Stowaway Beetles: Carl Lindroth, the Ballast Theory, and Transatlantic Science in the Cold War
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Matthew Evenden
Stowaway Beetles: Carl
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Lindroth, the Ballast Theory,
and Transatlantic Science in
the Cold War
Abstract
In his Faunal Connections between Europe and North
America (1957), a landmark study of ecological introduc-
tions from Europe to North America that prefigured Alfred
Crosby’s Columbian Exchange by three decades, Carl
Lindroth sought to explain the distribution of fauna and
particularly insects across the North Atlantic. It was the cul-
mination of a multi-year investigation begun in 1949 that
had seen the Swedish ecologist and entomologist collect
insects in Newfoundland, compare his findings with other
North American and European regions, and trace the ori-
gins of introduced insects to various sites in southwestern
England. Through this work, he identified the ballast of sail-
ing ships as a primary vector of insect introduction to the
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Matthew Evenden, “Stowaway Beetles: Carl Lindroth, the Ballast Theory, and
Transatlantic Science in the Cold War,” Environmental History 26 (2021): 508–532
doi: 10.1093/envhis/emab022
Advance Access Publication Date: 9 April 2021
Stowaway Beetles 509
Americas, argued for the importance of recurring introduc-
tions of breeding pairs in successful colonization, and dem-
onstrated the importance of examining a broad range of
species introductions, not just those that directly mattered
to human history. Conceived and executed in the context of
the Cold War, Lindroth’s project benefited from govern-
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mental and institutional support that sought to promote
transatlantic scientific cooperation and northern research.
His path to researching and writing Faunal Connections
reveals both the influence of geopolitics in shaping environ-
mental ideas and the capacity for scientists to benefit from
strategic funding opportunities while contributing little of
strategic value. Lindroth’s work deserves reexamination,
both for its substance and for what it can tell us about the
ideas behind foundational texts like The Columbian
Exchange and, indeed, the making of ideas about global
ecological change.
In his landmark study Faunal Connections between Europe and North
America (1957), Swedish entomologist Carl Lindroth explained that
“Newfoundland more than any other part of North America has re-
ceived an introduced element of animals and plants from Europe.”1 It
is a striking point even today, perhaps especially for environmental
historians. Accustomed to beginning discussions of transatlantic bio-
logical exchange with Alfred Crosby’s classic The Columbian Exchange,
a study in which Newfoundland does not appear by name, environ-
mental historians have tended to emphasize the sites and stories that
highlight biological exchanges as a force in human history without
reckoning with the wider landscape of ecological introductions.2 This
anthropocentric perspective elides some important features in the
history of biological exchange and has led to the relative neglect of
some species and regions.
Lindroth’s work deserves reexamination not only for its substance
but also for what it can tell us about the ideas behind foundational
texts like The Columbian Exchange and, indeed, about the making of
ideas about global ecological change. Lindroth’s careful ecological
and historical study pivoted on his research specialty in ground bee-
tles and focused on the case of Newfoundland, but he situated both
within much wider biological and geographical spheres. He did not
use the colorful language of his more influential contemporary
Charles Elton, but his work generated a more grounded and precise
body of ecological evidence and, in so doing, made the case for the
centrality of transportation vectors in biological exchanges and the
importance of recurring introductions of breeding pairs to sustain
510 Environmental History 26 (July 2021)
introductions—what ecologists would later refer to as propagule pres-
sure. Faunal Connections thus invites examination as a foundational
text in the history of biological exchange.
And there is no question that Lindroth set out to explain issues of
wide ecological significance: when, where, and how had European
species traveled to, and become established in, North America?3 He
approached the problem across a spectrum of species but with a keen
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eye on his specialty in ground beetle (Carabidae) taxonomy and ecol-
ogy. This focus allowed him to demonstrate conclusively that a large
number of species had been introduced to Newfoundland over the
past four hundred years and to create a framework that both identi-
fied those species and demonstrated their plausible entry points. The
chief vector for faunal introduction to Newfoundland, he argued,
had been the solid ballast carried by ships in the age of sail, which,
when dispensed on land, served as a landing ground for introduced
species. He contrasted these recent arrivals with a host of other shared
species that originated in circumpolar dispersals in earlier geological
eras. In working out this argument, Lindroth established an enduring
account of regional faunal history, demonstrated the vectors of inter-
continental biological exchange, and prefigured some major concepts
in the field of invasion ecology.
Lindroth’s Faunal Connections was researched and written at a par-
ticular moment in the postwar history of transatlantic science. The
Swedish entomologist conducted fieldwork with a team of Finnish
colleagues in Newfoundland in 1949 and again on his own in 1951,
benefiting from research funding provided by the Arctic Institute of
North America (AINA), the Rockefeller Foundation, and the Swedish
Natural Science Research Council.4 His closest intellectual collabora-
tors were Ernst Palmen, a Finnish entomologist and terrestrial arthro-
pod specialist, W. J. (Bill) Brown, a Canadian government
entomologist who worked principally on coleoptera, and Philip J.
Darlington, a Harvard zoologist and fellow carabid specialist. He also
corresponded with lecturers at Memorial University in St. Johns,
Newfoundland, hired local field assistants, and depended on the
guidance and advice of harbor officials and retired mariners in south-
west England. Like the island of Newfoundland, Lindroth’s scientific
network connected different corners of the North Atlantic.
Conceived and executed in the context of the Cold War, and benefit-
ing from governmental and institutional support that sought to pro-
mote transatlantic scientific cooperation and northern research,
Lindroth’s path to researching and writing Faunal Connections reveals
both the influence of geopolitics in shaping environmental ideas and
the capacity for scientists to benefit from strategic funding opportuni-
ties while contributing little of strategic value.
Studies of transatlantic scientific cooperation in the Cold War have
focused primarily on the physical sciences and the alignments amongStowaway Beetles 511
science policy, funding, and American foreign policy goals.
