesian
Islands; two further species were subsequently described from Hispaniola (Haiti)
and Puerto Rico (Darlington, 1935). A quite similar distribution is recorded for
the otherwise Old World Carabid genera Stylulus, with the species nasulus Schauf.
on the island of St. Thomas, and Lymnastis, with two species in Cuba and Guate-
mala.

These cases were stressed by Jeannel (1937, p. 356 a.f.) as evidence of an earlier
land-connection between the West Indies and the Mediterranean region. Darling-
ton (1938) has strongly opposed this view. To his mind, all of them (or their
ancestors) have been carried across to America with the trade winds. Though this
may be true for Perileptus and Lymnastis, being constantly or (Lymnastis) individ-
ually long-winged, I cannot agree that this possibility exists for the flightless,
blind Stylulus, subterranean in habit. The almost constantly very restricted
distribution of the individual species of subterranean beetles, for instance in
the Alps, where there are often heavy "Fohn" -winds, shows that their powers of
dispersal are extremely poor. On the other hand, they are among the most difficult
insects to collect and I am quite convinced that the actual distribution of genera
and higher groups of subterranean Carabidae is known only in fragments. It
should also be observed that the single West Indian species of Stylulus hitherto
known belongs to a subgenus of its own.

However, Darlington is certainly right in rejecting Jeannel's idea of land-con-
nection, and his suggestion that small flying Carabids may be carried by air currents
at high altitudes was strongly confirmed by the investigations of "aerial plankton"

The relationship between the palaearctic and nearctic faunas 283

by Glick (1939, p- 29-30), as only small species, of the genera Bradycellus (Steno-
cellus), Micratopus, Microlestes (Blechrus), Tachystodes, and Tachys, were captured
at 5,000 feet or higher, one specimen of Microlestes pusio Lee. (less than 2 mm.)
at 10,000 feet.

Vandel (1945) has drawn attention to the almost Amphiatlantic distribution of
the genera Porcellio and Metaponorthus and some other Terrestrial Isopods, and
regards this (p. 265-266) as a proof of Wegenerian connection. His maps show,
to my mind, that the world distribution of Wood-lice, notably in southern and
eastern Asia, is not yet sufficiently investigated to allow for conclusions of this
kind.

Jeannel (1942, p. 115) regards the Palpigrad Arachnid genus Koenenia as a case
of direct transatlantic faunal exchange. Since one species is known also from Siam
(Kastner, 1932, p. 98), this argument has lost its force.

The Cockroach (Blattodea) genus Arenivaga contains three subgenera, Arenivaga
s. str. being strictly American (southern U.S.A., Mexico), whereas subgg. Psammo-
blatta and Heterogamisca are chiefly Mediterranean (S Europe, N & E Africa,
Hither Asia). Bey-Bienko (1950, p. 300-323), who made a rearrangement of the
genus, regards its distribution (p. 302) as the result of a direct contact, in Wegener's
sense, across the present Atlantic, but the occurrence of species of Psammoblatta
as far east as Baluchistan, Afghanistan and West Turkestan makes this assumption
somewhat adventurous.

The Oleacinidae, a family of terrestrial Gastropod Molluscs, is restricted to
America north to southern U.S.A., and the Mediterranean region (Pilsbry, 1946,
p. 188). However, it is a very old type, known in fossil state right back to Upper
Cretaceous of Europe, and it must be assumed that the family has undergone
great, still unknown changes of area during the course of the Tertiary.

It is not surprising, with the above examples in mind, that several of the earlier
zoogeographers, especially, have tried to explain this type of disjunct distribution
by the assumption of some kind of land-connection also within the southern parts
of the North Atlantic, usually thought to have joined the Iberian Peninsula and
North Africa with the present West Indies {vide Scharff, 191 1, pp. 173, 214, 220,
271 a.f., fig.i4). Ihering (1927, pp. 26, 220) even boldly constructed one of "his"
bridges, the "Archatlantis", exclusively for the convenience of Sirenians of the
genus Trichechus, inhabitants of the sea, though as a rule, at least, of brackish
water (vide Holdhaus, 1927-28, p. 1051). In later years the supporters of land-
connections in this part of the Atlantic have usually been pronounced Wegenerians
who in cases of distributional pattern of the type exemplified above find biological
evidence in favour of the hypothesis of continental drift (vide next paragraph).

Yet it seems to me that there is no need at all for a land-connection to explain

284

CHAPTER III

the cases here treated. We have to reahze that they indicate an old contact, only in
some few insects expressed through identity of species, nota bene, of a primitive
and no doubt old type, otherwise in genera and families in common, and that we
must go well back in Tertiary' time for an explanation. Great faunal changes indeed
have taken place since then.

This may be illustrated by an example. The Cryptobranchidae among the Sala-
manders are represented in the recent fauna of the earth only by the well-known
"Giant Salamander", Megalobatrachus maximus Schleg., of China and Japan, and
Cryptobranchus alleghaniensis Daud. from the Mississippi basin. But the European
fossils from Miocene referred to genus Andrias (the famous A. scheuchzeri Tschudi,
"Homo diluvii testis", and others), according to Herre (1935, p. 48), are generically,
possibly even specifically, identical with the still living Megalobatrachus. Supposing
the contrary had happened, that the European form had happened to survive but
the East Asian had become extinct and no fossil remains discovered,— and we
would have had a perfectly Eur-American family of Salamanders, with several
false theories as a consequence.

The Baltic amber of Europe, usually considered of late Eocene age, contains
numerous examples of genera, families or higher taxonomic categories, parti-
cularly of insects, now absent from the continent or even from the entire Holarctic
region, examples of which are given below. As argued by Darlington (1948, p.
2 a.f.), recession and extinction plays a role equal to evolution and spreading in
faunal history, and extinction usually starts with the division of one large area into
small, separate ones.

The Amphiatlantic animals of the more southern type here treated apparently
all have entered this stage of recession. They are remnants of a large, though not
necessarily contemporaneous, area across the Asian continent. Thus I can add very
little to the views expressed by Matthew (191 5), Reinig (1937), Schmidt (1946),
Simpson (1947), Darlington (1948), and others.

The present distribution of these animals is of pronounced relict character and
necessitates no other change in extension of the present continents than the unani-
mously accepted Bering land bridge.

Wegener — pro and contra

The hypothesis of continental drift was in part based on biogeographical data by
Wegener himself (1929, p. 99-124) and subsequent students accepting his idea
have often used examples of animal distribution as confirmation.

Several zoologists and some botanists at first more or less enthusiastically accepted

The relationship between the palaearctic and nearctic faunas 285

the theory, for instance Michaelsen (1922), Jaschnow (1925), Brehm (1926),
Herre (1935), and others; among botanists, Bocher (1938), Steffen (1941), Walter
(1954, p. 53 a.f.); vide also "Atlantisheft" (1939). In later years the critical voices
have been in the majority but some zoologists are still ardent supporters. Thus,
Modell (1943) thinks there is evidence for a Tertiary transatlantic connection in
Wegenerian sense from the fossil Mollusc fauna of Europe, but the facts brought
forth by him are quite in accordance with the late Tertiary European flora, as
investigated by Szafer (1954), which needs no such explanation (below, p. 310).

His most loyal advocate among zoogeographers Wegener has found in Jeannel
who, in a long series of books and papers, the most comprehensive being that of
1942, has tried to explain any imaginable pattern of animal distribution as the
result of continental drift.

Jeannel's method is not one of science, but of faith. He simply "believes" in
Wegener and, refusing to discuss any other explanation, makes "dogmatic state-
ments with no distinction between fact and opinion". The last citation is from
Darlington's excellent review (1949) of Jeannel's "Genese des Faunes Terrestres"
(1942), to which the interested reader is referred.

Since, according to Wegener (1929, p. 20), the Atlantic was the ocean latest
formed and the continents bordering its northern part were the last to separate
(in Pliocene or even Pleistocene time), one would expect to find particularly clear
evidence of direct faunal exchange between Europe and North America, provided
the theory is true. But this is not so.

As shown on preceding pages of this chapter, the number of Eur- American ani-
mal species lacking in Asia, which cannot easily be explained either by human
transport or as the result of dispersal across the Atlantic under conditions similar
to those of the present day, is so exceedingly small that any idea of general
contact between the European and American faunas in Pleistocene or Late Tertiary
time is sharply contradicted, other than through Asia and the present Bering
Strait. This has already been pointed out, in part, by 0kland (1927, pp. 361,
363) but, according to him, facts speak in favour of the "bridge theory".
Actually, not even the assumption of a late (Pleistocene), complete transatlantic
land-bridge, only of one joining Greenland and Iceland with the mainland of
Europe, seems to be a zoogeographical necessity.

It must be frankly admitted that several single biogeographical facts would be
more easily understood by resorting to Wegener's theory of continental drift: i.e.,
the distribution of the genera Hydromantes (p. 281) and Umbra (p. 281), of Cepaea
hortensis O. F. Miill. (p. 234 a.f.) and Tylos latreilli Ad. & Sav. (p. 280), provided,
in the latter cases, the idea of introduction has to be abandoned. But why should
the (superficially regarded) simpliest explanation always be the true one? Widely

286 CHAPTER III

scattered relict occurrences (Hydromantes, Umbra, &c.), usually a step towards
extinction, do not necessarily imply earlier connection at shortest distance, the
fossils "often showing us the contrary.

It is questionable if, from the stand-point of a biologist, there is any need at all
for transatlantic land-connection in any geological period. Simpson, on palaeo-
zoological evidence, first (1940, p. 149) postulated "the inference of a wide-open
corridor" between Europe and North America in the Lower Eocene but later
(1947, pp. 658-659, 666, Footnote) regarded a substituting Pacific connection
during this period as at least equally probable.

Whether the theory of continental drift is true, is a problem for geologists
and geophysicians. The only contribution that can be delivered by a biogeographer
is the declaration that, at least as far as the North Atlantic area is concerned, the
continental drift, if considered a reality, took place in so early a period that its
biological consequences cannot be traced. Also, the annual rate of drift supposed
by Wegener for the North Atlantic is extravagantly high (Zeuner, 1950, p. 355).

Circumpolar animals

The word "Circumpolar" should not be interpreted too literally. No animal
species inhabits all continents and islands surrounding the North Pole. The
concept is here used to cover species occurring in the northern parts of Europe,
Asia and North America without conspicuous interruptions of area. The term
"Holarctic" is not a synonym because it only means that a species is indigenous
in both the Palaearctic and the Nearctic region, without any demand for continuity.
Yet all Circumpolar species are, of course, Holarctic at the same time.

Another question is how pronounced northerly a species' area is bound to be
in order to be called Circumpolar. The "Circumtropical" species, such as the Dra-
gonfly Pantala flavescens F. (Rensch, 1950, p. 136, fig. 109), must be excluded.
A subtropical animal distributed round the earth at least belongs to either Hemi-
sphere and may be included here, but the instances of this geographical type are
so few that their placing has almost no interest.

As a matter of fact, the zonation of the circumpolar species from north to south,
as well as vertically in the mountains, reveals interesting features. Carabid beetles
are sufficiently known in this respect to serve as an illustration. The distribution
of the 45 known circumpolar species among the vegetation belts as defined by
Scandinavian botanists {vide Lindroth, 1949, p. 436-448), and corresponding
zones of Asia and North America, is shown in table 8.

The table 8 gives clear evidence that the subarctic (subalpine) region contains

The relationship between the palaearctic and nearctic faunas

287

TABLE 8. Circumpolar Carabid beetles and their distribution among the
climatically determined zones of vegetation.

Species with incomplete circumpolar area in brackets. ^
Cross in brackets indicates rare, and perhaps not regular, occurrence.
The first column gives the state of the hind-wings, according to the following
categories: m = macropterous (probably flving); b = brachypterous (flightless, doubt-
ful cases in brackets); d = dimorphic (both forms represented within the species).

Agonum bogemanni Gyll. . .

A. consimile Gyll

(^A. exaratiim Mnh.) ....

A. tnannerheimi Dej

A. quadripunctatum DeG. . .

A. thoreyi Dej

Amara alpina Payk

A. erratica Dft

A. hyperborea Dej

{A. interstitialis Dej.) ....

A. lunicollis Schio

A. quenseli Schnh

A. torrida 111

{Bembidion daiiricum Mtsch.) .

B. grapei Gyll

B. hasti C. R. Sahib

{B. hyperboraeorum Munst.) .

B. lapponicum Zett

B. obscurellum Mtsch. . . .

B. petrosiim Gebl

B. quadrimaculatum L. . . .

B. transparens Gebl

{B. yukonum Fall)

BLethisa multipunctata L. . .
Calathus micropteriis Dft. . .
(Carabus trwicaticollis Eschz.)
Diachila arctica Gyll

D. polita Fald

Dyschiriits helleni J. Miill. . .
Elaphrus lapponicus Gyll. . .

E. ripariiis L

Wings

Regio arctica

Reg. sub-
arctica

Reg. coni-
ferina

Reg. quer-
cina

Southern
regions

superior

media

inferior

m

_

_

+

+

+

m

—

—

+

+

( + )

—

—

m

—

—

+

+

—

—

—

d

—

—

—

+

+

(+)

—

m

—

—

—

+

+

+

+

m

—

—

—

+

+

+

+

(b)

+

+

+

+

(+)

—

—

m

—

+

+

+

+

—

—

m

—

—

+

+

+

—

—

m

—

—

—

+

+

(+)

—

m

—

—

( + )

+

+

+

+

d

—

+

+

+

+

+

+

m

—

—

+

+

+

—

—

d

—

—

+

+

—

—

—

d

—

—

+

+

+

( + )

—

m

—

+

+

+

(+)

—

—

m

—

—

+

+

(+)

—

—

m

—

—

+

+

( + )

—

—

m

—

—

—

+

+

+

—

m

—

—

—

+

+

—

—

m

—

—

—

—

+

+

+

d

—

—

—

+

+

+

+

d

—

—

+

+

—

—

—

m

—

—

—

+

+

+

+

b

—

—

+

+

+

+

+

b

+

+

+

( + )

—

—

—

m

—

—

+

+

+

—

—

b

—

+

+

+

—

—

—

b

—

■ —

+

+

+

—

—

m

—

—

+

+

+

(+)

—

m

—

—

+

+

+

+

4-

1 The following species are possibly circumpolar though their distribution is insufficiently
known: Bembidion cremdatum F. Sahib., B. mckinleyi Fall, Dyschirius politus Dej.

288

CHAPTER

III

{Harpalobrachys leiroides

Wings

Regio arctica

Reg. sub-
arctica

Reg. coni-
ferina

Reg. quer-
cina

Southern
regions

superior

media inferior

Mtsch.)

ni

—

—

+

-r

+

—

—

Harpalus fuliginosus Dft. . .

m

—

—

+

-r-

4-

+

+

(//. nigri tarsi's C. R. Sahib.) .

in

—

— •

4-

-t-

'^

—

~

Miscodera arctica Payk. . . .

in

—

+

-i-

4-

+

4"

+

Nebria gyllenhali Schnh. . .

(b)

(+)

+

+

4-

4-

+

4-

N. nivalis Payk

(b)

-i-

+

+

(+)

—

—

—

Notiophilus aquaticiis L. . . .

d

-t-

+

+

4-

4-

—

4-

Patrobus septentrionis Dej. . .

m

+

-r

+

4-

+

—

—

Pelophila borealis Payk. . . .

m

—

( + )

+

4-

4-

(+)

■ —

Pterostichus adstrictus Eschz. .

m

—

+

+

4-

(+)

—

P. brevicornis Kby

b

—

—

+

4-

4-

—

—

(P. vermiculosus Men.) . . .

b

+

+

+

4-

—

—

Tachyta nana Gyll

m

—

—

—

( + )

4-

4-

( + )

Trichocellus cognatus Gyll. . .

m

—

—

+

4-

4-

4-

+

Total 45 (36) species

—

6 + (i)

12+{l)

.33+(i)

44 + (2)

33 +(5)

17 + (6)

i3+(i)

most circumpolar species, actually all but one {Bembidion quadrimaculatum L.),

The unwooded arctic (alpine) zone is inhabited by 33 (or 34) species, that is 73

(or 76) per cent, the conifer belt (the "taiga" or highboreal zone) likewise by 33

(or even 38) species, that is 73(-84) per cent. The number sinks rapidly as the

climate grows milder toward the south (and downward).

A clearer and more correct picture is given if the circumpolar species are re-

garded as an element of the zone in which they live and their number is calculated

as a ratio of the total fauna of this zone. This is possible only for the Carabids

of Fennoscandia in northwestern Europe. Three species lacking there {Carabus

truncaticoUis, Harpalobrachys leiroides, Pterostichus vermiculosus) must be excluded.

In table 8, as well as in the corresponding table 30 by Lindroth, 1949 (p. 440-448),

crosses in brackets are counted.

The circumpolar Carabids of Fennoscandia constitute:

in regio arctica superior, 5 species of 5, that is 100 per cent

,, regio arctica media, 11 ,, ,, 17, ,, ,, 65 ,, ,,

,, regio arctica inferior, 31 ,, ,, 75, ,, ,, 41 ,, ,,

,, regio subarctica, 41 ,, ,, 97, ,, ,, 42 ,. ,,

,, regio coniferina, 37 ,, ,, 271, ,, ,, 14 ,, ,,

,, regio quercina, 23 ,, ,, 315, ,, ,, 7 >. ..

,, more sou

thern r

egions

14

„ 31

4, M ,,

4 ..

> »

The relationship between the palaearctic and nearctic faunas

289

(From Rosenberg, Faglar i Sverige.)

FIG. 49. Breeding area of the Snowy Owl, Nyctea scandiaca L., an arctic, circumpolar
bird. The southern limit varies considerably according to supply of rodents.

(Compiled from Ekman, 1922, Pleske, 1928, "Check List", 193 1, Stegmann, 1938, TiM-

MERMANN, 1 949, SaLOMONSEN & GiTZ, 195O.)

Thus, the circumpolar type dominates under high-arctic conditions and de-
creases relatively with decreasing latitude.

Examples of circumpolar distribution are here shown by maps of one arctic
species, the Snowy Owl {Nyctea scandiaca L., fig. 49), one subarctic-highboreal
species, the Three-toed Woodpecker [Picoides tridactylus L., fig. 50), and one
generally boreal species, the Dragonfly Libellula quadrimaculata L. (fig. 51). This
latter is an excellent flyer with strong migratory tendencies; it is able to cross even
broad stretches of sea.

Several reasons may be given which could explain why no faunal element has
been able to spread all around the globe to such a great extent as that of the High
North: —

(a) The way along a given parallel is shorter the nearer it lies to the Pole.

