the enigma of guanacos in the falkland islands: the legacy of john hamilton
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ORIGINALARTICLE
The enigma of guanacos in the FalklandIslands: the legacy of John Hamilton
William L. Franklin1* and Melissa M. Grigione2
1Department of Natural Resource Ecology and
Management, Iowa State University, Ames, IA,
USA and 2Department of Environmental
Science and Policy, University of South Florida,
Tampa, FL, USA
*Correspondence: William L. Franklin, 404 8th
Street N. W., Nora Springs, IA 50457, USA.
E-mail: wlf@omnitelcom.com
ABSTRACT
Aim To address the biogeographical enigma of why guanacos (Lama guanicoe)
are in the Falkland Islands we investigated the following questions: (1) What was
the origin of the introduced guanacos? (2) What were the initial population sizes?
(3) Why are they found only on one island? and (4) Who was John Hamilton and
what role did he play?
Location The Falkland Islands are located in the South Atlantic Ocean 600 km
east of Patagonia at the southern end of South America. While dominated by East
and West Falkland Islands, the archipelago is composed of some 750 islands.
Sedge and Staats Islands, two small outlying islands of West Falkland, are the
focus of this paper.
Methods Historical information was collected from known relevant documents
housed at the Falkland Islands Government Archives in Stanley, and personal
interviews conducted with past and present residents of West Falklands. Research
expeditions were made to Staats Island in 1999, 2002 and 2003 to assess the
guanaco population size, distribution and social structure.
Results Guanacos were unsuccessfully introduced in 1862 to East Falkland south
of Mt Pleasant where Prince Alfred hunted them in 1871. John Hamilton, Scottish
immigrant to the Falklands and Patagonia of southern Argentina and Chile, was the
driving force in the introduction of guanacos from the region of Rio Gallegos,
Argentina during the 1930s. The guanaco was one of several wildlife species he
introduced, however, only the guanaco, Patagonia grey fox (Dusicyon griseus) and
perhaps the sea otter (Lutra felina) survive. Hamilton’s acting agent, Jimmy Miller,
imported four shipments totalling 26 guanacos from 1934 to 1939. In 1934 the
Falkland Government authorized Miller to introduce guanacos to Sedge Island, all
11 of which disappeared. Whether intentional or accidental, 15 guanacos were
taken to Staats Island, an islet of 500 ha on the western edge of the archipelago.
Historically, guanacos are unexpected on Staats Island because documentation
authorizing their introduction is unknown. Guanaco numbers have fluctuated
widely on Staats Island for 65 years primarily due to culling. In 1959 the population
was dangerously close to extirpation, but today 400 thrive there. A severely reduced
gene pool and genetic bottlenecking were suggested by recent field studies,
revealing preliminary evidence of deleterious consequences of inbreeding.
Main conclusions John Hamilton, spirited and visionary Scottish immigrant to
the Falklands in the early 1880s, was responsible for the introduction of guanacos
into the Falkland Islands. While there are some gaps in the historical events, the
enigma of how and why guanacos were introduced to a single island in the South
Atlantic Ocean is understood. Today, Staats Island, as a closed system, is a rare
natural experiment in progress. It offers unique opportunities for addressing
advanced questions in ungulate population, behavioural and genetic ecology. The
population potentially also represents breeding stock for farming the guanaco’s
Journal of Biogeography (J. Biogeogr.) (2005) 32, 661–675
ª 2005 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi doi:10.1111/j.1365-2699.2004.01220.x 661
INTRODUCTION
The guanaco (Lama guanicoe) (Fig. 1), one of two wild
Neotropical camelids and progenitor of the domestic llama
(Lama glama), is the dominant aridland ungulate of South
America (Prichard, 1902; Franklin, 1983). It is found in a wide
variety of habitats from sea level to 4000 m, including deserts,
shrublands, grasslands, clumped savannahs and forests (Miller
et al., 1973; Franklin, 1982). With as many as four subspecies,
the guanaco’s distribution spans from the dry west-facing
slopes of the Andes in northern Peru down the coast to central
Chile, the arid east slopes of the southern Andes, across the
Patagonian foothills and plains, and onto the austral islands of
Tierra del Fuego and Navarino (Miller et al., 1973; Franklin,
1975, 1983).