Historians of science have conducted a lively debate about the degree
to which US science policy and philanthropic foundation support
aimed to enroll European scientists into Western-oriented scientific
networks of mutual benefit and exchange.5 A related body of scholar-
ship considers how a perceived “Arctic Front” in the Cold War shaped
governmental and philanthropic funding priorities in a broadly con-
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ceived north. Driven by the need to make southern military technol-
ogies and military personnel adapt to cold environments, the
American military invested heavily in research to gain a better under-
standing of, and control over, the environment.6 This drive for north-
ern research arose not merely from American foreign policy but also
at the behest of Canada and the Nordic countries as well as several
newly minted research institutes.7
Transatlantic scientific cooperation in the Cold War circumpolar
north reflected multiple scientific and institutional relationships at
play. The AINA, for example, emerged in the late 1940s as a coopera-
tive endeavor between American and Canadian scientists, with the
clear ambition to remain independent of government, even as
American naval funding shaped much of the early research program.
Founders of the institute also sought to include participants from
Greenland and Newfoundland and identified these regions as part of
a wider north.8 When Lindroth pursued funding from the AINA,
therefore, he became enmeshed in an effort to define and analyze
northern environments for strategic ends, but he chose his own fo-
cus, built his own networks, and practiced his science without inter-
ference. In many respects, Lindroth drew on funding arising from the
Cold War to conduct research that had little if any bearing on the
conflict.
The inclusion of Lindroth’s research in funding streams that sup-
ported northern investigations speaks to the importance of imagined
geographies in science and its institutions. But place also mattered.
When Lindroth first visited Newfoundland with his Finnish col-
leagues in 1949, they entered a recently transformed political envi-
ronment, touring outport communities only a few months after the
island had joined Canadian Confederation. Newfoundland had re-
cently experienced a transformation in its external relations as a stra-
tegic site in the Second World War.9 The creation of an important
airbase at Gander, and the regular traffic of transatlantic convoys,
transformed the island’s economy after a crushing era of economic
depression and government bankruptcy. While Canadian govern-
ment and philanthropic interest in conducting research on
Newfoundland and its resources rose during the Cold War, Memorial
University in St. John’s had only just been founded, and the back-
ground of entomological collections and research on the island was
meager. Conducting research in and on Newfoundland came with its512 Environmental History 26 (July 2021)
share of logistical challenges; it also offered a major research opportu-
nity where many basic questions in ecology had yet to be addressed.
Faunal Connections appeared at a moment of rising interest in bio-
logical invasions and the agency of humans on earth. Published the
same year as Philip Darlington’s global survey Zoogeography, William
Thomas Jr.’s edited tour de force Man’s Role in Changing the Face of the
Earth, and just a year before Charles Elton’s Ecology of Invasions by
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Animals and Plants, Lindroth’s Faunal Connections epitomized a new
postwar vision of an increasingly globalized ecology.10 While much
of our understanding of ecology during the Cold War is based on the
intellectual production of metropolitan centers in the Anglo-
American world, Lindroth’s work, linking Europe and
Newfoundland, provides an opportunity to examine broader trends
from the margins of the North Atlantic.11 Lindroth’s questions dif-
fered to some degree, and the problems and experiences motivating
them proved less explicitly linked to the Cold War military power of
the United States or the United Kingdom than was the case with
many of the era’s major ecological texts.12 Lindroth did not deploy
military metaphors or elaborate a Malthusian perspective. Where
many global ecological texts of this era highlighted ruptures and cri-
ses, Lindroth emphasized long-term patterns and processes.12
Perhaps for these reasons, Lindroth’s work has not figured much in
the historiography of ecology or environmental thought.14
Whatever the reason, Lindroth’s work has also not been addressed
in the wider environmental historiography, despite its influence on
Alfred Crosby’s portmanteau biota thesis and its own significance as
a work of environmental historical interpretation. Crosby’s argu-
ment that the New and Old Worlds were connected by a set of dis-
ease, plant, and animal exchanges that set uneven terms for
encounter and profoundly shaped European imperialism in the
Americas drew on a range of ecological ideas, from the work of
Charles Elton to the cultural geography of Carl Sauer. Arguably,
none of those works pursued the ecological dimensions of faunal ex-
change as deeply as Lindroth’s nor demonstrated as clearly the vec-
tors of faunal dispersal. To be sure, Crosby cited Lindroth’s work, but
he largely glossed over Lindroth’s arguments and evidence. In doing
so, he neglected some important questions raised by Lindroth, in-
cluding the role of ballast in biological exchanges and the impor-
tance of propagule pressure. We are all, of course, indebted to
Crosby, but calling attention to Lindroth’s Faunal Connections never-
theless promises a more robust understanding of the mechanisms at
work in the Columbian exchange. Moreover, it offers a salient re-
minder that as environmental historians we need to attend not only
to species that have impacted human societies but also the broad
range of ecological introductions.Stowaway Beetles 513
MAKING CONNECTIONS
Lindroth’s research in Newfoundland pursued biogeographical
questions that had intrigued him since his doctoral research in
Iceland in the late 1920s. How had insects moved from Europe to the
islands of the North Atlantic? In Iceland, he determined that the
island’s insect fauna had arrived during the Pleistocene by a land
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bridge from the boreal regions of northwestern Europe, leading him
to question whether this biogeographical process extended further
westward across the Atlantic.15 A letter to a Newfoundland contact
suggests that he expected to find that the conclusions of his Icelandic
work applied in Newfoundland as well: “We are all interested in bio-
geography, ie the Pleistocene history of the animals and plants as
well as the possibility of former land connections between Europe
and North America.”16 But after several months of collecting in
Newfoundland in 1949, it became clear that the island was not
Iceland’s twin. As he later wrote in a typically understated way, “the
results obtained were unexpected.”17
At the time of his Newfoundland investigations, Lindroth had an
established scientific reputation but only the beginnings of an aca-
demic career. After obtaining a doctoral degree at the University of
Upsala in 1931, he spent the next decade and a half teaching high
school science. He nevertheless continued researching and, by the
1940s, had emerged as one of Sweden’s most prominent entomolo-
gists, serving as the president of the Entomological Society in
Stockholm, the Swedish Society Oikos (a scientific ecology society),
and the Society of Biology Teachers of Sweden. In 1947, he secured a
position at the University of Stockholm and, in 1951, took up a profes-
sorship at the University of Lund.18 After his appointment at
Stockholm, he discussed conducting research in Newfoundland with
Ernst Palmen of the University of Helsinki as well as Philip Darlington
at Harvard University, but he lacked the funding to execute it.19
Quite unexpectedly, the situation changed. In 1948, Palmen found
an advertisement in the Swedish language Finnish newspaper,
Hufvudstadsbladet, for research grants sponsored by a new organiza-
tion, the AINA, and enthusiastically alerted Lindroth to the opportu-
nity.20 Together with Palmen, Harry Krogerus (another
entomologist), and Risto Tuomikoski (a botanist), who were both at
the University of Helsinki, Lindroth duly applied to the institute for
funds to conduct fieldwork in Newfoundland in the summer of 1949
(see figure 1). In the adjudication process, the AINA sought out the
advice of Darlington who explained that, while he had never met
Lindroth, he had corresponded with him about scientific matters
since 1930 and judged the project to be “interesting and
important.”21 The AINA extended a grant of $6,500 to the team,
while the Swedish Natural Science Research Council provided514 Environmental History 26 (July 2021)
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Figure 1.Carl H. Lindroth, 1948. Credit: Passport photo appended to “Grant-in-Aid Application form,”
file 22, Arctic Institute of North America fonds, LAC.