(b) The continents lie everywhere closer together about the latitude of the Polar
Circle than farther south. The most important distances are:

Bering Strait 90 km.
Labrador-Greenland 750 km.

Greenland-Iceland 330 km.
Iceland-Norway ca. 900 km.

19 — 565597 Lindroth

290

CHAPTER III

^

< /

x^

VV^

>

%

X

/\

^

~/w

^X^~7) /\ /C^""V''- \

/.' ^\/

/ "

S

>

so

"7^

"yC^^

^^'■^

7^

Nryi^ J^aA/ ft--'''''!^^\) ^XJ'^

1^

i_^

t/

/^^

^^^\^Xt^

»9^^J^'

/^ > ~"~~~^^ /"

/

^^^

K^/

P

trr^

KJf

i

M

jp*^^ —

r \ "y^^

J

$

Z/

ff

\'\fyF\^

Ci,(

^

p^^/~-

\

FIG. 50. The Three-toed Woodpecker {Picoides tridactylus L.).
Breeding range of a subarctic-highboreal, circumpolar bird,
boreoalpine in Eurasia.

(Mainly according to "Check List", 193 1, Stegmann, 1938, Peters,
1948, and HoLDHAUS, 1954.)

(From Rosenberg, Faglar i Sverige.)

Yet these favourable conditions do not in themselves provide an exhaustive
explanation, except for flying or otherwise easily dispersed forms. This was shown
above (p. 264) regarding the Davis Strait which, though only 350 km. at its nar-
rowest, has functioned as a highly efficient barrier against the dispersal of soil-
bound animals.

The relationship between the palaearctic and nearctic faunas

291

/^

^

^

j

7

yyyf

s

>0:

K

vO^

. /^

\^

a

>^

^#

/ ^ \

Y^k

^l\ y

yC^V

i^\

>^

w

^^

^

^^

/_ ^/7^

T^v

%\tt

vQC^

^^^

1 — r~"

7^"

2^>;

\

^^^

^^g^

[^

■^t'^^

:\^~~~ ~~~/

-^

\X^

3^^^^l\^

i?o/

^nX^V)

^:^^

-4^

1^^

^

ilk

^

l?d%

Pp^P-j^LX/

^^'pp

\/\^^^

'f^^^

t

\\)fe^

^^J^^l

^^^M

^S

iTx

1

\^S

^^^V^

WM

VA^P

^^

7 -l/-

I

v^

W^^

"^^s

^^

-^ 7

Vv

"^M

l-/^^"--.^ /)v^

^» ^/ ^

/

\ J^

/ /%/ -

/w\"^. t/\

^^/^

)y-^

\ r /y<

.-•■■ /'^s/

FIG. 51. Libellula qiiadrimaculata L., a generally
boreal, circumpolar Dragonfly with excellent means
of dispersal by active flight combined with wind-
drift. It is a well-known migratory insect.

(According to Bartenef, 1935, Valle, 1952, Needham
& Westfall, 1955.)

'-V,

(From Svensk Insektfauna.)

(c) The sea and the sounds between the continents of the North are regularly
closed by drifting ice, at least in the first half of the year, joining the Siberian coast
with Novaja Zemlja and via Spitzbergen with Greenland and, usually, northern
and eastern Iceland. On the American side, continuous drifting ice every spring
goes right down to Newfoundland and the Bering Strait is regularly closed by ice.

The ice of the Polar basin, stable or drifting, is often traversed in winter by

292

CHAPTER III

large warm-blooded animals. The Arctic Fox {Alopex lagopus L.) has been tracked
on the ice at about 85° north (Braestrup, 1941, p. 19), also half-way between Green-
land and Iceland (Saemundsson, 1939, p. 3). Proofs of still longer distances
covered are available for the Reindeer (Rangifer tarandus L.) since animals with
owner's mark on the ears have repeatedly been observed in Spitzbergen (Ingstad,
1948, p. 239 a.f.), which they must have reached travelling on the ice from Novaja
Zemlja, at least, a distance of about 770 km. With this in mind it seems astounding
that a special race of the Reindeer, sbsp. pearyi All., could establish itself on Elles-
mere Island (fig. 53), separated from Greenland only by narrow ice-covered sounds.
Perhaps this is due to a more complete isolation in earlier periods of the Pleisto-
cene, and stragglers may not be accepted in the herds.

Cold-blooded and, probably, small warm-blooded animals, such as the Lem-
mings, cannot traverse the sea-ice actively. They may do so by passive ice-transport
carried by currents, as has been suggested for the Waterbeetle Colymhetes dolabra-
tus Payk. (from Greenland to Iceland; Lindroth, 193 1, p. 529) and is still more
probable for the high-arctic circumpolar Staphylinid Micralymma brevilingue
Schie. {dicksoni Makl.), inhabiting the tidal zone. Yet a general application of
this kind of dispersal to the soil-bound fauna is made highly dubious by the existing
faunal barrier between Greenland and Baffin Island, existing in spite of the fact
that surface currents rather favour westward transport across the Davis Strait
(Orchymont & Brown, 1940, fig. i).

(d) Extension of land in the far north may have been more favourable during
earlier parts of the Pleistocene, or in Tertiary time. This, to my mind, is the clue
to the circumpolar type of distribution, or at least to the frequent occurrence of
animals so distributed.

In a previous paragraph of this chapter (p. 253 a.f.) I have tried to show that Ice-
land and Greenland are so firmly linked to Europe, from a zoogeographical point of
view, that the assumption of an uninterrupted land-connection (or almost so) with
the European mainland seems unavoidable. At the same time, however, it was
stressed that a westward prolongation of this "bridge", to Baffin Island, could
not be traced on faunal evidence. This will be still more evident if we consider the
pattern of subspeciation among circumpolar animals (below). It is a fact beyond
any doubt that the circumpolar area of very few animal species, if any, has
incorporated America from Eurasia, or vice versa, by way of Greenland-Baffin
Island.

This being admitted, the Bering Strait, or rather the passage now occupied by
this strait, is the historical link between the Palaearctic and Nearctic regions.

The idea is not at all revolutionary. It has been advocated by many skilled bio-
geographers (Matthew, 1915; Mayr, 1946; Simpson, 1947; &c.). And the conditions

The relationship between the palaearctic and nearctic faunas 293

for a repeated firm connection between Asia and North America by this route
are more favourable than between any second pair of continents on the earth.

The Bering Strait is only 90 km. at its narrowest and quite shallow, a land rise
of 60-65 meters would be enough to shape a continuous bridge. This is no high
figure. It lies within the limits of what is thought to be the normal world-wide
eustatic regression of the sea during a glaciation. Even for the last glaciation ( Wiirm,
Wisconsin) this regression is usually calculated as something between 90 and 100
meters, somewhat more during at least one of the earlier ones (Zeuner, 1950, p.
129; Woldstedt, 1954, p. 289-291.

For other reasons, too, it is very likely that the Bering land-bridge existed during
a period (or several periods) of glaciation. Alaska was little affected by the land-ice,
the major part of it remained ice-free throughout the entire Pleistocene period
(Flint, 1952) as did the opposite part of eastern Siberia (fig. 52). Therefore, no
isostatic sinking of land caused by pressure of an ice-cap could affect the Bering
area. The bridge furthermore prevented cold polar water from entering the North
Pacific, and the south-coast of the isthmus, as well as the chains of islands off the
coast, the present Aleutians, &c., were more under the influence of the warm Kuro
Shivo current.

On the other hand, the Bering Isthmus was a pronounced "filter bridge"
(Simpson, 1940, p. 148), certainly in the first place due to climate. It allowed
passage to arctic and subarctic organisms but, as we just learned, there are very
few circumpolar animals not able to endure at least subarctic conditions. This is
well in accordance with the assumption made, that the Bering bridge developed
and existed dnring glacial periods of the Pleistocene. Therefore very few organisms
demanding a temperate, or warmer, climate are common to North America and
Asia. For instance the Tiger-beetles, genus Cicindela, are very abundant in North
America, amounting to almost one hundred different species (north of Mexico),
but they do not enter the Arctic and none of them has reached countries outside
the American continent.

In Tertiary time conditions may have been different. It is generally accepted
that the Bering land-bridge existed throughout the main part of this period
(Simpson, 1947) and thus under comparatively favourable climatic conditions. It
remains doubtful, however, if they were warm enough to explain the distribution
of Pantropical genera or groups (Mayr, 1946, p. 36). But, as we have seen, the
actually living species which are supposed to have crossed the Bering bridge are
northern in distribution, have low demands on climate and are no doubt Pleisto-
cene migrants. Blethisa eschscholtzi Zoubk., among Carabid beetles (p. 280), may
be a single surviving representative of Tertiary dispersal.

The following additional attempts (e-g) to explain the wide east- west distribu-

294

CHAPTER III

^^,^^

:^5;i.

FIG. 52. Pleistocene glaciation of North America regardless of time.

(Reproduced by permission from "Textbook of Geology", Part i, by Longwell, Knopf, &

Flint, published by John Wiley & Sons.)

tion of arctic-subarctic species are mainly hypothetical but are mentioned as a
possible incentive for further research.

(e) Arctic animals are more "hardy" than others, especially as regards low
temperature. They may therefore be more liable to survive passive transport,
above all as "aerial plankton" at high altitude. There would be no difficulty in
testing this experimentally.

(f) Arctic (and subarctic) animals may generally be older as species and thus
have had more time at their disposal for dispersal. This may be a direct conse-
quence of climate (Van Dyke, 1939, p. 258), either because the rate of mutation
decreases with temperature, as shown in Drosophila (Timofeeff-Ressovsky, 1947,
p. 243), or because the number of generations per unit of time is smaller. The
latter suggestion is more than a hypothesis: it is a common experience among

The relationship between the palaearctic and nearctic faunas

295

^J

Species 42

DIAGR. 9. Percentage of macropterous (white), dimorphic (striated) and brachy-
pterous (black) species of Carabid beetles inhabiting Fennoscandia, arranged into dif-
ferent groups of distribution.

a = exclusively European
b = Euro-Caucasian & Euro-Medi-
terranean
(From Lindroth, 1949, p. 435.)

c — West-Palaearctic
d — generally Palaearctic
e = Circumpolar

entomologists that widespread insects in the northern part of their area or in high
mountain regions turn to biennial development.

(g) Speciation may also be delayed, as an indirect result of climate, by decreased
selection within the rather uniform arctic region with its generally low population
density (reduced "competition"). This assumption, however, at least as far as
insects are concerned, is purely theoretical.

A question worth raising is why some species reached the outermost limits of
the continents, Scandinavia and/or the British Isles in Eurasia, Labrador and/or
Newfoundland in America, and acquired a perfectly circumpolar area, and how
these were selected from the far more numerous stock of other arctic species. The
catch-word "aggressive" species, &c., gives no real explanation.

An investigation of the means of dispersal of every separate species may do so,
at least in part. Circumpolar species of Carabid beetles as a rule have fully devel-
oped, functionary hind-wings, as shown in an earlier book (Lindroth, 1949, p. 435)
on the Fennoscandian fauna. The diagram here reproduced (diagr. 9) is not quite
up to date with respect to later discoveries of several species in North America,
but it demonstrates the tendency.

296

CHAPTER III

TABLE 9. Circumpolar Carabidae, 45 species {vide table 8). Western limit in
Europe plotted against eastern limit in North America.

W. limit in Europe

40-60 E.

„
20-40 E.

W. of 20° E.

Brit. Isles

130-160 w.

3

I

2

species

80-130° w.

2

I

2

I species

E. of 80° W.

I

3

I species

Newfoundl.

3

9

16 species

If the extension of area is primarily dependant on powers of dispersal, it is to
be expected that a species occurring far east, in Newfoundland or Labrador, in
North America, is usually at the same time among those which have spread farthest
west, to the British Isles or Scandinavia in Eurasia. This is confirmed by the follow-
ing simple arrangement (table 9).

Though circumpolar Carabids are thus, generally speaking, characterized by
well developed powers of dispersal, they by no means constitute a homogenous
group in this respect. A division of the species of each vegetational zone according
to the development of wings, into macropterous (flying, or probably flying),
brachypteroiis (flightless), and dimorphic forms (as in table 8), gives the following
picture: —

Circumpolar Carabidae

species macropt. brachypt. dimorph.

total

Regie arctica superior

7

I = 14 %

5=71%

I

,, ,, media

13

5 = 38 %

6 = 46%

2

,, ,, inferior

34

20 = 59 %

9 = 26 %

5

,, subarctica

44

28 = 64 %

9 = 20 %

7

,, coniferina

38

28 = 74 %

5 = 13%

5

,, quercma

23

16 = 70%

2= 9%

5

More southern regions

14

10 = 71 %

I = 7 %

3

Total 45

29 = 64 % 9 = 20 %

The decreasing number of flightless circumpolar species in the direction north-
south is very striking. The explanation is probably that selection on the wind-

The relationship between the palaearctic and nearctic faunas 297

swept barren land of the arctic region works in favour of flightless forms and species
(the dominance of the constantly brachypterous subgenus Cryobius, of the Carabid
genus Pterostichus, on the Siberian and Alaskan tundras is a good illustration); but
also, that the Bering Isthmus was available as a trafficable bridge at any period of
existence for animals enduring an arctic climate, and consequently that flightless
arctic forms were at less disadvantage crossing it than the more fastidious forms
of the forest regions. It is interesting to observe in this connection that no species
of Conifer trees (provided Picea glauca Moench. is specificially distinct from
abies L. and obovata Ledeb.) occurs on both sides of the Bering Strait (Hustich,

1953)-

Many circumpolar animal species seem to be uniformous, from a taxonomical
point of view, within their entire vast area, though this may be largely due to
imperfect study of sufficient series from different localities. In others, two or more
geographically separated forms, usually termed "subspecies", have been described.
One instance of a multiformous complex is the Reindeer {Rangifer tarandus L.;
fig. 53), in which the relationship between the different subspecies seems rather
obscure.

The usual trend among circumpolar animals is that subspeciation, or whatever
it may be termed, has the character of a sliding change of certain morphological
properties which thus can be expressed as dines (in Huxley's sense) running from
east to west. A good instance is provided by the Moose or European Elk {Alces
alces L.; fig. 54), as investigated by Peterson (1952). Here, with respect to the
shape of the palate bones, a double cline is formed, running in both directions
from the E. Siberian— Alaskan area (diagr. 10).

The result is, in this case and in the character mentioned, that the forms at
the periphery of the species' area, in Europe and eastern North America, are converg-
ing. A similar instance is provided by Davenport (1941) who, in his monograph of
the butterfly genus Coenonympha, the "Ringlets" or "Heaths", analyses the distri-
bution of the circumpolar, extremely multiformous C. tullia O. F. Miill. (fig. 55).
He found two points of general interest: (a) that northwestern North America and
northeastern Siberia are inhabited by the same form (sbsp. mixturata Alph.); and
(b) that the Nearctic form occurring farthest east (sbsp. inornata Edw.) is strikingly
similar to the forms of western Europe, especially to the Fennoscandian form
(sbsp. suevica Hemm.). Davenport is probably right in his conclusion that both
of these, geographically speaking, peripheral subspecies are similar, not because of
direct historical connection, but because they have remained primitive and like
the original ancestor of the species, whereas other forms situated closer to the
centre and origin of its area (probably in eastern Asia), but near the climatic

298

CHAPTER III

H O

Q O

< I _

-

NORTHERN EURASIA

NORTHERN NORTH AMERICA

-45'

/

/

ALASKA

'^EASTERN
ASIA

—40

/

+

EUROPE AND
„ WESTERN ASIA

WESTERN / >r\.

CENTRAL N.

-

" \

-35

■°

EASTERN

+

-

1

pfiz

1
enmayeri

1 1 1 1

gigas shirasi andersoni amencana

DIAGR. 10. Geographical relationships of the subspecies of Moose or European Elk
(Alces alces L.) as indicated by the average relative shapes of the palates.
(From Peterson, 1952.)

periphery, were exposed to heavier selection pressure and changed. The result of
this procedure would be the same as through "elimination", in the sense of Reinig
(1938), but attained in a different way.

The small Carabid Bemhidion (Chrysobracteon) lapponicum Zett. behaves in a
similar way (Lindroth, 1954a, p. 129-130), the ''forma typica'' of northwestern
Europe and western Siberia being more like the form of the Mackenzie River
region {"bryanti Carr") than specimens from Alaska and eastern Siberia (sbsp.
latiusculum Mtsch.).

FIG. 53. Distribution of the Reindeer and Caribou (Rangifer tarandus L.) and its
numerous subspecies.

1. forma typica

2. sbsp. platyrhynchus Vrol.

3. sbsp. pearsoni I. yd.

4. sbsp. sibiricus Murr.

5. sbsp. valentinae Fler.

6. sbsp. angustirostris Fler.

7. sbsp. phylarchus Holl.

8. sbsp. arcticus Rich.

9. sbsp. daivsoni Seton

10. sbsp. niontanus Seton

11. sbsp. caribou Gmel.

12. sbsp. caboti Allen

13. sbsp. terranovae Bangs

14. sbsp. pearyi Allen

15. sbsp. groenlandicus Gmel.

(According to Ekman, 1922, Anderson, 1946, Ellerman & Morrison-Scott, 1951, Burt &
Grossenheider, 1952, and Flerow, 1952.)

The relationship between the palaearctic and nearctic faunas

299

/\.

>r \ \___^''' /Vi _-/^ '^^^^C '^~-

jfy^

'//

\\(\jQ4J^^l^^-

S-

^^:::]

Tk^W

j

t

^P

—A

B

"^ ^

^^r^^^i]

\^}J^Wt>-'

^ ,..'

(From Burt & Grossenheider, 1952.)

300

CHAPTER III

Mi

1 Alccb alecs alces

2 Alces alc£s camelotdes
5 Alces dices pfitenmaven
4- Alccj. dices ^iq as

5 Alces alces shirasi

6 Alces alces andersoni

7 Alces alecs amei-icana.

FIG. 54. World distribution of the Moose or
European Elk (Alces alces L.) and its sub-
species.

(From Peterson, 1952.)

(From Burt & Grossenheider, 1952).

The relationship between the palaearctic and nearctic faunas

301

/

"y^cw\}3^

<^

&

/ /'

'Mm

'■ / /

wM

t

B

^

v^^^^^m^

\

}^3k^S^

FIG. 55. The distribution of Coenonympha tullia O. F.
Miill. {tiphon Rott.) (The Ringlet or Large Heath),
an extremely multiformous butterfly.

The figured butterfly is of forma typica from S.
Sweden.