On the Patagonian steppe the guanaco was historically the
dominant and most common large mammalian herbivore
(Prichard, 1902; Franklin, 1982). When Darwin (1845) visited
South America in the early 1830s he noted ‘The guanaco… an
elegant animal… is the characteristic quadruped of the plains
of Patagonia… It is very common over the whole of the
temperate parts of the continent’. Based upon twentieth
century stocking rates, there were an estimated 30–50 million
guanacos when the Europeans first arrived to the southern
cone of South America in the late 1500s (Raedeke, 1979).
The guanaco’s wide and successful distribution has been
made possible by its flexible social organization and adaptable
ecology. Some populations are sedentary and others migratory,
and its versatile foraging strategies include being both grazers
(grasses and forbs) and browsers (shrubs and trees) (Franklin,
1982). Habitation of dry environments is surely related to its
ability to go for long periods without drinking water when
forage moisture is sufficient, and its observed capacity to drink
brackish and saline water, including water from ocean surf and
tide pools (Musters, 1871; Franklin, 1983).
How important is the guanaco to the mammalian fauna of
the Neotropical Region? Despite the region’s high mammalian
diversity (second only to the Ethiopian-African Region) and its
highly valuable wool on other islands. Thus, among his many efforts to practice
land stewardship and promote economic diversity through the introduction of
Patagonian wildlife, a remaining legacy of John Hamilton to the Falkland Islands
is unmistakably the guanacos of Staats Island.
Keywords
Falkland Islands, genetic bottlenecking, Guanaco, introduced species, John
Hamilton, Lama guanicoe, Patagonia wildlife, Staats Island.
Figure 1 Family group of guanacos composed of an adult male, numerous adult females, and chulengos (young < 1 year) at Torres del
Paine National Park, Chile during the austral summer.
W. L. Franklin and M. M. Grigione
662 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
highest proportion (43%) of endemic mammalian families
among all zoogeographical regions of the world, South
America is impoverished in terms of hoofed mammals. Both
South American and African regions feature high climatic and
ecological biodiversity from stark deserts to dense tropical
jungles, yet, South America has only 19 species of wild
ungulates compared with 95 in Africa (Franklin, 1983; Nowak,
1999; Prothero & Schoch, 2002). On the high plateau, cold
deserts of Patagonia in southern Chile, the guanaco is the only
wild ungulate (Johnson et al., 1990).
Guanacos have not only played a notable role in the ecology
and biogeography of South America, but in the cultural and
social histories of this ungulate-poor continent. Indigenous
and subsistence peoples of the Patagonia used guanacos for
food, clothing and shelter, and inculcated them into religious
beliefs and ceremonies (Bridges, 1950; Franklin, 1982; McEwan
et al., 1997). The Selk’nams (Onas), a guanaco-dependent
culture and the southernmost inland tribe in the world,
survived the harshness and extremes of Tierra del Fuego
because of the presence of this wild camelid. With the arrival of
Europeans and human occupation of the steppes of southern
South America, guanaco numbers severely declined due to
overhunting and competition with livestock (Franklin et al.,
1997). The guanaco was viewed as an unwelcome competitor
for the limited forage of this aridland and newly pioneered
corner of the world.
In the late 1800s and early 1900s the recently established
sheep industry was gaining momentum as the economic base
of Patagonia and the Falkland Islands, an archipelago off the
east coast of southern South America. It was also a time when
whalers, sealers and penguin hunters were ravaging the wildlife
of the region, and conservation was a new concept in a land
where terrestrial and marine wildlife was viewed as unlimited,
if not expendable. It was at this time that John Hamilton, a
progressively minded Scottish emigrant to Patagonia and the
Falkland Islands, ventured onto the scene. His novel ideas were
to have both short- and long-term effects on the islands
(Bernhardson, 1988, 1989).