additional travel funds.22 Swedish restrictions on US dollar exchange
limited Lindroth’s plan to examine the insect collection at the
Museum of Comparative Zoology at Harvard. In 1949, the focus
remained on Newfoundland, but two years later, with the support of
a Rockefeller Foundation grant, Lindroth would also visit Labrador,
Nova Scotia, Ontario, and the eastern United States.
As Lindroth planned his first trip, he sought out local authorities.
In January 1949, he wrote to Memorial University College (soon to be
renamed Memorial University) asking about insect collections and
any publications relating to Coleoptera (beetles) in Newfoundland.23
Lindroth’s correspondence was forwarded to William Rees Wright, an
Anglican clergyman from Wales who taught biology at Memorial.
Rees Wright responded regretfully that he had little information to
offer Lindroth “because very little work seems to have been done on
the Zoology and Botany of the island apart from economic and sport-
ing aspects (and in fact a very great deal of the island is quite unex-
plored scientifically).”24 Having only been in Newfoundland for a few
months himself, Rees Wright conferred with the acting president of
the university who also drew a blank before suggesting that Lindroth
might have better luck with museum collections in Britain and the
United States. Although he could offer little assistance in locating in-
sect collections, Rees Wright advised Lindroth on travel matters and
offered to publicize his research and to facilitate his work in outport
communities.25
The scientists covered great distances around Newfoundland in the
summer months, camping to keep costs down (see figure 2). After
landing in Gander in June, Lindroth and his colleagues collected in
the vicinity, then took a train to St. John’s where they worked for
over a week on the Avalon Peninsula before moving west to collect atStowaway Beetles 515
various points between Port aux Basques and Pass Island. The Western
Star published in Corner Brook reported their departure on the SS
Baccalieu in late June.26 Heading north, they split into two parties to
cover more ground, examining the island’s northwestern penin-
sula.27 In the middle of August, they returned to Corner Brook and
traveled into the interior, collecting at eight points along the railway
line. Faunal Connections includes a photograph of this leg of the jour-
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ney, near St. Fintans, showing a meadow landscape “dominated by
Marguerites (Chrysanthemum leucanthemum L.) and Red Clover
(Trifolium pretense L.).”28 All told, Lindroth and the Finnish scientists
collected specimens at seventy-five separate locations and amassed a
vast collection of spiders, moths, butterflies, beetles, and other insects
as well as mosses and vascular plants. Whereas before the expedition
seventy-five species of Carabidae had been known to exist in
Newfoundland, Lindroth estimated in his field progress report that,
when his work was completed, the number would be over 130.29
While Lindroth and colleagues encountered immigrant fauna from
the Pleistocene and earlier, they also catalogued many introduced
species. When Rees Wright heard the result, he questioned their find-
ings. Surely, he wrote, nobody in Newfoundland’s history had been
so eccentric as to introduce hundreds of ground beetle breeding pairs
“like an entomological Noah!”30
Just where had these insects come from? Lindroth learned from his
colleague Ernst Palmen that a Canadian entomologist, W. J. Brown,
had written on related matters.31 Almost a decade earlier, Brown had
published on introduced Coleoptera in eastern Canada and noted the
many locations near ports where introduced beetles had been found.
To explain this, he referenced nineteenth-century naturalists who
had identified so-called ballast plants in eastern North America or in-
troduced plants found to be growing on or about ballast piles left by
vessels travelling across the Atlantic without a paying cargo. Since
this material came from many places and plausibly contained plants,
seeds, and animals, particularly insects, Brown thought they offered
one way of explaining more or less random introductions.32 Lindroth
was intrigued. As it happened, Brown had also conducted specimen
collections in Newfoundland in 1949 as part of the Northern Insect
Survey, a research project of the Canadian Departments of
Agriculture and National Defense.33 When Lindroth contacted him,
Brown offered to share specimens and do what he could to support
his research.34
Lindroth’s second North American research trip in 1951 funded by
the Rockefeller Foundation included trips to museum collections as
well as a second collecting season in Nova Scotia, Labrador, and
Newfoundland. In January, he examined the LeConte collection at
Harvard’s Museum of Comparative Zoology as well as collections at
the Museum of Natural History in New York, the Department of516 Environmental History 26 (July 2021)
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Figure 2.Sites and routes of the Swedish-Finnish expedition, 1949. Credit: Cartography by Eric
Leinberger based on Harry Krogerus, Carl H. Lindroth, Ernst Palm en, and Risto Tuomikoski, “Field
Progress Report: Arctic Institute of North America, Project Nos. ONR-29, 30, 33, 35, Biological Studies in
Newfoundland in 1949,” Arctic Institute of North America fonds, University of Calgary Special
Collections and Archives; Carl H. Lindroth, “Field Progress Report: Arctic Institute of North America,
Project Np. ONR-30,” file 22, Arctic Institute of North America fonds, MG28-I79, 104, LAC; Lindroth to
Darlington, July 1, 1949, box 2, Darlington Papers, Harvard University Archives; institute research
reports in Arctic 3 (1950): 122–23.