(According to Davenport, 1941, Ford, 1946, &c.)

(Photo P. Ardo.)

I.

sbsp. scotica Stgr.

15

2.

sbsp. philoxenus Esp. (davus

16

auctt.)

17

3-

sbsp. suevica Hemm. (incl.

18

orstadii Wahlgr.)

19

4-

forma typica

20

5-

sbsp. italica Vty.

21

6.

sbsp. bosniae Davenp.

22

7-

sbsp. occupata Reb.

23

8.

sbsp. rhodopensis Ehv.

24

9.

sbsp. chatiparae Shelj.

25

10.

sbsp. caeca Stgr.

26

II.

sbsp. eupompus Stauder

27

12.

sbsp. subcaeca Heyne

13-

sbsp. zvitimensis Davenp.

28

14-

sbsp. sibirica Davenp.

sbsp. viluiensis Men.

sbsp. mixturata Alph.

sbsp. kodiak Edw.

sbsp. tnackenziei Davenp.

sbsp. Columbiana McD.

sbsp. ampelos Edw.

sbsp. eryngii H. Edw.

sbsp. calif ornica Westw.

sbsp. furcae Brns. & Benj.

sbsp. subfusca Brns. & Benj.

sbsp. ochracea Edw.

sbsp. benjamini McD.

sbsp. inornata Edw. (incl.
mcisaaci dos P.)

sbsp. nipisiquit McD.

302

CHAPTER III

(From Faune de France.)

FIG. 56. Distribution of two species of the Carabid genus Blethisa.

Hatched area = Blethisa multipiinctata L., in Europe and western Siberia /or?«a
typica, in North America sbsp. aurata Fisch.; overlapping or transgrading area in
eastern Siberia.

Black dots = Blethisa eschscholtzi Zoubk.

The figured specimen belongs to multipunctata f. typ.

However, these are exceptions. The rule is that, provided a circumpolar species
is at all inclined to subspeciation, the most differing forms appear in western or
northern Europe on the one hand, in North America east of Hudson Bay on the
other. This applies to the following Carabid beetles: Agonum mannerheimi Dej.,
Amara alpina Payk., Blethisa multipunctata L. (fig. 56), Miscodera arctica Payk.,
Nebria gyllenhali Schnh., A'^. nivalis Payk., and Pelophila borealis Payk.

Unfortunately, the lack of sufficient series of these species from northern Asia
does not permit a description of existing cline systems, but the gradients as a rule
seem not to be evenly sloping but to form a distinct step within a transitional zone
on either side of the Bering Strait: in eastern Siberia at least for Amara alpina,
Blethisa multipunctata (fig. 56), Nebria gyllenhali and nivalis; in Alaska or adjacent
parts of Canada for Miscodera arctica and Pelophila borealis. Whether this is due
to overlapping after previous isolation or to the transitional zone indicating the
original centre of the species, I am not able to decide. But the presence, at least in
other animal groups, of a common subspecies (Coenonympha tullia, fig. 55; also
the sbsp. leptura Hbs. & Schtz. of the Burbot, Lota lota L.), or of two closely
related subspecies (Moose, diagr. 10), on both sides of the Strait, from which other

The relationship between the palaearctic and nearctic faunas 303

FIG. 57. Bembidion trans-
parens Gebl. Scandinavian
specimens.

a = macropterous form (fly-
ing)

b = intermediate form (flight-
less) (very rare)

c = brachypterous form
(flightless).

Short wing is a dominant
and long-winged specimens
are homozygotes.

subspecies radiate toward Europe and eastern North America, argues in favour
of the "overlapping area" as an old centre.

When pronounced and geographically concentrated, this area is situated west
of the Bering Strait, which supports the generally accepted idea that the main
Holarctic centre of speciation was in northeastern Asia and that the traffic over
the Bering land-bridge, irrespective of period, was most intense from the west
to the east (Simpson, 1947, p. 628 a.f.).

Yet it is important to realise that a circumpolar area is not the "climax" stage
of a perpetual and even dispersal from an ancient evolutionary centre of the species.
It is but one stage in a complex procedure involving intermittent expansion and
regression. Species may once have been circumpolar which now no longer are so,
for instance the Muskox {Ovibos moschatus L.). The repeated glaciations prevented
any species from being continuously circumpolar during the entire Pleistocene
period, but at the same time they caused pulsating advances toward the south which
in many cases, left permanent outposts in southern mountains. This gave rise to
boreoalpine types of distribution (Holdhaus & Lindroth, 1939; Holdhaus, 1954), for
instance in the Three-toed Woodpecker {Picoides tridactylus L.; fig. 50). It is
difficult to establish the period in which the area of a boreoalpine species was
formed, but it is reasonable to assume that the time of the largest ice-sheet, which,

304

CHAPTER III

FIG. 58. Bembidion transparens Gebl. Distribution of long-winged (white) and short-
winged (black) specimens in North America. The size of the circles is in proportion
to the number of individuals examined. The circle for Newfoundland is strongly
underdimensioned {vide fig. 59).

in Europe at least, was the last but one glaciation (Riss), had the greatest effect
(Lindroth, 1935, p. 627).

The boreoalpine type of distribution is most frequent in Europe because a wide,
continuous plain separates the southern mountains from the arctic-subarctic
regions of the north. But it occurs also in Asia, as shown by the Three-toed Wood-
pecker (fig. 50) and the Carabid Diachila polita Fald. (Lindroth, 1954 c, p. 9),
though information as to its extent is not obtainable because of insufficient know-

The relationship between the palaearctic and nearctic faunas

305

FIG. 59. Bembidion trans-
parens Gebl. Distribution of
long- and short-winged speci-
mens in Newfoundland. For
explanation, vide figs. 57, 58.

ledge. The Cordilleras of western North America are against boreoalpine distribu-
tion but in the northeast the St. Lawrence Valley cuts off from the main area an
arctic-subarctic fauna on the highest mountains of New England, especially in
the White Mountains of New Hampshire (Darlington, 1943). In some Diurnal
Lepidoptera (Butterflies) the southern component of a boreoalpine species is
supplemented or replaced by one or more subspecies ecologically connected with
mires and bogs (Petersen, 1954).

The distribution of wing-dimorphic insects, here as in many other cases, may
allow partial reconstruction of the troublesome history of circumpolar species.

The tiny Carabid beetle Bembidion transparens Gebl. [sulcatum Lee.) (fig. 57)
apparently has an uninterrupted transamerican distribution and a traditional map
of its area does not preclude the interpretation of a successive spread in a west-
east direction across North America in Postglacial time. The distribution of
long- and short-winged individuals (fig. 58) gives a contrary impression. From
the western half of North America, west of longitude 100°, only macropterous
specimens are known; the westernmost brachypterous specimen seen is from
Aweme, Manitoba. A predominance of this form occurs only in Nova Scotia and
Newfoundland and this region should therefore be regarded as a secondary centre
of distribution. The picture may become slightly, but not decisively, altered if

20 — 565597 Lindroth

3o6

CHAPTER III

more abundant material available reveals the presence of brachypterous individuals
in Alaska.

The following reconstruction of the history of Bembidion transparens in North
America is the most plausible:— Immigration from northeastern Asia in early
Pleistocene and subsequent spread to the Atlantic coast. Interruption of area
during one or more glacial periods in the western and central parts. Survival, at
least throughout the last glaciation (Wisconsin), in the Northeast. Postglacial re-
colonization of lost area from the east, possibly also from the west.

The distribution in Newfoundland (fig. 59) suggests glacial refuges on the coast,
in the northwestern, northeastern and southeastern parts. —A similar coastal refuge
during the last glaciation was assumed for this species in northeastern Fennoscandia
(Lindroth, 1949, p. 389 a.f., fig. 45).

The fossil evidence

The importance of reliably dated fossils to the reconstruction of faunal and floral
history is immense. Our knowledge of what actually happened, and thus the
interpretation of the present pattern of distribution, has often been radically
altered by the discovery of a single fossil specimen.

The difficulties lie in a competent identification of the often fragmentary re-
mains, but also in the scope of conclusions. Above all, considering what a futile
fraction of organisms existing during a certain geological period was by chance
fossilified, and how still more futile a fraction of them was discovered and came to
the notice of a specialist, it is extremely dangerous to conclude anything from
negative facts, from the seeming lack of a certain type of animal, in a certain area,
from a certain period.

Yet fossils may show that many gaps in the present area of an animal group or
species are secondary and that faunal connections once existed which are impossible
to reconstruct from the recent distribution.

This applies very much to the question of faunal exchange between Europe
and North America. The fossil faunas of both of these continents are the best
known in the world and, partly— at least — for this reason, many fossil forms were
regarded as Eur-American or a group of animals or plants now restricted to either
continent was discovered as fossil in the other.

The resemblance of these faunas was apparently greater in remote times and
this, as described above (p. 286), originally led Simpson to conclude, mainly from
fossil Mammalia, that a broad transatlantic connection existed in the Lower
Eocene; later he found this assumption unnecessary. It is worth mentioning in
this connection that the insect fauna described by Henriksen (1922) from Lower

The relationship between the palaearctic and nearctic faunas

307

(From Traite de Zool.)

(From Ander, 1942.)

FIG. 60. Distribution of the beetle family Cupedidae, recent and (in Europe) in Baltic
amber. The figured specimen is of Cupes rajjrayi Fairm. from Madagascar.

Eocene Diatom-earth in Denmark has its closest geographical connection with
Southern Asia.

Now, the distribution of existing species, and usually even of genera, is little
affected by what happened in the early Tertiary. The first period in which possibly
still existing species had developed within the terrestrial fauna was the time of the
Baltic amber, usually referred to Upper Eocene (or Lower Oligocene), that is
more than 40 million years ago. It happens that one of the most famous insect
specimens of the amber, a Tiger-beetle of genus Tetracha (Megacephala), has a
special bearing upon our problem. It is often regarded as the single instance of an
insect species still living unchanged since that time (Ander, 1942, p. 26)^, since W.
Horn (1906), though with some doubt, declared it as identical with the American
T. Carolina L., distributed from Virginia through Central America to northern
Chile. The genus has a wider occurrence (S Spain, Africa, Hither Asia, Australia)
but the group or subgenus to which Carolina belongs is strictly American and Walter
Horn was so skilled a specialist in Tiger-beetles that he could not possibly have
identified the wrong group. It is therefore easily understandable that some students
(at least verbally!) expressed their doubts as to the authenticity of the specimen, ^ —
On other proposed North American elements from Baltic amber, vide Kolbe

(1925)-

The main impression of the Baltic amber fauna, if climatic differences from pre-
sent European conditions are disregarded, is its importance as a connecting link

^ However also a small Silphid beetle, Nemadus colonoides Kr., is reported to have
remained unchanged since the time of the Baltic amber (Jeannel, 1942, p. 192).

^ The specimen belonged to the Berendt collection of the Geol.-Palaeont. Inst., Humboldt
University, Berlin, but seems to have been lost.

3o8 CHAPTER III

between highly disjunct present areas of old taxonomical groups. The primitive
beetle family Cupedidae (fig. 60) is a good example.

As -more and more recent species appear from the course of the Tertiary, the
geographical character of the different faunas and floras becomes more easy to
fix; and elements, often identified to species, of an indisputable "American"
type— judged from present distribution— were discovered in Europe.

The Tertiary development of the European fauna of terrestrial and limnic
Molluscs is notably well known and has been summarized by Ehrmann (1914).
From the Eocene no recent species is known; the connections, according to present
distribution of genera or groups, were largely towards the Tropics, including,
but not favouring. South and Central America. Of particular interest is the family
Oleacinidae (p. 283), now restricted to tropical America, subtropical North America
and the Mediterranean, but found as fossil in Europe as early as in Upper Creta-
ceous.

In the OUgocene more decidedly American types appear, among these genus
Strobilops {Strobilus auctt.), now chiefly American but also occurring in East
Asia (fig. 61). It persisted in Europe at least to Middle Pliocene. Pilsbry (1948,
p. 853) assumes an Asiatic centre of origin for Strobilops, with subsequent radia-
tion, west into Europe, east across the Bering bridge into America.

Other Oligocene Molluscs also show relationship with America, for instance
several freshwater Bivalves. Modell (1943) therefore postulates a direct Trans-
atlantic land-connection in this period, which was probably of "Wegenerian" type.
Yet the so-called American forms of Oligocene Molluscs in Europe are constantly
associated with other extra-European elements, mainly with recent occurrence in
Asia.

In the Miocene and Pliocene periods genera and species groups suggesting Ameri-
can and South or East Asian connections gradually and contemporaneously
disappear. The Upper Pliocene contains an almost entirely European fauna and
no "American" Molluscs; nor are any known from the Pleistocene.

A most interesting light has quite recently been thrown upon the late Tertiary
and early Pleistocene history of the European flora through the intense investiga-
tions in Polonia by Szafer (1954). In Lower Pliocene the American element was
twice— but the East Asian four times (!)— as numerous in species as the generally
Eurasian group. The balance changed gradually in favour of Eurasian plants but
not until the first Pleistocene glaciation (Giinz) did this element take a clearly
dominating position. In the first part of the following Interglacial period the East
Asian group was well represented (11 species) but it disappeared completely
during the course of this period, whereas two species {Tsuga sp. and Osmunda

The relationship between the palaearctic and nearctic faunas

309

FIG. 61. Recent and (in Europe) Tertiary distribution of the
Mollusc genus Strobilops (Strobilus auctt.) in the Northern
Hemisphere. (Outside the map found in South America only.)

1. Strobilops s. sir. (America) 3. sxxhg. Discostrobilops {America)

2. subg. Eostrobilops (E.Asia) 4. subg. Enteroplex (Philipp.)

The figured specimen is of the North American Strobilops
(5. str.) labyrinthica Say.

(After PiLSBRY, 1948.)

claytoniana L.^) of the American group survived the whole Giinz-Mindel Inter-
glacial.

From investigations in other parts of Europe the persistence of some of the
American plants can be followed into later parts of the Pleistocene. Three species
of the sedge genus Dulichium, now restricted to America, are recorded from Upper
Pliocene (Vlerk & Florschiitz, 1953; Szafer, 1954), two of these survived the
first glaciation and one, D. spathaceum Rich., still grew in Denmark during the
last (Riss-Wiirm) Interglacial (Jessen & Milthers, 1928, p. 348). The water-fern
Azolla filiculoides Lam. likewise persisted into Pleistocene time. Similarly, the
fern Osmunda claytoniana, just mentioned, or some closely related non-European
form, has quite recently (Lundqvist, 1955, p. 321) been discovered in Swedish
deposists from the last interglacial period,

A very interesting discovery was that reported from "glacial" deposits (without
exact dating)^ at Deuben, Saxonia, (Nathorst, 1894), of an elytron of Carabus

^ Osmunda claytoniana is not strictly American. It still grows in Himalaya and East Asia.
2 Woldstedt (1954, p. 248) dates the Deuben fossils as early Wiirm or, possibly, Riss.

310

CHAPTER III

chamissonis Fisch. {groenlandicus Dej.), a beetle now restricted to arctic-subarctic
North America (the record for Greenland being erroneous). Since the elytral
sculpture is very characteristic in this species, a wrong identification seems out
of the question.— For a review of the late Tertiary and Pleistocene fossil insects of
Europe, vide Henriksen (1933, p. 265 a.f.).

The key to a proper understanding of the now extinct American element, not
only of the European flora but also of the contemporaneous/awna, seems to be given
by the balance between American and East Asian plants in the different periods
investigated by Szafer (diagr. 11). Until, and inclusive of, the first part of the
Giinz-Mindel Interglacial the curves for these tzvo elements run perfectly parallel,
the East Asian element being clearly in the majority. This is definitely not what
could be expected if the presence of the American plants was due to a Transatlantic
land-connection. In that case, the American element would have increased at
the cost of the East Asian. The parallel development of the said elements can only
be explained under the assumption that the "American'' plants were simply part of
the East Asian group, that they invaded Europe together from the east.

The longer persistence in Europe of some few members of the American type
{Duiichium, Osmunda, Sec.) does not contradict this opinion, because it must be
expected that these plants, being able to reach both Europe and eastern North
America from a centre in Asia, belonged to the most hardy and most easily dis-
persed plants of the group which, in full, ought to be called "East Asian". The
only obscure point is why they later, entirely or in part, became extinct in Asia.

The opinion here stressed, that northeastern Asia has served as an evolution
centre for circumpolar animals, is quite in accordance with Hulten's view (1937)
of the development of the Holarctic flora. But to my mind, Huhen has underesti-
mated, (a) the possibility of transatlantic dispersal without the aid of land-connec-
tions, and (b) the importance of an element of original (endemic) species in Europe
and North America. Then it is evident that the mountains of both regions possess
a rich element of indigenous species though, for some reason, these have almost
failed to spread outside their area of origin.

The reasons may be tentatively indicated as follows: —The main areas of specia-
tion were mountains. They possess a great variety of habitats and the strong short-
way zonation shapes small isolated population areas, thereby favouring speciation.
The Tertiary mountain ranges, the Alps, the North American Cordilleras, &c.,
are young. They have existed long enough to allow for speciation but not for the
development of really new "inventions", higher taxonomic units, essentially new

The relationship between the palaearctic and nearctic faunas

3"

DiAGR. II. Three main elements of the Pliocene-Lower Pleistocene flora of SW
Polonia.

Black dots = the American element. Open dots = the East Asia element.

Crosses = the Eurasian element
Elements of a generally Holarctic or more southern character omitted.
(After Szafer, 1954.)

types of organisation. And the species arisen in these mountains had comparatively
too short time to spread.

Older mountains, of Palaeozoic (Carboniferous-Permian) age, the Hercynians
of Europe, the Appalachians, &c., have been more important centres of evolution.
Apparently it was the fauna of such mountains, in northeastern Asia, that provided
the main source of circumpolar animals. Considering that the Bering Strait during
the entire Tertiary and Quaternary periods has formed only a temporary obstacle
to dispersal, the fauna (and flora) of northeastern Asia was in the best imaginable
position, in the centre of the Holarctic region, for sending branches in both direc-
tions, towards Europe and towards North America.

But this is not sufficient explanation. Why, for instance, did the Appalachians

312 CHAPTER III

not provide a similar old faunal element, spreading west across the Bering land
bridge?

The most acceptable hypothesis seems to be that the old mountains of north-
eastern Asia, simply because they were situated farther north, the Hercynian ranges
reaching at least 60° north in the Jablonoj Mountains, and probably farther
northeast (Obrutschew, 1926, p. 445), and because they probably always had a
more continental climate, gave rise to a Tertiary fauna with a high amount of
hardy species^ which, more than any other contemporaneous fauna, were fit for
subsequent dispersal across two continents, and later to surv'^ive the critical glacial
periods. Thus they became circumpolar.

The original faunas of Europe and North America— and it should be realized
that these regions were certainly never "empty" — were less adapted to arctic-
subarctic conditions, were perhaps largely inhabitants of mountain woods, and
therefore had no chance of becoming members of a cold-adapted Holarctic fauna.
It is possible, though, that the Ural Mountains once served as a centre similar to
that of northeastern Asia but, if so, the following Pleistocene glaciations swept
away the traces of their original fauna.

^ Kusnezov (1935, p. 130) tries to explain this phenomenon by proposing a Miocene
glaciation of Alaska and NE Siberia.

Bibliography of Chapter III

Alexander, W. B. & Fitter, R. S. R., 1955. American land birds in western Europe.

— Brit. Birds. 48. Watford, Herts., p. 1-14.
Ander, K., 1942. Die Insektenfauna des baltischen Bernsteins nebst damit verkniipf-

ten zoogeographischen Problemen. — K. Fysiogr. Sallsk. Handl. (N.S.) 53:4.

Lund, p. 1-83.
Anderson, R. M., 1946. Catalogue of Canadian recent Mammals. — Bull. Nation.

Mus. Can. 102. Ottawa, pp. i-v, 1-238.
Arndt, W., 1928. Der Susswasserschwamm Heteromeyenia ryderi Potts auf den

Far Oern. Zugleich ein tjberblick iiber die Spongilliden-Fauna der Inseln des

Atlantik und seiner Neben- und Randmeere. — Zool. Anz. 77. Leipzig, p. 1 56-166.
Atlantisheft, 1939. — Geol. Rundschau. 30. Stuttgart, p. 1-400.
Atlas til Bind I og II, 1921. Gronland i 200-aaret for Hans Egedes Landing. —

Copenhagen.
Bartenef, a., 1935. iJber die Gattung Libellula und besonders iiber ihre pala-

arktischen Arten. — Zeitschr. wissensch. Zool. (Arch. f. Naturgesch.) (N.S.) 4.

Leipzig, p. 274-290.
Basilewsky, p., 1954. Description d'un Carabique nouveau du Ruanda-Urundi,

representant d'une sous-famille inedite pour TAfrique Noire. — Ann. Mus.

Congo. 4. Zool., I. Tervuren, p. 301-303.
Bey-Bienko, G. J., 1950. Blattodea. — Fauna SSSR (N.S.) 40. (Russian.) Moscow &

Leningrad, p. 1-343-
Blair, K. G., 1933. Coleoptera collected by the Oxford University Expedition to

Akpatok Island, Ungava Bay, Aug. -Sept. 1931. — Ann. Mag. Nat. Hist. (10)

12. London, p. 93-96.
Blasius, W., 1903. Der Riesenalk, Alca impennis L. — Naumann: Naturgesch. Vogel

Mitteleuropas. 12. Gera-Unte'rmhaus, p. 169-208.
Bocher, T. W., 1938. Biological distributional types in the flora of Greenland. —

Medd. om Gronl. 106:2. Copenhagen, p. 1-339-
— 1948. Contributions to the flora and plant geography of West Greenland.

I. Selaginella rupestris and Sisyrinchium montanum. — Ibidem. 147:3, p.

1-26.
Boettger, C. R., 1950. Analyse einer bemerkenswerter Population der Schnirkel-

schnecke Cepaea hortensis Miiller. — Abh. Braunschw. Wiss. Gesellsch. 2. Braun-
schweig, p. 1-12.
Boycott, A. E., 1936. The habitats of fresh-water Mollusca in Britain. — Journ.

Anim. Ecol. 5. London, p. 116-186.
Braendegaard, J., 1946. The spiders (Araneina) of East Greenland. A faunistic and

zoogeographical investigation. — Medd. om Gronl. 121:15. Copenhagen, p.
1-128.
Br^estrup, F. W., 1941. A study on the Arctic Fox in Greenland. — Ibidem. 131,

p. i-ioi.

314

CHAPTER III

Brehm, v., 1926. Amerikanische Typen in der Organismenwelt Europas. — Die

Erde. 4. Braunschweig, p. 315-325-
Brinck, p., 1940. Beitrag zur Kenntnis der Wasserkafer Gronlands. — Opusc. Ent.

5. Lund, p. 37-41-
Brooks, S. T. & B. W., 1940. Geographical distribution of the recent MoUusca of

Newfoundland. — Ann. Cam. Mus. 28. Pittsburgh, p. 53-75-
Brown, W. J., 1937. The Coleoptera of Canada's Eastern Arctic. — Can. Ent. 69.

Guelph, p. 1 06-1 1 1.
Brundin, L., 1943. Zur Kenntnis einiger in die Atheta-Untergattung Metaxya M. &

R. gestellten Arten (Col. Staphylinidae). — K. Fysiogr. Sallsk. Handl. (N.S.)

54:4. Lund, p. 1-38.
Burt, W. H. & Grossenheider, R. P., 1952. A field guide to the Mammals. —

Boston, p. 1-200.
Casey, T. L., 1900. Review of the North American Corylophidae, Cryptophagidae,

Tritomidae and Dermestidae, with other studies. — Journ. N. Y. Ent. Soc. 8.

New York, p. 51-172.
Check-List of North American Birds, 1931. 4. ed. — Amer. Ornith. Union.

Lancaster, Pa., p. 1-525.
Clapham, a. R., Tutin, T. G. & Warburg, E. P., 1952. Flora of the British Isles.

— Cambridge, pp. i-Li, 1-1591.
Creighton, W. S., 1950. The ants of North America. — Bull. Mus. Comp. Zool. 104.

Cambridge, Mass., p. 1-585.
Crosby, C. R. & Bishop, S. C, 1936. Aeronautic spiders with a description of a new

species. — Journ. N. Y. Ent. Soc. 44. New York, p. 43-49-
Dall, W. H., 1910. Land and fresh water Mollusks. — Harriman Alaska Exp. 13.

Smiths. Inst. Washington, D. C, pp. l-vn, 1-171.
Darlington, P. J., Jr., 1934. New West Indian Carabidae, with a list of the Cuban

species. — Psyche. 41. Cambridge, Mass., p. 66-131.

— 1935. West Indian Carabidae II.: Itinerary of 1934; forests of Haiti; new species;
and a new key to Colpodes. — Ibidem. 42, p. 167-215.

— 1938. Was there an Archatlantis? — Amer. Naturalist. 72. New York, p. 521-533.

— 1943- Carabidae of mountains and islands: data on the evolution of isolated
faunas, and on atrophy of wings. — Ecol. Monogr. 13. Durham. N. C, p. 37-61.

— 1948. The geographical distribution of cold-blooded vertebrates. — Quart. Journ.
Biol. 23. Baltimore, pp. 1-26, 105-123.

— 1949. Beetles and continents. (A review of La Genese des Faunes Terrestres by
R. Jeannel.) — Ibidem. 24, p. 342-345.

Davenport, D., 1941. The butterflies of the Satyrid genus Coenonympha. — Bull.

Mus. Comp. Zool. 87:4. Cambridge, Mass., p. 215-349.
Deevey, E. S., Jr., 1949. Biogeography of the Pleistocene. I. — Bull. Geol. Soc. Amer.

60. New York, p. 1315-1416.
Degerbol, M., 1940. Mammalia. — Zool. of the Faroes. 111:2. Copenhagen, p.

I-I33-
Ehrmann, P., 19 14. Grundziige einer Entvvicklungsgeschichte der Tierwelt Deutsch-

lands. (Lit.-Gesellsch. Neue Bahnen.) — Leipzig, pp. l-viii, 1-213.
Ekman, S., 1922. Djurvarldens utbredningshistoria pa skandinaviska halvon. —

Stockholm, pp. i-xvii, 1-614.

Bibliography 315

Ellerman, J. R. & Morrison-Scott, T. C. S., 1951. Checklist of Palaearctic and

Indian Mammals. — (Brit. Mus.) London, p. 1-810.
Emden, F. van, 1936. Bemerkungen zur Klassifikation der Carabidae: Carabini und

Harpalinae piliferae. — Ent. Blatter. 32. Krefeld, p. 12-52.
Fernald, M. L., 1925. Persistence of plants in unglaciated areas of Boreal America.

— Mem. Amer. Ac. Arts & Sci. 15:3. Lancaster, Pa., p. 237-342.

Fisher, J., 1952. The Fulmar. — (The New Naturalist.) London, pp. i-xv, 1-496.

Fisher, J. & Vevers, H. G., 1943. The breeding distribution, history and population
of the North Atlantic Gannet (Sula bassana). — Journ. Anim. Ecol. 12. Cam-
bridge, p. 173-213.

Flerow, C. C, 1952. Mammalia. L2. — Fauna SSSR. 55. Moscow & Leningrad, p.
1-256.

Flint, R. F., 1952. The Ice Age in the North American Arctic. — Artie. 5. Ottawa,
p. 135-152.

Ford, E. B., 1946. Butterflies. — The New Naturalist. London, pp. i-xiv, 1-368.

Fries, T. C. E., 1913. Botanische Untersuchungen im nordlichsten Schweden, &c.

— Uppsala, p. 1-36 1.

Gibson, A., 1920. Lepidoptera. — Rep. Can. Arct. Exp. 1913-18. Ill: i. Ottawa, p.

i-S8.
GiSLEN, T., 1940. The number of animal species in Sweden with remarks on some

rules of distribution especially of the microfauna. — K. Fysiogr. Sallsk. Handl.

(N.S.) 51:2. Lund, p. 1-23.
Glick, p. a., 1939. The distribution of insects, spiders and mites in the air. — U.S.

Dept. Agric, Techn. Bull. 673. Washington, D. C, p. 1-150.
Haanshus, K., 1933. Fortegnelse over Norges Lepidoptera. — Norsk Ent. Tidsskr.

3. Oslo, p. 165-216.
Hammer, Marie, 1953. Investigations on the microfauna of northern Canada. II.

CoUembola. — Acta Arctica. 6. Copenhagen, p. 1-108.
Hansen, A. M., 1929. Bre og biota. — Norsk. Vid. Akad. Skr., Mat.-Naturv. Kl. 5.

Oslo, p. 1-255.
Henriksen, K. L., 1922. Eocene insects from Denmark. — Danm. Geol. Unders.

(II) 37. Copenhagen, p. 1-36.-

— 1933. Undersogelse over Danmark-Skanes kvartaere Insektfauna. — Vid. Medd.
Dansk Naturh. Foren. 96. Copenhagen, p. 77-355.

— 1939. A revised index of the insects of Greenland. — Medd. om Gronl. 1 19: 10.
Copenhagen, p. 1-112.

Henriksen, K. L. & Lundbeck, W., 1917. Gronlands Landarthropoder (Insecta et
Arachnida Groenlandicae). — Ibidem. 22, p. 483-823.

Herre, W., 1935. Die Schwanzlurche der mitteleocanen (oberlutetischen) Braun-
kohle des Geiseltales, &c. — • Zoologica. 87. Stuttgart, pp. i-vi, 1-85.

Heslop-Harrison, J., 1953. The North American and Lusitanian elements in the
flora of the British Isles. — • In: Lousley, J. E. (editor), The changing flora of
Britain. — Arbroath, p. 105-123.

Holdhaus, K., 1927-28. Die geographische Verbreitung der Insekten. — In: Schro-
der, Handb. d. Ent. 11:6. — Jena, p. 592-1058.

— 1954. Die Spuren der Eiszeit in der Tierwelt Europas. — Abh. Zool.-bot.
Gesellsch. 18. Wien, p. 1-493.

3l6 CHAPTER III

HoLDHAUS, K. & LiNDROTH, C. H., 1 939- Die europaischen Koleopteren mit boreo-

alpiner Verbrcitung. — Ann. Naturh. Mus. 50. Wien, p. 123-293.
Horn, W., 1906. Ober das Vorkommen von Tetracha Carolina L. im preussischen

Bernstein und die Phylogenie der Cicindela-Arten. — Deutsch. Ent. Zeitschr.

Berlin, p. 329-336.
HuBENDiCK, B., 1947. Die Verbreitungsverhaltnisse der limnischen Gastropoden in

Siidschweden. — Zool. Bidr. 24. Uppsala, p. 419-559.
HuLTEN, E., 1937. Outline of the history of Arctic and Boreal Biota during the

Quaternary Period. — Stockholm, p. 1-168.

— 1950. Atlas of the distribution of vascular plants in NW. Europe. — Stockholm,
pp. i*-ii9*, 1-5 12.

HusTiCH, I., 1953. The boreal limits of conifers. — Arctic. 6. Montreal, p. 149-162.
Hynes, H. B. N., 1954. Identity of Gammarus tigrinus Sexton 1939. — Nature. 174.

St. Albans, p. 563.
Ihering, H. von, 1927. Die Geschichte des Atlantischen Ozeans. — Jena, pp. i-ix,

1-237-
Ingstad, H., 1948. Landet med de kalde kyster. (Spitzbergen.) — Oslo, p. 1-422.
Jaschnow, W. a., 1925. Crustacea von Nowaja Zemlja. — Ber. Wissensch. Meeres-

inst. 12. Moscow, p. 50-76.
Jeannel, R., 1937. Les Bembidiides endoges. — Revue Frang. d'Ent. 3. Paris, p.

241-399-

— 1 941. Coleopteres Carabiques. I. — Faune de France. 39. Paris, p. 1-571.

— 1942. La genese des faunes terrestres. — (Bibl. Inst. Marit. & Colon.) Paris,
pp. i-viii, 1-513.

Jensen, A. S., 1928. Gronlands Fauna. Et Forsog paa en Oversigt. — Festskr. Kbhvns

Univ. Copenhagen, p. 1-88.
Jessen, K., 1949. Studies in late Quaternary deposits and flora-history of Ireland.

— Proc. R. Irish Ac. (B) 52. Dublin, p. 85-290.
Jessen, K. & Milthers, V., 1928. Stratigraphical and palaeontological studies of

interglacial fresh-water deposits in Jutland and northwest Germany. — Danm.

Geol. Unders. (II) 48. Copenhagen, p. 1-380, Atlas.
JOHANSEN, H., 1955. Die Jenissei-Faunenscheide. — Zool. Jahrb., Abt. Syst., Okol.,

Geogr. 83. Jena, p. 237-247.
Johnson, C. W., 1906. On the distribution of Helix hortensis Mueller, in North

America. — Nautilus. 20. Philadelphia, p. 73-80.

— 191 5. Helix hortensis from a Maine shell heap. — Ibidem. 28, p. 131.
Kastner, a., 1932. Palpigradi Thorell. — In: Kiikenthal, Handb. d. Zool. 111:2, i.

Berlin, p. 76-98.
Kevan, D. C. McE., 1943. Study of an introduced North American freshwater

Mollusc, Stagnicola catascopium (Say). — Proc. R. Soc. B. 61. Edinburgh,

p. 430-461.
Kloet, G. S. & HiNCKS, W. D., 1945. A check list of British insects. — Arbroath,

pp. i-Lix, 1-483.
Kolbe, H., 1925. Vergleichender Blick auf die rezente und fossile Insektenwelt

Mitteleuropas, &c. — Deutsch. Ent. Zeitschr. Berlin, p. 147-162.
Krogerus, H., 1954. Investigations on the Lepidoptera of Newfoundland. I. Macro-

lepidoptera. — Acta Zool. Fenn. 82. Helsingfors, p. 1-80.

Bibliography 317

KuLLENBERG, B., 1946. tJber die Verbreitung und Wanderungen von vier Sterna-

Arten. — Ark. f. Zool. 38 A :i7. Stockholm, p. 1-80.
KusNEZOV, N. J., 1935. The origin of the Lepidopterous fauna of the Arctic Eurasia.

— Arctica. 111:3. Leningrad, p. 1 15-136.

Lauterborn, R., 1927. Beitrage zur Flora der oberrheinischen Tiefebene und der

benachbarten Gebiete. — Mitt. Bad. Landesver. Naturk. & Naturschutz (N.S.)

2. Freiburg i. Br., p. 77-88.
Leng, C. W., 1920. Catalogue of the Coleoptera of America, north of Mexico. —

Mount Vernon, N.Y., pp. i-x, 1-470.
LiNDROTH, A., 1942. Garfageln i Sverige. — Var Fagelvarld. i. Stockholm, p. 138-

140.
LiNDROTH, C. H., 193 1. Die Insektenfauna Islands und ihre Probleme. — Zool.

Bidr. 13. Uppsala, p. 105-589.

— 1935. The Boreo-British Coleoptera. A study of the faunistical connections
between the British Isles and Scandinavia. — Zoogeogr. 11:4. Jena, p. 579-634.

— 1946. Inheritance of wing dimorphism in Pterostichus anthracinus 111. — Hered-
itas. 32. Lund, p. 37-40.

— 1948. Interglacial insect remains from Sweden. — Sver. Geol. Unders. C 492.
Stockholm, p. 1-30.

— 1954a. Random notes on North American Carabidae (Coleopt.). — Bull. Mus.
Comp. Zool. 111:3. Cambridge, Mass., p. 117-161.

— 1954b. Carabidae common to Europe and North America. — Coleopt. Bull. 8.
Washington, D. C, p. 35-52.

— 1954c. A revision of Diachila Motsch. and Blethisa Bon. — K. Fysiogr. Sallsk.
Handl. (N.S.) 65. Lund, p. 1-28.

— 1955. The Carabid beetles of Newfoundland, including the French islands St.
Pierre and Miquelon. — Opusc. Ent., Suppl. XII. Lund, p. 1-168.

LoHAMMAR, G., 1954. Matsmaltningens inverkan pa Potamogeton-fronas groning.

— Fauna o. Flora. 49. Uppsala, p. 17-32.

Lowe, A., 1950. Some innovations and nomeclatural suggestions in the Icelandic

flora. — Bot. Notiser. Lund, .p. 24-60.
LuNDQViST, G., 1955. Stocken i Oje. Ett sakert interglacialfynd. — Geol. Foren.

Forh. 77. Stockholm, p. 317-322.
Maklix, F. W., 1878. Diagnoser ofver nagra nya sibiriska insektarter. — Ofvers.

Finsk. Vet.-Soc. Forh. 19 (1876-77). Helsingfors, p. 15-32.

— 1881. Coleoptera insamlade under den Nordenskiold'ska expeditionen 1875, &c.

— K. Sv. Vet. Ak. Handl. 18:4. Stockholm, p. 1-48.