Agriculture in Ottawa, and the National Museum in Washington,
DC.35 Plumbing the depths of these collections, he tried to work out
the mainland distribution of some of the species found inStowaway Beetles 517
Newfoundland and compared European and North American speci-
mens.36 Along the way, he had the opportunity to discuss his findings
with Philip Darlington at Harvard, his major interlocutor on ques-
tions of insect migration, as well as Bill Brown in Ottawa. The visit to
Harvard occurred just as Darlington was completing a major text,
Zoogeography, and initiated a close working relationship. In the fol-
lowing years, Darlington served as a patron for Lindroth, encouraging
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him to submit articles to North American journals and then manag-
ing all the editing and correspondence on his behalf; a few years later,
Darlington put in a good word for Lindroth with the publisher Wiley
and Sons and wrote the foreword for Faunal Connections.37 Lindroth
also struck up a friendly relationship with Brown that grew in the
1960s as Lindroth pursued carabid studies across Canada.38
Travelling to Newfoundland in the summer months of 1951, but
this time without his Finnish colleagues, Lindroth hired John Steele,
a Dalhousie University undergraduate from Newfoundland, to assist.
Rees Wright brokered the appointment and explained that Steele
would be the right sort, “the son of a businessman in St. John’s, and
accustomed to meeting people like ourselves”—much better, he
thought, than a “man from the outports . . . [who] would be servile or
insolent!”39 Lindroth appears not to have commented on Rees
Wright’s prejudices, though he later informed him that Steele had
worked out well.40 His summer travels took him to the north shore to
Twillingate, Fogo Island, and Hampden and, from there, by boat to
Labrador.41 For comparison, Lindroth also collected in Nova Scotia
and visited St. Pierre and Miquelon, two French islands, south of
Newfoundland in the Gulf of St. Lawrence. This was “the climax” of
the collecting season, he told Brown, even if “I failed to find any spe-
cies obviously imported from France.”42 By the end of the summer,
Lindroth reported to the Rockefeller Foundation that he had come to
the conclusion that, since the Pleistocene, introduced species could
be traced either to introductions “by shipment (ballast)” or to repre-
sentatives of circumpolar varieties (see figure 3).43
The final phase of Lindroth’s research on carabid introductions to
Newfoundland occurred in 1954 on the other side of the Atlantic. In
1950 and 1951, Lindroth had received advice on the history of ballast
from Hubert J. Squires, a Newfoundland scientist completing a degree
at McGill and based at the Newfoundland Biological Station.44
Reading into these sources, Lindroth determined that several south-
west English ports had played a significant role in the Newfoundland
fishery and corresponded with English archivists, local historians,
and port authorities to learn more. He then traveled to these ports in
the summer of 1954 where he located the historical ballast grounds.
Local historians rounded up retired sailors and mariners who
explained where they had collected ballast, when, and how—testi-
mony that Lindroth quoted almost verbatim in Faunal Connections.518 Environmental History 26 (July 2021)
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Figure 3.Lindroth’s Newfoundland collecting sites over two field seasons, 1949 and 1951. Credit:
Cartography by Eric Leinberger based on Carl Lindroth, “The Ground Beetles of Canada and Alaska,”
Opuscula Entomologica Supplementum 12 (1955): 160, unnumbered plates.
“To visit the Crow (opposite Instow),” explained one such correspon-
dent, Vernon Boyle of Westward Ho!, “you may ask for Mr. Fred
Johns the ferryman, an old sailor, who will tell you a lot. If you miss
seeing me you could call Capt. W.J. Slade, Bridgeland Street,
Bideford, an old schooner man who knows a lot about ballast.”45 At
Poole (Dorset), Topsham (North Devon), Dartmouth (South Devon),
Plymouth (South Devon), Appledore (North Devon), FremingtonStowaway Beetles 519
(North Devon), Barnstaple (North Devon), and Bristol (Somerset),
Lindroth collected insects at the former ballast places. To his delight,
Lindroth discovered that at least half the introduced species that he
had identified in Newfoundland also occurred at these sites.46 This
suggested a possible empirical foundation for what Brown had re-
ferred to in 1940 as the “Ballast theory.”47
After reporting some of this research in the early 1950s, Lindroth
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decided that the subject warranted a more general book, limited to
neither beetles nor Newfoundland but looking at the problem of
transatlantic exchange generally. Lindroth reached out to specialists
in North America and Europe as he sought to develop lists of
European species introduced to North America and vice versa. This
work depended on a vast network of expertise. Eugene Munro, a
Canadian entomologist, at the Division of Insects in the Department
of Agriculture, confirmed that “Dr. Freeman has checked the butter-
flies, Dr. Hardwick has checked Noctuidae, and I have done the
Geometridae and Bombyces.”48 I. H. H Yarrow of the British Museum
of Natural History commented on ants and hornets.49 W. J. Giertsch
of the American Museum of Natural History in New York offered ad-
vice on spiders. Philip Darlington at Harvard offered Lindroth a
lengthy set of observations comparing Lindroth’s specimens with
those held at the Museum of Comparative Zoology.50 Douglas
Ferguson of the Nova Scotia Museum of Science and a doctoral stu-
dent at Cornell offered responses to his lists of Macrolepidoptera.51
Melville Hatch at the University of Washington provided advice on
Lindroth’s questions about possible introductions in the Pacific
Northwest.52 These consultations helped Lindroth to develop a sub-
stantial list of introduced fauna and revealed the extent of his net-
work on insect taxonomy in England, Canada, and the United States,
which was not formally connected through coauthorship or funding
but which had arisen at least partly from Lindroth’s ability to travel
in North America with the AINA and Rockefeller Foundation support.