Matthew, W. D., 1915. Climate and evolution. — Ann. N.Y. Ac. Sci. 24. New

York, p. 171-318.
Mayr, E., 1946. History of the North American bird fauna. — Wilson Bull. 58.

Ann Arbor, Mich., p. 3-41.
McDuNNOUGH, J. H., 1950. Species of Euxoa of eastern North America, &c. —

Bull. Amer. Mus. Nat. Hist. 95:6. New York, p. 355-408.
Mertens, R. & MuLLER, L., 1940. Die Amphibien und Reptilien Europas. — Abh.

Senck. Naturf. Gesellsch. 451. Frankfurt a. M., p. 1-56.
MiCHENER, C. D. & M. H., 195 1. American social insects. — New York, pp. i-xiv,

1-267.

3l8 CHAPTER III

MiCHAELSEN, W., 1 922. Die Verbreitung dcr Oligochaten im Lichte der Kontinental-

verschiebung, &c. — Verb. Naturw. Verein. (3) 29 (1921). Hamburg, p. 45-79.
MoDEl,L, H., 1943. Tertiare Najaden. III. Nordamerikanische Najaden im bayrischen

Oligozan. — Arch. f. Molluskenk. 75. Frankfurt a.M., p. 107-117.
Nannfeldt, J. A., 1940. On the polymorphy of Poa arctica R. Br., with special

reference to its Scandinavian forms. — Symb. Bot. 4. Uppsala.
Nathorst, a. G., 1894. Die Entdeckung einer fossilen Glacialflora in Sachsen, am

aussersten Rande des nordischen Diluviums. — Ofvers. K. Vet. Ak. Forh. 51.

Stockholm, p. 519-543.
Needham, J. G. & Westfall, M. J., Jr., 1955. A manual of the Dragonflies of North

America (Anisoptera). — Berkeley & Los Angeles, p. 1-615.
NORDHAGEN, R., 1916. Ranunculus Cymbalaria Pursh. fundet i Norge. — Nyt Mag.

f. Naturv. 55. Oslo, p. 1 19-145.

— 1933. De senkvartaere klimavekslinger i Nordeuropa og deres betydning for
kulturforskningen. — Inst. Sammenlign. Kulturforskn. A. 12. Oslo, p. 1-246.

— 1935. Om Arenaria humifusa Wg. og dens betydning for utforskningen av
Skandinavias eldste floraelement. — Berg. Mus. Arbok. Bergen, p. 1-183.

Obrutschew, W. a., 1926. Geologic von Sibirien. — Fortschr. Geol. & Palaeont. 15.

Berlin, pp. l-xi, 1-572.
Oekland, F., 1925. Die Verbreitung der Landgastropoden Norwegens. — Skrift.

Norsk. Vid.-Ak., I. Mat.-Naturv. Kl. 8. Oslo, pp. i-viii, 1-168.

— 1927. Einige Argumente aus der Verbreitung der nordeuropaischen Fauna mit
Bezug auf Wegeners Verschiebungstheorie. — Nyt. Mag. f. Naturv. 65. Oslo, p.

339-367-
Orchymont, a. de & Brown, W. J., 1940. Helophorus arcticus Brown, a living

fossil. — Can. Ent. 72. Ottawa, p. 1-4.
Ostenfeld, C. H., 1926. The flora of Greenland and its origin. — Biol. Medd. K.

Dansk. Vid. Selsk. 6:3. Copenhagen, p. 1-71.
Palmen, E., 1944. Die anemohydrochore Ausbreitung der Insekten als zoogeogra-

phischer Faktor, &c. — Ann. Zool. Soc. Zool. Bot. Fenn. Vanamo. 10:1. Hel-

singfors, pp. i-v, 1-262.
Peters, J. L., 1948. Check-List of birds of the world. VI. — Cambridge, Mass.,

pp. i-xi, 1-259.
Petersen, B., 1954. Some trends of speciation in the cold-adapted Holarctic fauna.

— Zool. Bidr. 30. Uppsala, p. 233-314.
Peterson, R. L., 1952. A review of the living representatives of the genus Alces. —

Contr. R. Ont. Mus. Zool. & Palaeont. 34. Toronto, p. 1-30.
PiLSBRY, H. A., 1894. Helicidae. — Manual of Conch. (Tryon & Pilsbry). 2. Pul-

monata. IX. — Philadelphia, pp. i-xlviii, 1-366.

— "939, 1946, 1948. Land Mollusca of North America (north of Mexico). I:i,
II: 1-2. — Ac. Nat. Sci., Monogr. 3. Philadelphia, pp. l-xvii, 1-573; i-vi, i-
1113.

PiROZNiKOV, P. L., 1937. A contribution to the study of the origin of the northern
elements in the fauna of the Caspian Sea. — C. R. Ac. Sci. URSS. 15. Moscow,
p. 521-524.

Pleske, T., 1928. Birds of the Eurasian tundra. — Mem. Bost. Soc. Nat. Hist. 6:3.
Boston, p. 11-485.

Bibliography 319

PouLSEN, E. M., 1939. Freshwater Crustacea. — Zool. of Icel. 111:35. Copenhagen,

p. 1-50.
Praeger, R. L., 1932. Recent views bearing on the problem of the Irish flora and

fauna. — Proc. R. Irish Ac. B. 41. Dubhn, p. 125-145.

— 1934- The botanist in Ireland. — Dublin.

Reinig, W. F., 1937. Die Holarktis. Ein Beitrag zur diluvialen und alluvialen Ge-
schichte der zirkumpolaren Faunen- und Florengebiete. — Jena, pp. l-vil,
1-124.

— 1938. Elimination und Selektion. — Jena, pp. l-viii, 1-146.

Reitter, E., 1875. Beitrag zur Kenntniss der aussereuropaischen Crytophagidae. —
Coleopt. Hefte (E. v. Harold). 13. Miinchen, p. 73-87.

— 191 1. Fauna Germanica. III. — Stuttgart, p. 1-436.

Rensch, B., 1950. Die Verteilung der Tierwelt im Raum. — Handb. d. Biol. V: i.

Potsdam, p. 125-172.
SiEMUNDSSON, B., 1939. Mammalia. — Zool. of Icel. IV: 76. Copenhagen, p. 1-38.
Sahlberg, J., 1880. Bidrag till Nordvestra Sibiriens insektfauna. Coleoptera, &c. —

K. Svensk. Vet. Ak. Handl. 17. Stockholm, p. 1-115.
Salomonsen, F., 1 95 1. The immigration and breeding of the fieldfare (Turdus

pilaris L.) in Greenland. — Proc. X. Intern. Ornith. Congr. (1950). Uppsala, p.

515-526.
Salomonsen, F. & Gitz-Johansen, 1950. Gronlands Fugle. The Birds of Green-
land. — Copenhagen, p. 1-608.
Sann, J., 1952. (On Penguinus [Alca] impennis.) — Fauna. 5. (Norsk Zool. Foren.)

Drammen, p. 74-75.
ScHARFF, R. F., 1907. European animals: their geological history and geographical

distribution. — London, pp. l-xiv, 1-258.

— 1909. On the evidence of a former land-bridge between Northern Europe and
North America. — Proc. R. Irish Ac. 28 B. Dublin, p. 1-28.

— 191 1. Distribution and origin of life in America. — London, pp. i-xvi, 1-497-
ScHERHAG, R., 1948. Neue Methoden der Wetteranalyse und Wetterprognose. —

Berlin, pp. i-xii, 1-424.
Schmidt, K. P., 1946. On the zoogeography of the Holarctic region. — Copeia. Ann

Arbor, Mich., p. 144-152.
Segerstrale, S. G., 1954. The freshwater Amphipods, Gammarus pulex (L.) and

Gammarus lacustris G. O. Sars, in Denmark and Fennoscandia, &c. — Comment.

Biol., Soc. Sci. Fenn. XV: i. Helsingfors, p. 1-91.
Semenov, a., 1935. Analecta coleopterologica. XXI. — Rev. Russ. d'Ent. 25.

Moscow, p. 275.
Simpson, G. G., 1940. Mammals and land bridges. — Journ. Wash. Ac. Sci. 30.

Washington, D.C., p. 137-163.

— 1947. Holarctic Mammalian faunas and continental relationships during the
Cenozoic. — Bull. Geol. Soc. Amer. 58. New York, p. 613-687.

Smith, M. R., 1951. Formicidae. — In: Muesebeck, Krombein, Townes, Hymeno-
ptera of America, &c. — U. S. Dept. Agric, Agric. Monogr. 2. Washington, D. C,

p. 778-875.
SpArck, R., 1934. Freshwater Sponges. — Zool. of the Faroes. I:i, 4. Copenhagen,

P- 1-3-

320

CHAPTER III

Stebbins, R. C, 195 1. Amphibians of Western North America. — (Univ. Calif.

Press.) Berkeley & Los Angeles, pp. i-Xvii, 1-539.
Stegmann, B., 193S. Principes generaux des subdivisions ornithogeographiques de

la region palearctique. (Russ.) — Fauna SSSR. (N. S.) 19. Moscow & Leningrad,

P- I-IS7-
Stephens, Jane, 1920. The fresh-water Sponges of Ireland. — Proc. R. Irish Ac. 35.

Dublin, p. 205-254.

Stephensen, K., 1921. Gronlands Dyreverden. — Medd. om Gronl. 60. Copen-
hagen.

Sterner, R., 1945. Nordiska havsstrandsvaxter. En vaxtgeografisk overblick. —
Fauna o. Flora. Uppsala, p. 28-43.

Szafer, W., 1954. Pliocene flora from the vicinity of Czorsztyn (West Carpathians)
and its relationship to the Pleistocene. — Prace Inst. Geol. 11. Warszawa, p.
1-238.

Szekessv, v., 1936. Revision der boreoalpinen Koleopteren auf vergleichend-ana-
tomischer Grundlage. II. — Ent. Tidskr. 57. Stockholm, p. 97-126.

Tambs-Lyche, H., 1937. Forekomsten av Ranunculus Cymbalaria Pursh i 0stfold
og Bohuslan. — Nyt Mag. f. Naturv. 77. Oslo, p. 15-38.

Taylor, J. W., 1906-14. Monograph of the land & freshwater MoUusca of the British
Isles. III. — Leeds, p. 1-522.

Thienemann, a., 1950. Verbreitungsgeschichte der Susswassertierwelt Europas. —
Stuttgart, pp. i-xvi, 1-809.

Thoroddsen, Th., 1 9 14. An account of the physical geography of Iceland, &c. —
Bot. of Icel. I, 2. Copenhagen.

Thorkelsson, Th., 1935. A fossiliferous interglacial layer at EUidaarvogur, Reyk-
javik. — Greinar (Soc. Scient. Isl.) I:i. Reykjavik, p. 1-14.

Timmermann, G., 1938-49. Die Vogel Islands. 1: 1-2. — Visindafelag Islendinga
(Soc. Scient. Isl.) 21. 24. 28. Reykjavik, p. 1-524.

Timofeeff-Ressovsky, N. W., 1947. Das Treffersprinzip in der Biologic. — Bio-
physik. I. Leipzig, pp. i-xn, 1-3 17.

Tuxen, S. L., 1944. The hot springs of Iceland, their animal communities and their
zoogeographical significance. — Zool. of Icel. 1: 11. Copenhagen, p. 1-216.

Valle, K. J., 1952. Die Verbreitungsverhaltnisse der ostfennoskandischen Odonaten.
— Acta Ent. Fenn. 10. Helsingfors, p. 1-87.

Vandel, a., 1945. La repartition geographique des Oniscoidea (Crustaces Isopodes
terrestres). — Bull. Biol. France & Belg. 79. Paris, p. 221-272.

Van Dyke, E. C, 1939. The origin and distribution of the Coleopterous insect fauna
of North America. — Proc. 6. Pac. Sci. Congr. 4. San Francisco, p. 255-268.

Vibe, C, 1950. Some insects new to Greenland. — Ent. Medd. 25. Copenhagen, p.
419-420.

Vlerk, I. M. van der & Florschutz, F., 1953. The palaeontological base of the sub-
division of the Pleistocene in the Netherlands. — Verb. K. Nederl. Ak. Wet., Afd.
Naturk. (I) 20:2. Amsterdam, p. 1-58.

Wahlgren, E., 1 91 9. Vastarktiska element i Skandinaviens fjarilfauna. — Ent. Tid-
skr. 40. Stockholm, p. 22-32.

Walter, H., 1954. Einfiihrung in die Phytologie. III. Grundlagen der Pflanzen-
verbreitung. II. Arealkunde. — Stuttgart, p. 1-245.

Bibliography 321

Wegener, A., 1929. Die Entsthehung der Kontinente und Ozeane. 4. Aufl. — Die

Wissenschaft. 66. Braunschweig, pp. l-x, 1-23 1.
West, A., 1930. Coleoptera. — Zool. of. the Faroes. 40. Copenhagen, p. 1-92.
Wheeler, W. M., 1900. The habits of Ponera and Stigmatomma. — Biol. Bull.

Marine Biol. Lab. Woods Hole. 2. Boston, p. 43-69.

— 1915- The ants of the Baltic amber. — Schr. Phys.-Okon. Gesellsch. Konigs-
berg. 55. Leipzig & Berlin, p. 1-142.

Williamson, K., 1954. American birds in Scotland in autumn and winter, 1953-54.

— Scott. Nat. 66. Aberdeen, p. 13-29.

Woldstedt, p., 1954. Das Eiszeitalter. Grundlinien einer Geologic des Quartars. 2.
Aufl. — Stuttgart, pp. i-xv, 1-406.

— 1955. Norddeutschland und angrenzende Gebiete im Eiszeitalter. (Geogr. Handb.)

— Stuttgart, pp. i-xii, 1-467.

WuRTEMBERG, W. J., 1919. HcHx hortensis. — Nautilus. 33. Philadelphia, p. 71.
Zeuner, F. E., 1945. The Pleistocene Period, its climate, chronology and faunal
successions. — (Ray Soc.) London, pp. l-xil, 1-322.

— 1950. Dating the past. An introduction to geochronology. — London, pp.
i-xviii, 1-474.

2 1 — 565597 Littdroih

Conclusions

The animal species common to Europe and North America are numerous. If
conditions among the groups here treated in Chapter I, comprising altogether
908 species in common, are regarded as representative, a calculation on the basis of
the Swedish fauna (Gislen, 1940), in which these groups make 17,3 per cent of the
entire terrestrial and limnic fauna, would lead to the conjecture that the actual
number of Eur- American non-marine animal species amounts to more than 5,000.
The true figure is probably higher because the microfauna, here disregarded, in-
cludes forms with a distribution above average.

A large part of the species in common are regarded as introduced with man into
either continent, or both, this element amounting to between 41 and 46 per cent
(375-417 species). As the birds, with no unintentionally introduced species, are
among the groups treated, these figures are probably not too high, if applied to
the calculated total of Eur-American species.

Species introduced from Europe into North America are about ten times as
numerous as those transported in the opposite direction. This is explained by
the peculiar character of ballast traffic in olden times, sailing vessels going almost
exclusively in ballast on their way west, to Newfoundland and the Maritime Pro-
vinces of Canada on the one hand, to the Pacific Northwest on the other. The
main area of departure for these animals, mostly belonging to the soil fauna, was
southwestern England.

A contributory reason why the contingent of introduced European animals is
particularly large in the coastal regions of northeastern and northwestern North
America is, of course, that the climate of these parts of the continent is most
similar to that of western Europe.

Among groups treated, the number of animal species undoubtedly indigenous
in both continents exceeds 500. Most of them possess a more or less continuous
circumpolar distribution. A special interest, however, is connected with the
so-called Amphiatlantic (incl. "Westarctic") species, which are lacking or have
a broadly interrupted area on the Pacific side of the globe. They have often been
used as an argument in favour of earlier transatlantic land-connections.

Attempts are made to demonstrate that the cases of Amphiatlantic, and similar,
distribution can be understood, partly by over-sea dispersal under present condi-

Conclusions 323

tions, partly as remnants of a broken-up area of circumpolar type. From a bio-
geographical point of view, the necessity for any kind of land-connection in the
past, directly joining the European with the North American continent, is thus
denied. Like Simpson (1947, p. 666, footnote) I started my investigation "with a
feeling that a north Atlantic bridge was probable" and like him I dare hope that
abandoning this prejudice reduces the subjectivity of opinion.

On the other hand, it seems unevitable to assume that part of the fauna of
Greenland and Iceland immigrated across a Pleistocene land-bridge from the
European mainland. An interesting point is that this invasion did not at all affect
the fauna of Baffin Island, which is completely devoid of a European element.
Actually, the narrow strait between Greenland and Baffin Island is the most
pronounced north-south barrier within the arctic region of the Holarctic area.

Since the Bering land-bridge, repeatedly at work during Tertiary and Pleistocene
time, was the most important link toward evolution of the circumpolar fauna, a de-
tailed comparative study of the Alaskan and Northeast Siberian faunas is the most
important task for the future.