FAUNAL CONNECTIONS
Published in 1957, Faunal Connections offered a sweeping explana-
tion of faunal exchanges between Europe and North America.
Although grounded in historical research, the book was not con-
structed as a historical narrative. Rather, it offered three substantial
chapters, each very different. The first comprised a list of fauna found
in both Europe and North America, but excluding animals introduced
deliberately. In addition to agricultural animals, pets, and acclimatized
species like English sparrows, Lindroth also excluded freshwater ani-
mals and unintentionally imported animals that did not become estab-
lished. He thus focused on the “species common to Europe and North520 Environmental History 26 (July 2021)
America, and breeding (or supposed to breed) in both continents,
which are not purposely brought across by man in either direction.”53
He was, in short, interested in stowaways that became established.
Even when excluding a range of fauna in this way, the list ran to over
100 pages. Broken down by broad taxonomic groupings (Mammalia,
Aves, Insecta, Macro-Lepidoptera, Diptera, Hymenoptera, Odanata,
Araneae, Chilopoda, Diplopoda), some with as many as nineteen sub-
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sections, the list provided information on an animal’s genus, species,
and subspecies, if relevant, as well as its distribution and whether or
not it was known to be introduced. The choice to introduce a book
with a long list may seem curious, but Lindroth saw this as a crucial
foundation. The whole study turned, to some extent, on the basic map-
ping of animal distribution, classification, and the identification of
unintentional introduction. The list concluded with a few important
facts: many more species had been introduced to North America from
Europe than vice versa, and of all the introductions, the most extensive
were the Coleoptera or beetles—between 162 and 170 species or subspe-
cies that accounted for between 65 and 68 percent of the species found
on both continents.
Although the first chapter indicated known and presumed intro-
duced species, it did not define what was meant by introduction.
That he left to the second chapter, which opened with a list of five
key criteria that provided the basis for the definition, albeit without
commenting on the nature of the metaphor. The first he labeled the
historical criterion or the capacity of the biologist to determine his-
torical evidence of the introduction of a particular species. The sec-
ond he labeled the geographical criterion, and it provided, by
contrast, spatial evidence of newness, either a bounded distribution
in some coastal area near a port or an “unnatural” distribution unre-
lated to ecology, soil, and climate. The three other criteria focused on
ecological, biological, and taxonomic clues that differentiated a spe-
cies and suggested a distant point of origin. In general, Lindroth ob-
served that introduced species often inhabited distinct ecological
zones: “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.”54 He illustrated these criteria with specific
examples of species that had spread significantly over time such as
the Colorado beetle in Europe and the starling in North America
alongside other species that, for various ecological and biological rea-
sons, had a much more circumscribed distribution, including the
muskrat in Central Europe and the alfalfa weevil in the western
United States. This systematic opening, laying out the criteria for in-
troduction across different groups at a continental scale set up the
rest of the chapter’s analysis.Stowaway Beetles 521
“From where did they come?” Lindroth then asked.55 Moving from
the criteria and examples, Lindroth considered how it was that intro-
duced carabids could be found in relatively high numbers in port
locations around the island of Newfoundland yet not on the main-
land of North America to the same degree. Even more striking, the
Avalon Peninsula, the island’s settlement hub and the location of
Newfoundland’s principal city, St. John’s, stood out as the core recep-
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tion area of introduced carabids. Offering examples to ground his
claims, Lindroth reinforced his criteria and explained how each
revealed the pattern of carabid introduction. With this framework in
place, he proceeded to lay out a historical explanation, offering a brief
history of the Newfoundland trade and drawing on classic works such
as D. W. Prowse’s History of Newfoundland.56
This historical review aimed to establish one key point: that ships
had carried insects to Newfoundland from Europe with the solid bal-
last in ship’s hulls as the crucial transmission medium. He outlined
the regulations laid down in the earliest settlements of the seven-
teenth century that outlawed the dumping of ballast—the sand,
gravel, and other heavy materials that had provided stability for ships
without heavy cargo—in harbors and the reiteration of these prohibi-
tions in later phases of Newfoundland history. This meant, he under-
lined, that ships traveling in ballast had to dispose of their loads on
land, providing an entry point for stowaways. And many ships did
travel in ballast in the fishery. Since settlements attached to the fish-
ery were seasonal and then very slow to arise, most vessels traveling
west across the Atlantic did so without cargo and had to take on bal-
last. While Lindroth understood that the fishery included vessels
from various locations, he focused his investigation on the southwest
English ports that were most involved and were thus the key sources
of ballast. He discussed the sites where ballast had been collected in
these ports and their different qualities from stone to fine-grained
river sands. Quoting mariners’ accounts, Lindroth offered compelling
evidence that the transatlantic fishery had delivered massive quanti-
ties of ballast to Newfoundland over centuries. The shipping lanes in
his telling were like conveyor belts of sand, stone, and dirt across the
North Atlantic.