Index

OF CHAPTER II AND III
Species, Genera and certain Families

Animals

aalge, Uria
Abax ater, 193, 203
Ablattaria laevigata, 186, 205
Acidota crenata, 260
acuminatus, Neoclytus
Acupalpus dubius, 193, 203

meridianus, 144, 170, 193, 195, 203
Adalia = Coccinella
adstrictus, Pterostichus
Aegialia arenaria, 169, 179, 188, 208,
fig. 26

rufa, 157
aenea, Amara

aeneus, Harpalus, Meligethes, Olibrus
Aeschna subarctica, 233
aestivum, Apion
aethiops, Notaris

affinis, Harpalus, Scolopostethus
Agabus congener, 259

moestus, 259

solieri, 259
Agelena labyrinthica, 190
agilis, Dicranomerus
Agonum atratum, 193, 203

bogemanni, 149, 287

consimile, 149, 287

dorsale, 184, 193, 203

exaratum, 287

mannerheimi, 149, 287, 302

micans, 193, 203

mijlleri, 139, 144, 149, 193, 203, 215

obscurum, 193, 203

quadripunctatum , 149, 287

ruficorne, 144, 149, 153, 177, 193, 203,

215
thoreyi, 149, 287

Agriotes lineatus, 138, 188, 207, 215

sputator, 169
Ag>'neta cauta, 231
alata, Islandiana
alauda, Cionus
albifrons, Anser
albipes, Tachypodoiulus
AIca, vide also Penguinus
Alca torda, 231

Alces alces, 297, fig. 54, diagr. 10
Aleochara bipustulata, 186, 205

obscurella, 260
Alfalfa Weevil =Phytonomus posticus
alleghanensis, Cryptobranchus
alliarius, Oxychilus
Allolobophora caliginosa, 201
Alopecosa pulverulenta, 190
Alopex lagopus, 292
alpestris, Vertigo
alpina, Amara, Erolia

Amara aenea, 142, 144, 150, 184, 193, 195,
203, 215, 220

alpina, 150, 258, 264, 287, 302, fig. 40

anthobia, 143, 144, 170, 192

apricaria, 142, 144, 150

aulica, 142, 144, 150, 151, 159, 193,
203, 258

avida, 142

bifrons, 142, 144, 150, 151

communis, 193, 203

convexior, 193, 203

convexiuscula, 193, 203, 209

equestris, 193, 203, 209

erratica, 150, 287

familiaris, 142, 144, 150, 193, 203

fulva, 142, 144, 150, 151, 154, 159

INDEX

325

Amara hyperborea, 150, 287

impuncticollis, 142

interstitialis, 287

latior, 142

lunicollis, 142, 144, 150, 193, 203, 287

majuscula, 138

montivaga, 193, 203

ovata, 193, 203

pallipes, 142

patruelis, 142

plebeja, 193, 195, 196, 203

quenseli, 150, 258, 287

similata, 193, 203

tibialis, 193, 203

torrida, 150, 287
Amischa analis, 186, 200, 201, 205, 260,
268

decipiens, 186, 205
Amphipyra tragopogonis, 237
Anacaena globulus, 259
analis, Amischa
Anarta, 243

funebris, 239

lapponica, 231, fig. 30

leucocycla, 233

melaleuca, 239

melanopa, 233

zetterstedti, 239
Anas crecca, 251
ancillaria, Physa
Andrias scheuchzeri, 284
Androniscus dentiger, 191
Anergates atratulus, 234
Angoumois Grain Moth = Sitotroga
anguineus, Proteus
angustatus, Xantholinus
angusticollis, Harpalus
angustula, Atheta
angustus, Polydesmus

Anisodactylus binotatus, 144, 170, 193, 203
annulipes, Dictyna
Anomogyna laetabilis, 239
Anser albifrons, 247
Anthicus humilis, 188, 207
anthobia, Amara
anxius, Harpalus
Apanteles glomeratus, 220, 222

Aphelinus mali, 223
Aphodius, 170

fasciatus, 214

fimetarius, 188, 208, 215

granarius, 188, 208

lapponum, 262

nodifrons = fimetarius

putridus = fasciatus

rufipes, 188, 208

tenellus = fasciatus
Apion aestivum, 188, 208

apricans, 188, 208

assimile, 188, 208

cruentatum, 262

flavipes, 188, 208

hookeri, 188, 208

miniatum = sanguineum

nigritarse, 188, 208

ononicola, 188, 208

ononis, 188, 208

sanguineum, 188, 208

ulicis, 188, 208

violaceum, 188, 208

virens, 188, 208
apricans, Apion
apricaria, Amara

aquaticus, Helophorus, Notiophilus
Araneus redii, 190
arboreus, Zonitoides
Arctia quenseli, 239

arctica, Diachila, Fratercula, Miscodera
arcticola, Pterostichus
arcticus, Helophorus, Otiorrhynchus
Arctosa leopardus, 190

perita, 190
Arenivaga, 283
Arenocoris falleni, 189
Argyroploce pomonella, 214, 222
arietis, Clytus
Arion ater, 201

circumscriptus, 148
Armadillidium depressum, 191

vulgare, 191, 215
Army Worm = Leucania
Arpedium brachypterum, 260
Asaphidion flavipes, 144, 193, 203
asellus, Oniscus

326

INDEX

Ashfordia granulata, 191

asperatus, Sciaphilus

aspersa, Helix

assimile, Apion, Bembidion

assimilis, Ceuthorrhynchus, Patrobus

Astenus longelytratus, 186, 205

Astilbus canaliculatus, 186, 205

astutiis, Philonthus

Atactogenus Cneorrhinus

ater, Abax, Staphylinus

Atheta angustula, i86, 205

atramentaria, 186, 205, 260

excellens, 260

exigua, 186, 205

fungi, 186, 205, 260, 267

graminicola, 260

hyperborea, 259, 260, 272

islandica, 259, 260

melanocera, 260

oblita, 186, 205

orbata, 186, 205

triangulum, 186, 205

vestita, 186, 205, 260

sp., 263
Atomaria fuscata, 188, 207
atomus, Malthodes
atramentaria, Atheta
atratulus, Anergates
atratum, Agonum
atricapillus, Demetrias
atriceps, Tachyporus
atroapterus, Otiorrhynchus
atrorufus, Patrobus
attenuatus, Harpalus
Auk = Penguinus
aulica, Amara
auratus, Carabus
auricularia, Forficula
auropunctatus, Otiorrhynchus
Autalia puncticollis 260
Autographa = Plusia
avida, Amara

baccarum, Dolycoris
Badister bipustulatus, 193, 203
banksi, Chrysomela
Barnacle-Goose Branta

Barynotus moerens, 168

obscurus, 215

schonherri squamosus

squamosus, 168, 262, 268
bassanus, Morus
Bathyphantes nigrinus, 231

pullatus, 240
Bathyplectes curculionis, 222
beddardi, Bimastus
Bembidion assimile, 193, 203

bipunctatum, 258

bruxellense, 144, 150, 151

concinnum, 177, 193, 203

crenuiatum, 287

dauricum, 287

grapei, 150, 258, 275-278, 287, figs. 44-
47, diagr. 8

grapeioides =yukonum

guttula, 193, 203

hasti, 287

hyperboraeorum, 287

iricolor, 193, 203

lampros, 138, 144, 150, 153, 154, 158
193-195. 203

lapponicum, 287, 298

laterale, 179, 193, 203

lunulatum, 193, 203

mckinleyi, 287

minimum, 193, 203

normannum, 193, 203

obscurellum, 287

petrosum, 150, 287

properans, 143, 144, 168, 193, 203

quadrimaculatum, 150, 194, 203, 287,
288

redtenbacheri, 258

rupestre = bruxellense

scandicum = mckinleyi

stephensi, 144, 150, 151

tetracolum, 139, 144, 150, 153, 154, 158,
194, 203, 214, 215, fig. 12

transparens, 150, 287, 305, figs. 58, 59

ustulatum = tetracolum

varium, 194, 203

yukonum, 287
Beosus maritimus, 189
bidentata, Clausilia, Diplocentria

INDEX

327

bifrons, Amara
biguttatus, Notiophilus
bimaculatum, Theridium
Bimastus beddardi, 213

tenuis, 201
binotatus, Anisodactylus
bipustulata, Aleochara
bipustulatus, Badister
bipunctata, Coccinella
bipunctatum, Bembidion
blandingi, Emys
Blechrus = Microlestes
Bledius spectabilis, 179, 187, 205
Blethisa eschscholtzi, 280, 293, fig. 56

multipunctata, 150, 287, 302, fig. 56
bogemanni, Agonum
boletophagum, Megasternum
boops, Quedius

borealis, Pelophila, Rhaebothorax
Boreaphilus nearcticus, 259, 260

nordenskioldi, 259
Brachyiulus littoralis, 191
brachypterum, Arpedium
Bradycellus, 283

harpalinus, 144, 170, 194, 204

verbasci, 194, 204
Branta leucopsis, 251
brevicollis, Gymnusa, Nebria
brevicornis, Pterostichus
brevilingue, Micralymma
brevipalpis, Helophorus
britannicus, Cylindroiulus
Broscus cephalotes, 194, 204
Brown-tail Moth = Euproctis
brunnipes, Stenus
bruxellense, Bembidion
Bryoporus rufescens, 260

rugipennis, 260

sp., 260
bucculentus, Stenomyphantes
Bucephala islandica, 253
buchanani, Colpodes
Burbot = Lota
Byrdia groenlandica, 256
Byrrhus fasciatus, 261, 272
Bythinus puncticollis, 261

validus = puncticollis

Cabbage Butterfly = Pieris rapae
Cabbage White = Pieris rapae
Caenoscelis ? cryptophaga, 257, 261, 263

macilenta, 263
caesareus, Staphylinus
caespitum, Tetrannorium
Cafius xantholoma, 260
Calathus erratus, 194, 204

fuscipes, 144, 170, 194, 204, 258

melanocephalus, 194, 204, 258

micropterus, 150, 287

mollis, 194, 204, 209
calcaratum, Stenodema
caliginosa, Allolobophora
Calosoma sycophanta, 222
canaliculatus, Astilbus, Stenus
cancellatus, Carabus
Candidula = Helicella
Cantharis rufa, 188, 207

rustica, 188, 207
cantiana, Monacha
caperata, Helicella
Carabus auratus, 143, 144, 169, 192

cancellatus, 143, 144, 169, 192

catenulatus = problematicus

chamissonis, 310

granulatus, 139, 142, 143, 144, 168, 169,
fig. 8

groenlandicus = chamissonis

nemoralis, 136, 139, 144, 150, 151, 153,
154, 169, 237, fig. 7

problematicus, 237, 258

truncaticollis, 287, 288

violaceus, 194, 204
Caradrina = Rhyacia
carbo, Phalacrocorax
carbonarius, Stenus
Caribou = Rangifer
Carolina, Tetracha
carolinensis, Eisenia
catascopium, Lymnaea
Catharacta skua, 231, 242
caudatus, Ephialtes
cauta, Agyneta
cavicollis, Gnypeta
cellarius, Oxychilus
Cepaea hortensis, 148, 234, 285, fig. 29

328

cephalotes, Broscus

Cepphus grylle, 231

Cercyon litoralis, 244, 259

cerealella, Sitotroga

Cernuella = Helicella

Ceuthorrhynchus assimilis, 170, 188, 208

hirtulus, 188, 208

troglodytes, 189, 208
chalceus, Pogonus
Chalcoides fulvicornis, 188, 208
chamissonis, Carabus
Charadrius hiaticula, 231, 242
Chartoscirta cocksi, 189
Cheiracanthium erraticum, 190
chiragra, Megalonotus
Chrysolina '-^ Chrysomela
Chrysomela banksi, 188, 202, 208

staphylea, 262
chrysomelinus, Tachyporus
chrysorrhoea, Euproctis
Cicindela, 293
Cidaria ruberata, 233

sabini, 239, 242
Cionella lubrica, 191

minima, 191
Cionus alauda, 189, 208

scrophulariae, 189, 208
circellaris, Sipalia
circumscriptus, Arion
clandestina, Spaelotis
clathrata, Linyphia
Clausilia bidentata, 191
clavicornis, Stenus

Clover Leaf Weevil = Phytonomus punctatus
Clivina collaris, 143, 144, 186, 192, 215

elongata ^ collaris

fossor, 139, 143, 145. 150. 153. 154. 158,
186, 192, fig. 13
Clubiona compta, 190

neglecta, 190

norvegica, 233

reclusa, 190

terrestris, 190

trivialis, 233
Clytus arietis, 188, 208
Cneorrhinus exaratus, 189, 208
coarctata, Ponera

INDEX

coarcticoUis, Stenus

Coccidula rufa, 188, 207

Coccinella bipunctata, 188, 207, 215
quatuordecim-(i4-)punctata, 188, 207
septem-(7-)punctata, 188, 207
transversoguttata, 256, 262, 263
undecim-(i i-)punctata, 142, 262, 268
vigintiduo-(22-)punctata, 188, 207

cochleariae, Phaedon

Cochlicopa Cionella

cocksi, Chartoscirta

Codling Moth = Argyroploce

Coenonympha tullia, 297, 302, fig. 55

coerulescens, Pterostichus

cognatus, Trichocellus

Colias, 243
hecla, 243
nastes, 240, 242, 243

collaris, Clivina, Stenichnus

Collinsia thulensis, 256

colonoides, Nemadus

Colorado Beetle = Leptinotarsa

Colpodes buchanani, 172

columella, Lymnaea

Colymbetes dolabratus, 253, 259, 263, 272,
274, 292

Colymbus = Gavia

communis, Amara

complanata, Eurynebria

complanatus, Pristonychus

complicatus, Leptyphantes

compta, Clubiona

concinnus, Philonthus

concolor, Sminthurus

confluens, Polydrosus

congener, Agabus

Conosoma lividum, 187, 205
pubescens = testaceum
testaceum, 187, 205

consimile, Agonum

convexior, Amara

convexiuscula, Amara

Coreus marginatus, 189

coriaceus, Polydesmus

Cormorant = Phalacrocorax

Corn Borer = Pyrausta

Corncrake = Crex

INDEX

329

Corticarina fuscula, 188, 207

gibbosa, 188, 207
corticinus, Tachinus
Corymbetes cupreus, 138

sjaelandicus, 237
crecca, Anas

Crematogaster lineolata, 213
crenata, Acidota
crenulatum, Bembidion
Crex crex, 251
cribricollis, Otiorrhynchus
Crino sommeri, 253, 255, fig. 37
cristatus, Xysticus
crocata, Dysdera
cruentatum, Apion
cruentatus, Philonthus
Crustulina sticta, 23 1

Crymodes exulis, 231, 243, 253, 255, fig. 38
Cryptobranchus alleghaniensis, 284
cryptophaga, Caenoscelis
Cryptophagus vini, 188, 207
Cryptops hortensis, 191

parisi, 191
Ctenium lividum, 189
cubensis, Lymnaea
cunea, Hyphantria
Cupedidae, 252, 308, fig. 60
Cupes rafFrayi, 307
cupreus, Corymbites, Pterostichus
curculionis, Bathyplectes
curtipennis, Quedius
Cydia = Argyroploce
cylindracea, Lauria
Cylindroiulus britannicus, 191

frisius, 191

silvarum, 191

teutonicus, 191
Cymindis unicolor, 258
Cytilus sericeus, 188, 207, 261

Danaus plexippus, 243
Dasychira = Byrdia
dauricum, Bembidion
decemlineata, Leptinotarsa
decipiens, Amischa
decoratus, Scolopostethus
deflexum, Liosoma

22 — 565507 Lindroth

degeeri, Pachygnatha
degenerata, Nebria
Demetrias atricapillus, 194, 204
Dendrobaena octaedra, 201
subrubicunda, 201
dentiger, Androniscus
depressum, Armadillidium
Deroceras laeve, 201
reticulatum, 148
Deronectes multilineatus, 259
destructor, Mayetiola
Diachila arctica, 240, 287

polita, 287, 304
Diaptomus minutus, 253
diasema, Plusia
Dichirotrichus pubescens, 179, 194, 204,

209, 258
dicksoni, Micralymma
Dicranomerus agilis, 189
Dictyna annulipes, 233
Dicymbium nigrum, 190
diligens, Pterostichus
Diplocentria bidentata, 233
discus, Trechus
dispar, Lymantria
Discus rotundatus, 191
Diver = Gavia
dolabratus, Colymbetes
Dolycoris baccarum, 189
domesticus, Passer
dorsale, Agonum
Dorytomus taeniatus, 262, 263
sp., 257, 263
dougalli. Sterna
draparnaldi, Oxychilus
Drassodes lapidosus, 190
drewseni, Rhyacia
Dromius linearis, 194, 204

melanocephalus, 194, 204
Drosophila, 294
Drymus sylvaticus, 189
dubius, Acupalpus, Otiorrhynchus
Dunlin = Erolia alpina
Dyschirius globosus, 194, 204

helleni, 287

politus, 150, 287
Dysdera crocata, 189

33°

INDEX

Earwig = Forficula
Eisenia carolinensis, 213

foetida, 201

rosea, 201
Eiseniella tetraedra, 201
Elaphrus lapponicus, 240, 287

riparius, 150, 287
Elk = Alces

Elleschus scanicus, 169
Elm Leaf Beetle = Galerucella
elongata, Clivina
elongatus, Phytonomus
Emys blandingi, 282

orbicularis, 282
English Sparrow = Passer
Enicmus transversus, 188, 207
Eperigone maculata, 213
Ephialtes caudatus, 222
ephippiatus, Lacinius
equestris, Amara
erichsoni, Nematus
Eriosoma lanigerum, 212, 223
Erolia alpina, 251

maritima, 231, 242
erratica, Amara
erraticum, Cheiracanthium
erratus, Calathus
erythrocephala, Theatops
erythrocephalus, Hydroporus
eschscholtzi, Blethisa
Estrandia grandaeva, 240
Euophrys frontalis, 190

molesta, 190
Eupithecia gelidata, 233
Euproctis chrysorrhoea = phaeorrhoea

phaeorrhoea, 222
Eurygaster testudinaria, 189
Eurynebria complanata, 194, 204, 209
Euxoa = Rhyacia
exaratum, Agonum
exaratus, Cneorrhinus
excellens, Atheta
excentrica, Vallonia
exigua, Atheta
extensa, Tetragnatha
exulis, Crymodes

Falagria obscura, 187, 205

Fall Webworm = Hyphantria

fallax, Ophyiulus

falleni, Arenocoris

familiaris, Amara

fasciata, Phlegra

fasciatus, Aphodius, Byrrhus, Gammarus

ferrugineum, Pachymerium

festivus, Phrurolithus

Fiber zibethicus, 135, fig. 5

fimetarius, Aphodius, Philonthus

flavescens, Sitona

flavipes, Apion, Asaphidion, Halobrecta,

Heliophanes, Helophorus, Meligethes,

Stenus
foetida, Eisenia
Forficula auricularia, 189, 215
forficatus, Lithobius
formicetorum, Stenus
fossor, Clivina
foveatus, Metabletus
Fox = Alopex
Fratercula arctica, 231
frigida, Hilaira
frigidaria, Scopula
frisius, Cylindroiulus
frontalis, Euophrys
fulgidus, Quedius
fuligineus, Stygnocorus
fuliginosus, Harpalus, Quedius
Fulmar = Fulmarus
Fulmarus glacialis, 231, fig. 27
fulva, Amara
fulvicollis, Quedius

fulvicornis, Chalcoides, Lamyctes, Stenus
fulvipenne, Lahtrobium
fulvus, Haliplus, Trechus
funebris, Anarta
fungi, Atheta
fuscata, Atomaria
fuscipennis, Philonthus
fuscipes, Calathus, Paederus
fuscula, Corticarina
fuscus, Oedothorax, Proteroiulus, Spelerpes