This material, he explained, was also alive. Earthen ballast provided
an ideal medium for plants and seeds as well as a host of hardy insect
invaders. Systematically accounting for each of the ballast sites he vis-
ited in southwest England, he reviewed the available historical record
for the port in question, the type of ballast typically used there, and
the insects and plants discovered in his survey of the historical ballast
places. He concluded by explaining that the lack of ballast carried in
the opposite direction helped to explain the low level of introduc-
tions from North America to Europe. As far as the historical exchange
of fauna, he argued, ballast held the key.522 Environmental History 26 (July 2021)
In a sharp turn from the emphasis in chapter 2 of Faunal
Connections on relatively recent introductions via ballast, the book’s
final chapter pushed the question of introductions into the deeper
past and outlined Lindroth’s answer to the question that had first ac-
tuated his research: what fauna was shared with Europe, and how
could one account for its dispersal from Europe in the Pleistocene
and earlier? He discussed a range of species shared between Europe
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and North America in a broad North Atlantic zone that he called
Amphiatlantic or Westarctic species, those that are found in north-
western Europe, but not in Asia, and extend west across the Atlantic
in places like Iceland, Greenland, and the North American mainland.
He developed two principal explanations for their dispersal. First, he
noted that many bird species could migrate across oceanic distances
under the right conditions and that many Amphiatlantic bird species
were strong flyers. This also held for Amphiatlantic Lepidoptera (but-
terflies and moths). In most cases, however, Lindroth imagined that
animals were carried across distances by the wind or on ocean cur-
rents, particularly if they were attached to floating debris. Many
plants would have traveled the same way or could have been ingested
as seeds in bird stomachs. Through such mechanisms, Pleistocene
introductions could be explained.
Lindroth concluded, however, with another look at the possibility
of a land bridge that might have connected Iceland and Greenland
and thus provided a pathway for some species that did not proceed to
mainland North America. He evaluated the faunal evidence to sup-
port such a contention and argued that the Davis Straight appeared
to have served as a barrier to the spread of some Amphiatlantic spe-
cies beyond the North Atlantic islands. In reviewing a draft manu-
script, Philip Darlington strongly disagreed with the idea of a land
bridge: “My instinct as a zoogeographer is against it.” Lindroth coun-
tered that Darlington had not addressed all the evidence that sup-
ported his position.57 Wading into the question of land bridges also
drew Lindroth into the debate over the continental drift hypothesis,
which in the mid-1950s remained unresolved.58 “The only contribu-
tion that can be delivered by a biogeographer,” he concluded on
something of an anti-climax, “is the declaration that, at least as far as
the North Atlantic area is concerned, the continental drift, if consid-
ered a reality, took place in so early a period that its biological conse-
quences cannot be traced.”59 Lindroth favored a more modest
explanation—the arrival of European species by a range of dispersal
mechanisms across an open ocean or via Pleistocene land bridges be-
tween North Atlantic islands in previous geological eras.
Faunal Connections concluded with a two-page summary, which did
not extend the previous arguments. The core of this book was to be
found in its parts: a scrupulously composed list of introduced species,Stowaway Beetles 523
an account of introductions particularly through the ballast vector,
and a review of species introduced in earlier geological times.
INTELLECTUAL CONNECTIONS
Lindroth’s book was in its own way an Amphiatlantic specimen:
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published simultaneously in Stockholm and New York, it treated a
transatlantic subject and engaged specialists in Europe and North
America. Widely reviewed, Lindroth’s work would prove to be a fun-
damental contribution to the faunal history of Newfoundland and
laid the foundation for Lindroth’s future research, which would cul-
minate in a multi-volume account of the Carabidae of Canada and
Alaska. Lindroth’s hopes that the book would build a larger audience
for his work in biogeography, however, went largely unmet. Many
reviewers commented that Faunal Connections focused primarily on
Newfoundland and carabids—as, indeed, it did—while perhaps miss-
ing the larger messages that it delivered about process and the con-
ceptualization of biological invasion.60 Several reviews of Lindroth’s
work published following his death in 1979 highlighted Faunal
Connections primarily from an entomological perspective and paid as
much, if not more, attention to his later research in Canada and
Alaska.61
The reach of Faunal Connections was also undercut by the publica-
tion of a popular book on ecological invasions by Oxford ecologist
Charles Elton that almost immediately overshadowed it. Elton’s
Ecology of Invasions by Animals and Plants had started as a British
Broadcasting Corporation radio script for a wide audience and used
colorful examples from around the world while deploying frighten-
ing language to highlight the risks. Elton devoted comparatively little
attention to mechanisms or vectors but emphasized consequences
and warned readers that they lived in a world shaped not only by nu-
clear bombs but also “ecological explosions.”62 By contrast, Lindroth
wrote carefully for a scientific audience, in precise, but uncolorful,
English. Whereas Elton chose vivid examples that captivated the
imagination, Lindroth focused mostly on species that had created lit-
tle notice even in the regions in which they had successfully colo-
nized, offered complex explanations of taxonomy and dispersal
mechanisms, and included a hundred-plus-page table as his first
chapter.
Neither Elton nor Lindroth knew of one another’s projects before
publication. As the ecologist James Carlton argues, had Elton known
of Lindroth’s careful work when he wrote the Ecology of Invasions, he
would have likely cited it and thereby provided further exposure for
Lindroth’s highly original and careful explanations.63 As it was,
Elton’s far more popular book became the key reference on biological524 Environmental History 26 (July 2021)
invasions. There is no record that Lindroth or Elton sought to engage
with one another following the publication of their books. Although
as Matthew K. Chew argues, Lindroth’s work “was in many ways
more overtly grounded in ecological explanation, historical research
and empirical natural history” than Elton’s, Elton’s work gained fame
while Lindroth’s did not. Lindroth subsequently turned to his multi-
volume study of Carabidae in Canada and Alaska, an excellent subject
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but one that was unlikely to expand his public audience.64 While
Lindroth’s work provided foundational insights for the identification
of water ballast as a vector of marine bio-invasions and continues to
be cited, it never attained the fame of Elton’s work.65
Nor, despite its historical method, did Faunal Connections gain an
audience amongst historians, excepting Alfred Crosby who paid it at
least perfunctory attention in his pioneering text, The Columbian
Exchange. Crosby cited Lindroth twice in his discussions of plant and
animal exchanges between the New and Old Worlds.66 Like many of
the initial reviewers, however, Crosby did not take up, or reflect
upon, the vectors of introduction that Lindroth discussed at length.