Gabrius Philonthus
Galerucella luteola, 215, 221

INDEX

331

Galerucella xanthomelaena = luteola
gamma, Plusia
Gammarus fasciatus, 252

lacustris, 252

tigrinus = fasciatus
Gannet = Morus
Gavia immer, 253
gelidata, Eupithecia
gemellatus, Sitona
geminum, Lathrobium
genei, Hydromantes
Geodromicus globulicoUis, 260
gibbosa, Corticarina
gibbus, Phylan
glacialis, Fulmarus
globosus, Dyschirius
globulicoUis, Geodromicus
globulus, Anacaena
glomeratus, Apanteles
Glomeris marginata, 191
Gnaphosa orites, 233
Gnypeta cavicollis, 260, 272
Goldeneye = Bucephala
Gonatium ruberis, 237
Grain Moth = Sitotroga
graminicola, Atheta
granarius, Aphodius
grandaeva, Estrandia
grandis, Lebia
granulata, Ashfordia
granulatus, Carabus
Grape Phylloxera = Phylloxera
grapei, Bembidion
grapeioides, Bembidion
Green-veined White ^ Pieris napi
griseus, Sitona
groenlandica, Byrdia
groenlandicus, Carabus
grylle, Cepphus
Guillemot = Cepphus and Uria
Gull = Larus
guttula, Bembidion
gyllenhali, Nebria
Gymnetron netum, 169
Gymnusa brevicollis, 257, 260
Gypsy Moth = Lymantria
Gyraulus parvus, 214

gyrina, Physa

Gyrinus opacus, 259, 263

gyrosicollis, Stomodes

Habrobracon, 223
haematopus, Pterostichus
Haliplus fulvus, 259
Halobrecta flavipes, 187, 205
Haplodrassus signifer, 237
Haplophilus subterraneus, 191
Hare = Lepus

Harlequin Duck = Histrionicus
harpalinus, Bradycellus
Harpalobrachys leiroides, 288
Harpalus aeneus = affinis

affinis, 145, 150, 153, 194, 195, 204, 214,

215
angusticollis = puncticeps
anxius, 194, 204
attenuatus, 194, 204
fuliginosus, 150, 288
latus, 194, 204
nigritarsis, 150, 288
pubescens = rufipes
puncticeps, 194, 204
quadripunctatus, 258
rubripes, 194, 204
rufipes, 145, 150, 151, 154, 159, 169,

194, 195

seladon, 194, 204

tardus, 194, 204

tenebrosus, 194, 204

viridiaeneus = affinis
Harpocera thoracica, 189
hasti, Bembidion
Hawaiia minuscula, 213
hecla, Colias
heeri, Lesteva
Helicella caperata, 191

virgata, 191
Helicodiscus parallelus, 213
Heliophanes flavipes, 190
Helisoma duryi, 214

tenuis, 214

trivolve, 214
Helix aspersa, 191
Helix, also = Cepaea

332

INDEX

helleni, Dyschirius
Helodes minuta, 261
Helophorus aquaticus, 259

arcticus, 259

brevipalpis, 186, 205, 259

flavipes, 259
helveticus, Oxychilus
Hessian Fly = Mayetiola
Heteromeyenia ryderi, 241, 245, 249, 251,

255
heterostropha, Physa
hiaticula, Charadrius
Hilaira frigida, 240
Hillia iris, 240
hirtula, Rhamphomyia
hirtulus, Ceuthorrhynchus
hirundo, Sterna
hispida, Hygromia
hispidulus, Sitona
hispidus, Trachodes
Hister purpurascens, 187, 206
Histrionicus histrionicus, 253
hoffmannseggi, Platyarthrus
hookeri, Apion
hortensis, Cepaea, Cryptops
House Sparrow ^ Passer
humeralis, Sitona
humilis, Anthicus
Hydnobius punctatus, 259
Hydrobia jenkinsi, 201
Hydromantes, 285

genei, 281, fig. 48

platycephalus, 282, fig. 48
Hydroporus erythrocephalus, 259

lapponum, 259

melanocephalus, 259, 263

memnonius, 259

nigrita, 259

palustris, 259

polaris, 259

pubescens, 259

sp., 272
Hygromia hispida, 191
striolata, 148, 191
hyperborea, Amara, Atheta
hyperboraeorum, Bembidion
Hyphantria cunea, 212

Hypnoidus riparius, 261
hypnorum, Tachyporus
Hypocyptus longicornis, 187, 205
Hypolithus = Hypnoidus

immer, Gavia
impennis, Penguinus
impressicollis, Ochthebius
impressifrons, Polydrosus
impressus, Stenus
impuncticollis, Amara
inconstans, Polydesmus
interjectus, Silometopus
inuncta, Podops
iricolor, Bembidion
iris, Hillia
Islandiana alata, 233

princeps, 256
islandica, Atheta, Bucephala, Oxypoda

jenkinsi, Hygrobia
juno, Stenus

Kittiwake = Rissa
kochi, Xysticus
Koenenia, 283
krameri, Umbra

labradorensis, Lepyrus
labyrinthica, Agelena
Lacinius ephippiatus, 190
lacustris, Gammarus, Umbra
laetabilis, Anomogyna
laevigata, Ablattaria
laevigatum, Stenodema
laeviusculum, Omalium
lagopus, Alopex
laminatus, Philonthus
lampros, Bembidion
Lampyris noctiluca, 187, 207
Lamyctes fulvicornis, 201
Land-Rail = Crex
lanigerum, Eriosoma
lapidosus, Drassodes
lapponica, Anarta
lapponicum, Bembidion
lapponum, Aphodius, Hydroporus

INDEX

333

Lapwing = Vanellus
Larch Sawfly = Nematus
Larus marinus, 231

minutus, 251
laterale, Bembidion
Lathrobium fulvipenne, 257, 260, 267

geminum, 187, 205
latior, Amara
latreilli, Tylos
latus, Harpalus
Lauria cylindracea, 191
Lebia grandis, 222
leiroides, Harpalobrachys
leopardus, Arctosa
lepidus, Sitona
Leptinotarsa decemlineata, 135, 142, 212,

218, 222, fig. 3
Leptyphantes complicatus 233

minutus, 231

tenuis, 190
Lepus timidus, 271
Lepyrus labradorensis, 262
Lesteva heeri, 187, 205

longelytrata, 260
Leucania unipuncta, 212
leucocycla, Anarta
leucopsis, Branta

Libellula quadrimaculata, 289, fig. 51
Licinus punctulatus, 145
Ligia oceanica, 191
ligneus, Otiorrhynchus
Limax marginatus, 148

maximus, 148, 236
limbata, Rhagonycha
limbatus, Zyras
limonii, Scymnus
linearis, Dromius

lineatus, Agriotes, Sitona, Stenomyphantes
lineolata, Crematogaster
Linyphia clathrata, 190

pusilla, 190
Liosoma deflexum, 189, 208
liquidus, Olibrus
listeri, Pachygnatha
Lithobius forficatus, 191
litoralis, Cercyon, Paederus
littoralis, Brachyiulus, Salda

litura, Rhizobius

lividum, Conosoma, Ctenium

lomvia, Uria

longelytrata, Lesteva

longelytratus, Astenus

longicornis, Hypocyptus

Loricera pilicornis, 150, 194, 204, 258

Lota lota, 302

lubrica, Cionella

luctuosum, Orygma

lunicollis, Amara

lunulatum, Bembidion

lurida, Oedemera

luteola, Galerucella

Lygus rugulipennis, 189

Lymantria dispar, 135, 222

Lymnaea, 201

catascopium, 214, 249

columella, 214

cubensis, 214

limosa = peregra

ovata = peregra

peregra, 148, 251
Lymnastis, 282

macilenta, Caenoscelis
macrodon, Mustela
maculata, Eperigone
madidus, Pterostichus
majuscula, Amara
Malachius viridis, 188, 207
mali, Aphelinus
Malthodes atomus = pumilus

mysticus, 261

pumilus, 261
mannerheimi, Agonum
marginata, Glomeris
marginatus, Coreus, Limax, Philonthus
marinum, Micralymma
marinus, Larus, Ochthebius
maritima, Erolia

maritimus, Beosus, Quedius, Tibellus
maurus, Microlestes
maximus, Limax, Megalobatrachus
Mayetiola destructor, 214, 222
mckinleyi, Bembidion
Megacephala = Tetracha

334

INDEX

Megalobatrachus maximus, 284
Megalonotus chiragra, 189

praetextatus, 189
MegaSternum boletophagum, 186, 205, 259
melaleuca, Anarta
melanarius, Pterostichus
Melanimon tibiale, 188, 207
melanocephalus, Calathus, Dromius, Hydro-

porus, Othius
melanocera, Atheta
melanogrammus, Strophosomus
melanopa, Anarta
Melanophthalma = Corticarina
Meligethes aeneus, 188, 207

flavipes, 188, 207

picipes, 188, 207
memnonius, Hydroporus
merianae, Meta
meridianus, Acupalpus
meridionalis, Otiorrhynchus
Mesoleius tenthredinidis, 222
mesomelinus, Quedius
Meta merianae, 190
Metabletus foveatus, 194, 204
metallica, Simplocaria
Metaponorthus 283
micans, Agonum
Micaria pulicaria, 190, 200
Miccotrogus = Tychius
Micralymma brevilingue, 256, 260, 263, 292

dicksoni, 261

marinum, 231, 244, 260, 263, fig. 32
Micrambe = Cryptophagus
Micraspis sedecim-(i6-)punctata, 188, 207
Micratopus, 283
Microlestes, 283

maurus, 194, 204
micropterus, Calathus
miniata, Xeroclysa
miniatum, Apion
minima, Cionella
minimum, Bembidion
minuscula, Hawaiia
minuta, Helodes

minutus, Diaptomus, Larus, Leptyphantes
mirabilis, Pisaura
Miscodera arctica, 150, 288, 302

moerens, Barynotus

moestus, Agabus

molesta, Europhrys

mollis, Calathus

molochinus, Quedius

Monacha cantiana, 191

Monocephalus parasiticus, 231

montivaga, Amara

Moose = Alces

Morus bassanus, 231, 242, fig. 28

moschatus, Ovibos

Mouse = Mus

Mud-minnows ^^ Umbra

Mud Puppies = Necturus

miilleri, Agonum

multilineatus, Deronectes

multipunctata, Blethisa

Murre = Uria

Mus musculus, 271

muscorum, Philoscia

musculus, Mus

Muskox = Ovibos

Muskrat ^ Fiber

Mustela macrodon, 234

Mycetoporus splendidus, 187, 205

sp., 260
myrmecoides, Nabis
myrmecophilus, Othius
Myrmecopora uvida, 187, 205
Myrmedonia = Zyras
mysticus, Malthodes

Nabis myrmecoides, 189

rugosus, 189
nana, Tachyta
nanus, Stenus
napi, Pieris
nastes, Colias
nasulus, Stylulus
nearcticus, Boreaphilus

Nebria brevicollis, 145, 150, 153, 157, 194,
195, 204

degenerata = salina

gyllenhali, 150, 257, 258, 271, 288, 302

nivalis, 150, 258, 288, 302

rufescens = gyllenhali

salina, 194, 204, 258

INDEX

335

Necturus, 281

neglecta, Clubiona

Nemadus colonoides, 307

Nematus erichsoni, 222

nemoralis, Carabus

Neoclytus acuminatus, 212

netum, Gymnetron

niger, Porcellio, Pterostichus

nigriceps, Pardosa

nigrinus, Bathyphantes

nigrita, Hydroporus, Pterostichus

nigritarse, Apion

nigritarsis, Harpalus

nigritulus, Philonthus

nigrum, Dicymbium

nitidiusculus, Stenus

nitidula, Retinella

nitidulus, Tachyporus

nitipennis, Quedius

nivalis, Nebria

nobilis, Oedemera

noctiluca, Lampyris

nodifrons, Aphodius

Nomius pygmaeus, 280

schoutedeni, 280
Nopoiulus pusillus = venustus

venustus, 214
nordenskioldi, Boreaphilus
normannum, Bembidion
Northern Army Worm = Leucania
norvegica, Clubiona
Notaris aethiops, 262
Notiophilus aquaticus, 150, 258, 288

biguttatus, 143, 145, 151, 154, 194, 204,
258, 268, fig. 14

palustris, 194, 204

substriatus, 194, 204
nubilalis, Pyrausta
nubilus, Peritrechus
Nyctea scandiaca, 289, fig. 49

oblita, Atheta
obscura, Falagria
obscurella, Aleochara
obscurellum, Bembidion
obscurum, Agonum
obscurus, Barynotus

obtusus, Tachyporus, Trechus, Tropipho-

rus
Ocalea picata, 260
oceanica, Ligia
Ochthebius impressicollis, 186, 205

marinus, 186, 205
Ocneria = Lymantria
octaedra, Dendrobaena
octona, Subulina
Oedemera lurida, 188, 207

nobilis, 188, 207
Oedothorax fuscus, 190

retusus, 190
Oleacinidae, 283, 308
olens, Staphylinus
Olibrus aeneus, 188, 207

liquidus, 188, 207
Olisthopus rotundatus, 194, 204, 258
Olm = Proteus
Omalium laeviusculum, 260

riparium, 260
Ondatra = Fiber
Oniscus asellus, 191, 215, 237
ononicola, Apion
ononidis, Phytonomus
ononis, Apion
opaciceps, Ponera
opacus, Gyrinus
Opeas pumilum, 213
operculella, Phthorimaea
Ophyiulus fallax, 191
opilio, Phalangium
orbata, Atheta
orbicularis, Emys
orbiculatus, Stilicus
orites, Gnaphosa
Orygma luctuosum, 244
Othius melanocephalus, 260, 278

myrmecophilus, 260

punctulatus, 260
Otiorrhynchus, 200, 209, 224

arcticus, 257, 262, 264, 268, 270, fig. 41

atroapterus, 262

auropunctatus, 168

cribricollis, 170

dubius, 257, 262

ligneus, 189, 208

336

INDEX

OtJorrhynchus meridionalis, 170

ovatus, 189, 208, 215

rugifrons, 262, 268

rugQsostriatus, 189, 208, 257

scabcr, 168

sulcatus, 189, 208, 215
ovata, Amara
ovatum, Theridium
ovatus, Otiorrhynchus
Ovibos moschatus, 245, 303
Owl = Nyctea
Oxychilus alliarius, 191

cellarius, 191

draparnaldi, 148, 191

helveticus, 191
Oxypoda islandica, 260

soror, 260
Oxytelus tetracarinatus, 187, 205

Pachygnatha degeeri, 190

listeri, 190
Pachymerium ferrugineum, 169
Paederus fuscipes, 187, 205

litoralis, 187, 205
pallipes, Amara
Palomena prasina, 189

palustris, Hydroporus, Notiophilus, Pardosa
Pantala flavescens, 286
parallelus, Helicodiscus
parasiticus, Monocephalus
Pardosa nigriceps, 190

palustris, 190

proxima, 190

puUata, 190

saltuaria, 233
parilis, Plusia
parisi, Cryptops
parvulus, Tachys
parvus, Gyraulus
Passer domesticus, 135, 219
Patrobus assimilis, 258

atrorufus, 258

septentrionis, 151, 258, 271, 288
patruelis, Amara
Pear Psylla = Psylla
pedestris, Phytonomus, Zelotes
pcllucida, Vitrina

Pelophila borealis, 151, 288, 302

Penguinus impennis, 244, fig. 33

peregra, Lymnaea

Perileptus, 282

perita, Arctosa

Peritrechus nubilus, 189

Petrobius, 177

petrosum, Bembidion

Phaedon cochleariae, 188, 200, 208

tumidulus, 188, 208
phaeorrhoea, Euproctis
Phalacrocorax carbo, 231
Phalangium opilio, 190
Philomachus pugnax, 251
Philonthus astutus, 187, 205

concinnus, 187, 205

cruentatus, 187, 205

fimetarius, 187, 205

fuscipennis, 187, 205

laminatus, 187, 205

marginatus, 187, 205

nigritulus, 187, 206

trossulus, 260

varians, 187, 206

varius, 187, 206
Philopedon plagiatus, 179, 189, 208, fig. 26
Philorinum sordidum, 187, 206
Philoscia muscorum, 191
Phlegra fasciata, 231
Phrurolithus festivals, 190
Phthorimaea operculella, 223
Phylan gibbus, 188, 207
Phyllodecta polaris, 262
Phyllotreta undulata, 188, 208
Phylloxera vastatrix = vitifoliae

vitifoliae, 212, 219, 223
Phymaphora pulchella, 257
Physa ancillaria, 214

gyrina, 214

heterostropha, 214
Phytometra = Plusia
Phytonomus elongatus, 257, 262

ononidis, 189, 208

pedestris, 262

posticus, 135, 169, 189, 208, 221

punctatus, 221

variabilis = posticus

INDEX

337

Phytophaga = Mayetiola

picata, Ocalea

picipennis, Quedius

picipes, Meligethes, Quedius, Stenus

picirostris, Tychius

Picoides tridactylus, 289, 303, fig. 50

pictus, Porcellio

Pieris napi, 220

rapae, 135, 220, 222
Piesma quadrata, 189
pilaris, Turdus
pilicornis, Loricera
Pisaura mirabilis, 190
plagiatus, Philopedon
Platyarthrus hoffmannseggi, 234
platycephalus, Hydromantes
plebeja, Amara
plexippus, Danaus
Plover = Charadrius, Vanellus
Plusia diasema, 240

gamma, 266

parilis, 231, fig. 31
Pocadicnemis pumila, 233
Podops inuncta, 189
Pogonus chalceus, 183, 194, 204
polaris, Hydroporus, Phyllodecta
polita, Diachila
politus, Dyschirius
Polydesmus angustus, 191

coriaceus =inconstans

inconstans, 191
Polydrosus confluens, 189, 208

impressifrons, 169
pomonella, Argyroploce
Ponera atavia, 252

coarctata, 231, 252, fig. 35

opaciceps, 213
Porcellio, 283

niger = scaber

pictus =spinicornis

scaber, 191, 215
spinicornis, 215
Porthetria = Lymantria
posticus, Phytonomus
Potamopyrgus = Hydrobia
Potato Tuber Moth = Phthorimaea
potentillae, Sibinia

praetextatus, Megalonotus

prasina, Palomena

princeps, Islandiana

Pristiphora = Nematus

Pristonychus complanatus, 145, 170

terricola, 145, 154, 158, 165
properans, Bembidion
Propylaea = Coccinella
Proteroiulus fuscus, 201
Proteus anguineus, 281
provisorius, Trichoniscus
proxima, Pardosa
Psylla pyricola, 215
Pterostichus, 297

adstrictus, 151, 258, 288

anthracinus, 275

arcticola, 258

brevicornis, 151, 288

coerulescens = versicolor

cupreus, 195, 204

diligens, 258, 271

haematopus, 258

madidus, 195, 204

melanarius, 136, 139, 145, 151, 154, 158

niger, 195, 204

nigrita, 195, 204, 224, 258

strenuus, 145, 151, 153, 154, 158, 195, 204

vermiculosus, 288

vemalis, 195 204

versicolor, 195, 204

vulgaris = melanarius
puberulus, Scolopostethus
pubescens, Conosoma, Dichirotrichus, Har-

palus, Hydroporus
Puffin = Fratercula
pugnax, Philomachus
pulchella, Phymaphora
pulicaria, Micaria
pullata, Pardosa
puUatus, Bathyphantes
pumicatus, Stomis
pumila, Pocadicnemis
pumilum, Opeas
pumilus, Malthodes
punctatus, Hydnobius, Phytonomus
puncticeps, Harpalus
puncticollis, Autalia, Bythinus, Sitona