Rather, he cited Lindroth in support of mundane factual points.
Crosby was interested in the range of introductions to the Americas
that impacted disease ecologies and the patterns of colonialism,
whereas Lindroth focused on synanthropic species in a region where
they had little if any noticeable impact on the colonial process.
Lindroth also excluded many of the introduced species that were
central to the problems of ecological imperialism. While Lindroth
used historical methods to understand ecological change, he neither
contributed to historiography as such nor attempted to shed light on
human perceptions of the natural world. Nevertheless, Crosby
missed an opportunity to reflect on the varieties of ecological imperi-
alism through Lindroth’s observations on carabids in Newfoundland
as well as the geographical and biological conditions that made
them possible.67 As the ecologist Daniel Simberloff argued in a re-
view of Crosby’s Ecological Imperialism, by failing to address
Lindroth’s ideas about transportation vectors as a critical factor in
ecological exchange, Crosby was left to assume (indeed, imply) an in-
nate dominance in European species.68 The field sometimes inadver-
tently reiterates that conceit. Not only did Lindroth’s work
foreshadow Crosby’s, but it also continues to provide a useful
corrective.
COLD WAR CONTEXT
Faunal Connections provides a revealing angle on a process of intel-
lectual exchange crystallizing in the early Cold War. While several
historians of science have considered the transatlantic traffic in ideasStowaway Beetles 525
in these years, they have focused primarily on the associations be-
tween scientific research and geopolitics.69 This is an important con-
nection to highlight as Cold War fears of Soviet influence drove
scientific research and helped to lay the foundations for new institu-
tions like the AINA.70 Newfoundland’s entrance into Canadian
Confederation in 1949 and the repositioning of Canadian interests in
the North Atlantic were also part of this process. It is no accident that
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the research conducted by Bill Brown in Newfoundland in 1949 was
sponsored partly by the Canadian Department of National Defense or
that the AINA received a block grant from the US Naval Research
Board (NRB), which helped to underwrite the grant-in-aid program
that funded Lindroth’s research and led the AINA to deposit a copy of
the Swedish scientist’s research reports with the NRB.71 The AINA’s
support for research on Newfoundland grew out of the wartime stra-
tegic interest in the Atlantic islands as well as the broader north and
the emerging priorities of the Cold War. It paralleled efforts by both
the US and Canadian governments to develop expertise and knowl-
edge about northern environments and contributed to what the US
Navy Vice Admiral H. G. Bowen described in 1946 as the need for
baseline data.72 Operating independently, yet with the NRB as its
largest patron, the AINA pursued new knowledge of the north and
promoted international cooperation inevitably tied to the politics of
the Cold War.73
While it is difficult to imagine that Lindroth’s research would have
been funded without such connections, research on insects in
Newfoundland represented a modest contribution to strategic knowl-
edge. When Vice Admiral Bowen argued for baseline data, he had in
mind the environmental data that might impact weapons systems
and communications in the Arctic, not ecology writ large.
Newfoundland’s inclusion in the AINA’s mandate came about in part
from an inclusive spirit amongst its founders, who also made sure
that Greenland would have a place at the table. No evidence exists to
suggest that Lindroth ever considered his work within a strategic con-
text. How Newfoundland or his research might connect to global geo-
politics appears not to have preoccupied him or influenced his
research questions. Lindroth found the AINA to be a generous and
convenient funding source, and he thought the same of the
Rockefeller Foundation. He flew into and out of Newfoundland from
Gander Airport, which had recently served as a key node in the trans-
atlantic military alliance, but he hardly participated in the project
with the aim of advancing American geopolitical interests. His exam-
ple suggests that a lot of basic science happened under the guise of
Cold War funding initiatives, with direct and indirect military aims,
without contributing much, if any, strategic knowledge. “Science
during the Cold War,” as Hunter Heyck and David Kaiser argue, “took
many forms in many places, with science–state relationships varying526 Environmental History 26 (July 2021)
markedly from nation to nation, agency to agency, institute to insti-
tute, and individual to individual.”74
This is not to say that Lindroth’s work did not reflect the Cold War
context from which it emerged. It is no coincidence that Lindroth’s
work appeared alongside other notable studies taking biological inva-
sion and the human impact on the earth as their subject. Some, like
Elton’s, drew on cataclysmic Cold War imagery to underscore their
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argument; many, including Lindroth’s, drew on military funding; all
of them reflected an increasing concern with the consequences of
globalization. Apart from the comparison with Elton’s work, it is
worth remembering that an interest in biological invasion ran
through the work of Carl Sauer and his students in these years.75
Andrew Hill Clark’s study of New Zealand published in 1949, for in-
stance, took a more cultural approach than Lindroth but addressed
similar questions, again with a focus on islands and an expanding
European flora and fauna.76 Faunal Connections also appeared in the
same year as Thomas’s wide-ranging Man’s Role in Changing the Face of
the Earth, which included several chapters treating biological invasion
as a pressing new field of scientific research.77 While Lindroth
sounded less of an alarm than Elton or others on the topic of intro-
ductions, he nevertheless walked a parallel path in seeking to explain
the globalization of ecology.
Lindroth’s Faunal Connections should thus be viewed partly as a
product of these political and intellectual forces, but it was also
rooted in a host of local circumstances in both Europe and North
America. His research emerged from questions generated in Sweden
and worked out first in Iceland, then in Scandinavia, and, finally, in
Newfoundland. Newfoundland’s settlement history and the use of
ballast in the North Atlantic trade in the age of sail meant that a pro-
cess that was global but dispersed assumed on the island a concen-
trated and observable form. The absence of existing institutional
expertise in entomology in Newfoundland also provided a relatively
wide field for investigation and coincided with rising Canadian scien-
tific interest in the subject from individuals like Brown at the time of
Newfoundland’s entrance into Confederation. If Lindroth’s research
operated in the context of Cold War investments in scientific institu-
tions, it did so from a vantage point that was outside the main centers
of metropolitan science and geopolitical power.
A SIDELONG VIEW OF THE COLUMBIAN
EXCHANGE
In retrospect, it is tempting to ask what might have been the im-
pact on the field of environmental history had Lindroth’s work more
strongly shaped Alfred Crosby’s field-defining vision of theStowaway Beetles 527
Columbian exchange. While the answer is necessarily speculative,
there are at least three possibilities. Most obviously, Lindroth’s work
provided empirical evidence to clarify a long-standing theory
amongst naturalists that the solid ballast of ships carried plants and
seeds to the Americas, and he extended that theory by establishing
that many insects had been stowaways. This ballast theory provided a
foundation for explaining why some plants and animals became
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established in the Americas and why many American plants and ani-
mals did not take the voyage in reverse. Ballast and all it carried went
to American shores while little was returned to Europe. Crosby
explained the differential of introductions across the Atlantic in
terms of the characteristics of animals and plants and the ecology of
reception areas. In doing so, he sought to undermine a narrative that
assumed the superiority of European colonizers over those they colo-
nized; the European colonists proved successful in Crosby’s telling
not because of a superior culture or even technology but, rather, be-
cause of evolutionary accidents over which they had little control.
Ironically, by not fully considering the modes of those organisms’ dis-
persal, he left open the implication that European species were more
dominant than their New World counterparts.
Lindroth’s work also clarified some of the population pressures that
helped to reinforce introductions in ways that Crosby often elided.
Carabids became established in Newfoundland, he noted, not when a
single breeding pair dropped to shore. A recurring process of intro-
duction and reinforcement increased the likelihood of establishment
by an introduced species in conjunction with the ecological condi-
tions of reception areas. Later, biological invasion scientists would re-
fer to this idea as propagule pressure.78 Crosby did not consider this
process systematically, but a careful examination of trade flows and
the intensity of shipping traffic would go some distance to frame a
more geographically and ecologically precise analysis of how intro-
duced species reached the Americas and why they became established
where they did.
Finally, Lindroth’s focus on unintentional introductions of mainly
synanthropic species raises questions about Crosby’s focus on
microbes, animals, and plants that shaped the conquests as well as
the New and Old World demographic processes. While Crosby radi-
cally decentered a political and military narrative of discovery and
conquest, he retained an anthropocentric valuation of which
microbes, plants, and animals crossing the Atlantic mattered.
Lindroth, by contrast, considered the insect species that he discov-
ered in fieldwork and then sought to extend his analysis by including
all unintentional faunal introductions. His focus on beetles might
seem arcane, but beetles do make up a quarter of all living species on
the earth and account for a majority of the introduced species across
the Atlantic. Had Crosby evaluated a wider range of species, the force528 Environmental History 26 (July 2021)
and focus of his arguments may have been diluted. But he also may
have offered a more systematic and less anthropocentric account of
ecological introductions and how they both shaped and did not
shape colonial processes.
Of course, Crosby’s work ranged widely and addressed a host of
issues that Lindroth did not. The Swedish scientist had nothing to say
about the annihilation of indigenous Beothuk in Newfoundland even
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as he analyzed the faunal consequences of colonialism. He ignored
beasts of burden, had little to say about flora, and nothing to say
about microbes. Crosby’s vision of exchange was broad and expansive
and trampled productively across traditional intellectual divisions.
Lindroth’s also challenged the bounds of ecological and historical re-
search but within more circumscribed boundaries, for the better in
terms of precision and probably for the worse in terms of visibility
and reception. The point is not to claim that Lindroth’s work held
some key that Crosby failed to deploy but, rather, to flag a trail not
taken and to underline the ways in which Lindroth’s work continues
to merit close reading and reflection.
Matthew Evenden is a Professor of Geography and Associate Vice-
President of Research & Innovation at the University of British Columbia
Notes
My thanks to the archivists at the University of Lund who guided me through the
Lindroth papers. The vast majority of Carl Lindroth’s correspondence regarding his
North American research is in English. For the Swedish correspondence between
Lindroth and Palmen, I thank Silva Kraal for offering translation support as well as
ABC language solutions translation service. Thanks to Maribeth Murray, executive di-
rector of the Arctic Institute of North America, for granting permission to examine
the Arctic Institute of North America (AINA) files held by Library and Archives
Canada (LAC) as well as archivists at LAC, the Rockefeller Foundation, Harvard
University, and the University of Calgary. Thanks also to Peder Roberts for sharing
insights about the AINA. Since this article delves into various entomological and eco-
logical matters, I benefited from the advice of my sister, Maya Evenden, an entomol-
ogist at the University of Alberta, as well as some of her colleagues whom she
introduced me to, especially John Acorn (University of Alberta) and David Larson
(Memorial University). Their comments on a draft provided excellent additional
insights, as did the comments from the journal’s peer reviewers. Environmental
History editors Stephen Brain and Mark Hersey offered excellent criticism and guid-
ance along the way. Thanks to Eric Leinberger for preparing the maps. I am also
pleased to acknowledge support for this research from the University of British
Columbia.
1 Carl H. Lindroth, The Faunal Connections between Europe and North America
(Stockholm: Almqvist & Wiksell; New York: John Wiley & Sons, 1957), 147.
2 John R. McNeill, “Biological Exchange in Global Environmental History,” in A
Companion to Global Environmental History, ed. John R. McNeill and Erin Stewart
Mauldin (Chichester: Wiley-Blackwell, 2012), 433–51; Alfred Crosby, The
Columbian Exchange: Biological and Cultural Consequences of 1492 (Westport:
Greenwood, 1973).You can also read