338

INDEX

punctulatus, Licinus, Othius

purpurascens, Hister

pusilla, Linyphia

pusillus, Nopoiulus, Tachyporus, Tricho-

niscus
putridus, Aphodius
pygmaeus, Nomius
Pyrausta nubilalis, 222
pyricola, Psylla

quadrangula, Rhyacia
quadrata, Piesma
quadrimaculata, Libellula
quadrimaculatum, Bembidion
quadripunctatum, Agonum
quadripunctatus, Harpalus
quadristriatus, Trechus
quatuordecim-(i4-)punctata, Coccinella
Quedius hoops, 257, 260, 267

curtipennis, 187, 206

fulgidus, 263

fuliginosus, 187, 206, 261

fulvicoUis, 261

maritimus, 187, 206

mesomelinus, 263

molochinus =picipennis

nitipennis, 261

picipennis, 187, 206

picipes, 187, 206

rufipes, 187, 206

schatzmayri, 187, 206

subfuliginosus = fuliginosus

tristis, 187, 206

umbrinus, 261
quenseli, Amara, Arctia

raflfrayi, Cupes

Rangifer tarandus, 292, 297, fig. 53

rapae, Pieris

Razorbill = Alca

reclusa, Clubiona

redii, Araneus

redtenbacheri, Bembidion

regensteinensis, Sitona

Reindeer = Rangifer

remota, Simplocaria

reticulatum, Deroceras

Retinella nitidula, 191
retusus, Oedothorax
Rhaebothorax borealis, 231
Rhagonycha linibata, 188, 207
Rhamphomyia hirtula, 253
rhombeus, Syromastus
Rhyacia drewseni, 256

quadrangula, 253, fig. 36
riparium, Omalium
riparius, Elaphrus, Hypnoidus
Rissa tridactyla, 231
Roeseliana roeselii, 223
roeselii, Roeseliana
romanus, Styloctetor
rosea, Eisenia

rotundatus. Discus, Olisthopus
rubens, Gonatium, Trechus
ruberata, Cidaria
rubripes, Harpalus
rufa, Aegialia, Cantharis
rufescens, Bryoporus, Nebria
Ruff = Philomachus
ruficorne, Agonum
rufinasus, Stenopelmus
rufipes, Aphodius, Harpalus, Quedius, Ta-

chinus
rugifrons, Otiorrhynchus
rugipennis, Bryoporus
rugosostriatus, Otiorrhynchus
rugosus, Nabis
rugulipennis, Lygus
rupestre, Bembidion
ruricola, Trochosa
rustica, Cantharis
rusticola, Scolopax
ryderi, Heteromeyenia

sabini, Cidaria

sabulosum, Schizophyllum

Salamander = Cryptobranchus, Hydroman-

tes, Megalobatrachus
Salda littoralis, 189
salina, Nebria
saltuaria, Pardosa
Sandpiper = Erolia maritima
sandvicensis, Thalasseus
sanguineum, Apion

INDEX

339

scaber, Otiorrhynchus, Porcellio
scandiaca, Nyctea
scanicus, Elleschus
scarabaeoides, Spaeridium
schatzmayri, Quedius
scheuchzeri, Andrias
Schizophyllum sabulosum, 191
schonherri, Barynotus
schoutedeni, Nomius
Sciaphilus asperatus, 262, 268
Scolopax rusticola, 251
Scolopostethus affinis, 189

decoratus, 189

puberulus, 189
Scopula frigidaria, 240, 242
scrophulariae, Clonus
Scymnus limonii, 262, 267
sedecim-(i6-)punctata, Micraspis
seladon, Harpalus
semistriata, Simplocaria
septempunctata, Coccinella
septentrionis, Patrobus
sericeus, Cytilus
Sibinia potentillae, 189, 209
signifer, Haplodrassus
Silometopus interjectus, 190
Silpha tristis, 186, 205
silvarum, Cylindroiulus
similata, Amara
similis, Stenus
Simplocaria metallica, 262

remota, 262

semistriata, 181, 188, 207, 261

tessellata, 261, 262, 263
Simulium vittatum, 253, fig. 39
Sipalia circellaris, 187, 206, 261
Sirenians, vide Trichechus
Sitona, 215

flavescens = lepidus

gemellatus, 168

griseus, 189, 209

hispidulus, 189, 209

humeralis, 189, 209

lepidus, 181, 189, 209, 215

lineatus, 170, 189, 209

puncticollis, 189, 209

regensteinensis, 189, 209

Sitona sulcifrons, 189, 209

Sitotroga cerealella, 214

sjaelandicus, Corymbites

skua, Catharacta

Small White = Pieris rapae

Sminthurus concolor, 256

solieri, Agabus

solutus, Tachyporus

sommeri, Crino

sordidum, Philorinum

soror, Oxypoda

Spaelotis clandestina, 240, 256

unicolor = clandestina
Sparrow, English or House, = Passer
spectabilis, Bledius
Spelerpes = Hydromantes
Sphaeridium, 170

scarabaeoides, 186
spinicornis, Porcellio
splendidus, Mycetoporus
sputator, Agriotes
staphylea, Chrysomela
Staphylinus ater, 187, 206

caesareus, 187, 206

olens, 187, 206
Starling = Sturnus
Stenichnus collaris, 259
Stenocellus = Bradycellus
Stenodema calcaratum, 189

laevigatum, 189
Stenolophus teutonus, 195, 204
Stenomyphantes bucculentus, 190

lineatus = bucculentus
Stenopelmus rufinasus, 212
Stenus brunnipes, 187, 206, 261

canaliculatus, 187, 206, 261

carbonarius, 261

clavicornis, 187, 206, 261

coarcticollis, 261

flavipes, 187, 206

formicetorum, 187, 206

fulvicornis, 187, 206

impressus, 187, 206, 261

juno, 187, 206

nanus, 187, 206, 261

nitidiusculus, 261

picipes, 187, 206

34°

INDEX

Stenus similis, 187, 206

subaeneus, 187, 206
stephensi, Bembidion
Sterna dougalli, 231, 242

hirundo, 231, 242
sticta, Crustulina
Stilbus testaceus, 188, 207
Stilicus orbiculatus, 187, 206
Stomis pumicatus, 143, 145, 181, 195, 204
Stomodes gyrosicoUis, 169
strenuus, Pterostichus
striatus, Thanatus
striolata, Hygromia
Strobilops, 308, fig. 6i
Strobilus = Strobilops
Strophosomus nielanogrammus, 262, 268
Sturnus sturnus, 135, 219, fig. 4
Stygnocorus fuligineus, 189
Styloctetor romanus, 190
Stylulus nasulus, 282
subaeneus, Stenus
subarctica, Aeschna
Subcoccinella vigintiquatuor-(24-)punctata,

188, 207
subfuliginosus, Quedius
subrubicunda, Dendrobaena
substriatus, Notiophilus
subterraneus, Haplophilus, Tachinus
Subulina octona, 213
Sula = Morus
sulcatus, Otiorrhynchus
sulcifrons, Sitona
sycophanta, Calosoma
sylvaticus, Drymus
Syromastus rhombeus, 189

Tachinus corticinus, 261, 272

rufipes, 187, 206, 261

subterraneus, 187, 206
Tachistodes, 283
Tachypodoiulus albipes, 191
Tachyporus atriceps, 261

chrysomelinus, 187, 206

hypnorum, 187, 206

nitidulus, 187, 206

obtusus, 187, 206

pusilius, 187, 206

Tachyporus solutus, 187, 206
Tachys, 283

parvulus, 145, 170
Tachyta nana, 288
taeniatus, Dorytomus
tarandus, Rangifer
tardus, Harpalus
Teal = Anas crecca
tenebrosus, Harpalus
tenellus, Aphodius
tenthredinidis, Mesoleius
tenuis, Bimastus, Helisoma, Lepthyphantes
Tern = Sterna, Thalasseus
terrestris, Clubiona
terricola, Pristonychus, Trochosa
tessellata, Simplocaria
testaceum, Conosoma
testaceus, Stilbus
testudinarea, Eurygaster
tetracarinatus, Oxytelus
tetracolum, Bembidion
tetraedra, Eiseniella
Tetragnatha extensa, 190
Tetramorium caespitum, 234
Tetrastichus, 221
teutonicus, Cylindroiulus
teutonus, Stenolophus
Thalasseus sandvicensis, 231, 242
Thanatus striatus, 233
Thea = Coccinella
Theba = Monacha
Theatops erythrocephala, 280
Theridium, 171

bimaculatum, 190

ovatum, 190
thoracica, Harpocera
thoreyi, Agonum
thulensis, Collinsia
Tibellus maritimus, 190
tibiale, Melanimon
tibialis, Amara
tigrinus, Gammarus
timidus, Lepus
tomentosus, Tropiphorus
torda, Alca
torrida, Amara
Trachodes hispidus, 157

INDEX

34^

tragopogonis, Amphipyra
transparens, Bembidion
transversoguttata, Coccinella
transversus, Enicmus
Trechus discus, 143, 145

fulvus, 258

obtusus, 143, 145, 170, 195, 204, 258

quadristriatus, 195, 205, 209

rubens, 145, 151, 154
triangulum, Atheta
Trichechus, 283
Trichia = Hygromia
Trichocellus cognatus, 151, 258, 288
Trichoniscus provisorius, 191

pusillus, 201
tridactyla, Rissa
tridactylus, Picoides
tristis, Quedius, Silpha
trivialis, Clubiona
trivolve, Helisoma
Trochosa ruricola, 190

terricola, 190
troglodytes, Ceuthorrhynchus
Tropiphorus obtusus, 168, 262, 268

tomentosus, 168
truncaticollis, Carabus
Tuber Moth = Phthorimaea
tullia, Coenonympha
tumidulus, Phaedon
Turdus pilaris, 266, fig. 42
Turtle, vide Emys

Tychius picirostris, 137, 168, 189, 209
Tylos latreilli, 280, 285
Tytthaspis = Micraspis

ulicis, Apion
Umbra, 285

krameri, 281

lacustris = krameri
umbrinus, Quedius
undecim-(i i-)punctata, Coccinella
undulata, Phyllotreta
unicolor, Cymindis, Spaelotis
unipuncta, Leucania
Uria aalge, 231

lomvia, 231

ustulatum, Bembidion
uvida, Myrmecopora

validus, Bythinus

Vallonia excentrica, 148, 237

Vanellus vanellus, 25 1

variabilis, Phytonomus

varians, Philonthus

varium, Bembidion

varius, Philonthus

venustus, Nopoiulus

verbasci, Bradycellus

vermiculosus, Pterostichus

vernalis, Pterostichus

versicolor, Pterostichus

Vertigo alpestris, 148, 236

vestita, Atheta

vigilax, Walckenaera

vigintiduo-(22-)punctata, Coccinella

vigintiquatuor-(24-)punctata, Subcoccinella

Vine Aphis = Phylloxera

vini, Cryptophagus

violaceum, Apion

violaceus, Carabus

virens, Apion

virgata, Helicella

viridiaeneus, Harpalus

viridis, Malachius

vitifoliae. Phylloxera

Vitrina pellucida, 191

vittatum, Simulium

vulgare, Armadillidium

vulgaris, Pterostichus

Walckenaera vigilax, 231, 244
Webworm = Hyphantria
White = Pieris
Woodcock = Scolopax
Woodpecker = Picoides
Woolly Aphis = Eriosoma

Xantholinus angustatus, 187, 206
xantholoma, Cafius
xanthomelaena, Galerucella
Xeroclysa miniata, 190
Xysticus cristatus, 190
kochi, 190

342

yukonum, Bembidion

Zelotes pedestris, 190
zetterst'edti, Anarta

INDEX

zibethicus, Fiber
Zonitoides arboreus, 213
Zyras limbatus, 187, 206

Plants

Achillea millefolium, 174, 180, 197
Agrostis sp., 181
Alchemilla filicaulis, 146

minor, 146

pastoralis, 146
Alliaria officinalis =petiolata

petiolata, 180, 197
Alopecurus ventricosus, 146

sp., 176
Ammophila arenaria, 178, 198
Anagallis arvensis, 178, 198
Anisantha sterilis, 174, 197
Artemisia vulgaris, 174, 176, 197, 198
Atriplex sp., 176
Azolla caroliniana, 212, 241

filiculoides, 309

Ballota nigra, 174, 197
Bellis perennis, 173, 180, 197
Beta vulgaris, 183, 198
Brassica nigra, 178, 197
Bromus = Anisantha

Calystegia sepium, 174, 197
Capsella bursa-pastoris, 174, 177, 197
Cardamine flexuosa, 146
Carduus crispus, 180, 197
Carex fulva = hostiana

hostiana, 251
Centaurea nigrescens, 146
Cerastium vulgatum, 174, 180, 197
Chaerophyllum temulum, 180, 197
Chamaenerium Epilobium
Chrysanthemum leucanthemum, 147, 180,

197
Cirsium arvense, 173, 174, 178, 180, 197,
198

lanceolatum vulgare

palustre, 146

vulgare, 174, 180, 197

Cochlearia danica, 146
Convolvulus = Calystegia
Coronopus didymus, 174, 197
Crepis sp., 176, 178, 180
Cynoglossum officinale, 178, 198
Cytisus = Sarothamnus

Dactylis glomerata, 173, 178, 180, 197
Daucus carota, 178, 197
Dulichium spathaceum, 309
spp., 309, 310

Elodea canadensis, 212, 241
Entomophthora sphaerosperma, 222
Epilobium adenocaulon = glandulosum

glandulosum, 212

hirsutum, 174, 197

latifolium, 253
Equisetum arvense, 174, 197
Erigeron canadense, 212
Eriocaulon septangulare, 240, 241, 250
Erodium cicutarium, 178, 198
Euphorbia paralias, 178, 198

portlandica, 178, 198

Festuca ovina, 178, 197
Fragaria vesca, 146
Funaria hygrometrica, 176

Galium aparine, 146, 180, 197

saxatile, 146
Geranium dissectum, 174, 180, 197

ibericum, 146

robertianum, 180, 197, 198
Glaux maritima, 183, 198
Gnaphalium sylvaticum, 146

Habenaria hyperborea, 253

straminea, 253
Halimione pedunculata, 183, 198

INDEX

343

Hieracium, 146
Holcus lanatus, 174, 180, 197
Hordeum murinum, 174, 176, 197, 198
Hypochaeris radicata, 174, 180, 197

Juncus acutiflorus, 146
bulbosus, 146
Dudleyi, 241
effusus, 176
macer = tenuis
tenuis, 212, 241

Kentranthus ruber, 178, 197, 198

Lappula echinata, 146
Lathyrus pratensis, 146
Lepidium densiflorum, 213
Leucorchis = Habenaria
Limosella subulata, 240, 250
Linum catharcticum, 146
Lobelia dortmanna, 241
Lolium perenne, 180, 197
Lotus corniculatus, 173, 178, 180, 197
Luzula campestris, 146
pallescens, 146

Matricaria discoidea =matricarioides
maritima, 176, 180, 197, 198
matricarioides, 177, 197, 198, 213
suaveolens = matricarioides

Melilotus altissima, 180, 197

Mertensia maritima, 244

Montia rivularis, 146

Myosotis arvensis, 180, 197

Myriophyllum alterniflorum, 240, 250

Najas flexilis, 241
Nardus stricta, 146

Obione = Halimione
Oenothera biennis, 213
Ononis arvensis, 168

repens, 178, 198
Osmunda claytoniana, 309, 310
Oxalis stricta, 213

Pedicularis palustris, 146

sylvatica, 146
Phragmites communis, 183, 198
Picea abies, 297

glauca, 297

obovata, 297
Plantago coronopus, 173, 197

lanceolata, 173, 174, 178, 197

major, 174, 177, 180, 197

media, 173, 180, 197
Poa annua, 174, 177, 180, 184, 197

pratensis, 173, 197

trivialis, 174, 180, 197
Polygonum hydropiper, 146

Raii, 146
Potamogeton epihydrus, 241, 250

oblongus = polygonifolius

polygonifolius, 251
Potentilla anglica, 146

erecta, 146

reptans, 178, 197

sterilis, 146
Puccinellia maritima, 183, 198

Ranunculus cymbalaria, 213, 244

repens, 174, 177, 180, 197
Reseda luteola, 178, 197
Rubus sp., i8i
Rumex crispus, 174, 180, 197

Salicornia sp., 179
Sarothamnus scoparius, 173, 197
Scrophularia nodosa, 146

sp., 180
Senecio jacobaea, 176, 178, 180, 197, 198

vulgaris, 176, 177, 178, 197
Sieglingia decumbens, 146
Silene cucubalus, 180, 197

vulgaris = cucubalus
Sisymbrium officinale, 176, 177, 197
Sisyrinchium angustifolium, 155, 241, 247,
249, 250, 251

montanum, 155
Solanum dulcamara, 180, 197

rostratum, 219

tuberosum, 219
Sonchus oleraceus, 180, 197

344

INDEX

Sper^jularia marina ^ salina

rubra, 146

salina, 176, 198
Spiranthes romanzoffiana, 241, 250, 251
Stellaria media, 177, 197

Triglochin maritimum, 183, i<
Tsuga sp., 308

Ulex europaeus, 173, 180, 197
Urtica dioica, 178, 180, 197

Taraxacum sp., 176, 178
Trifolium, 154

campestre, 173, 178, 180, 183, 197

hybridum, 176, 197

pratense, 147, 173, 176, 180, 183, 197

procumbens = campestre

Veronica officinalis, 146

peregrina, 213
Vicia angustifolia, 173, 176, 180, 197

cracca, 173, 197
Vitis spp., 219

vinifera, 219

repens, 176, 177, 178, 180, 181, 183, 197 Vulpia bromoides, 173, 197

Marine Biological Laboratory

Received,

July 26, 1958

A M 73810

Accession No.

Given By_

John Wiley and Sons, Inc.

Place N®^ York City