Today, hundreds of miles from South America, on a small
island in the Falkland Islands, there is a thriving population of
guanacos (Strange, 1972; Franklin, 2003, 2004). The enigma is
how they got there. Why are they on this island? What role did
the pioneer John Hamilton and his colleagues play? Thus the
biogeographical objectives of this paper were to clarify the
origin, numbers, and islands where guanacos were introduced
into the Falkland Islands. If the original gene pool was small, it
Figure 2 The southern tip of South America
and important locations associated with the
exportation of guanacos in the 1930s to the
Falkland Islands 600 km east of the Patagonia
coast.
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 663
could have important implications for the genetic diversity and
health of the guanacos surviving today.
METHODS AND STUDY AREA
A search was made of historical records housed at the Falkland
Islands Government Archives in Stanley, including personal
correspondences, government memoranda, government stock
returns, Colonial Secretariat livestock files and shipping
records. In addition, personal interviews were made with past
and present residents of West Falklands. Strange (1992) was
used for the nomenclature reference for all taxa mentioned.
The Falkland Islands are located in the South Atlantic Ocean
600 km east of the South American mainland and of the
Patagonian desert at the far southern end of South America
(Fig. 2). East and West Falkland Islands are the two main large
islands, surrounded by an archipelago of some 750 islands
(Woods, 2002). Sedge and Staats Islands, two small outlying
islands of West Falkland, are the focus of this paper (Fig. 3).
Three research expeditions were made to Staats Island:
14–23 November 1999, 9–18 December 2002 and 7–17
December 2003, during which counts and surveys were made
of the guanaco population found there (Franklin, 2003, 2004).
In addition, historical inscriptions and writings found on the
walls of Waldron’s Shanty on Staats Island were transcribed
(Franklin, unpubl. Shanty data).
A relatively small (500 ha) island of the Weddell and Beaver
Island group, Staats Island, is at the far western edge (51�53¢ S
latitude and 61�11¢ W longitude) of West Falkland (Fig. 3).
The south-west end of Staats is a large plateau surrounded by a
coastline of formidable cliffs, with the balance of the island
dominated by a series of distinct domed peaks of up to 140 m,
separated by steep slopes and narrow valleys of short grass and
forb meadows (locally called ‘greens’ and vegas in Spanish on
the mainland) (Kerr & McAdam, 2003) (Fig. 4). Oceanic
Heath and grass are the island’s dominant plant formations.
A serious loss of biodiversity occurred when the once
abundant and ecologically important stands of tussac grass
(Poa flabellata) were destroyed by fire and overgrazing by
historically present sheep and more recently guanacos (Moore,
1968; Strange, 1989, 1992; Kerr & McAdam, 2003).
RESULTS AND INTERPRETATIONS
Early history
The introduction of guanacos into the Falkland Island has long
been contemplated. Lever (1985) in his review of naturalized
mammals of the world, reported that in the mid-1840s the first
appointed governor of the Falklands, Richard C. Moody, noted
the similarity of vegetation and climate between the islands
and the eastern Patagonia. Moody suggested because of the
value of its wool and skins, that the introduction of guanacos
into the Falklands could be ‘worthy of the alternative
consideration of the enterprising settler’. Over the next
100 years two major efforts were made to do so.
The first introduction of guanacos was in 1862, to East
Falkland (Lever, 1985), in the area just south of today’s
Mt Pleasant International Airport. Nine years later in February
1871 HRH Prince Alfred, Duke of Edinburgh, while in
command of the non-combat H.M.S. Galatea, made the first
royal visit to the Falkland Islands (Smith, 1973). A dispatch
written by then Governor George D’Arcy to the Secretary of
State for the Colonies states: ‘On Monday Captain Packe
organized an Expedition for the Prince to Mare Har-
bour, 50 miles distant from the Settlement [Stanley] to the
southward, here his Royal Highness remained till the following
Wednesday evening shooting Guanacos, wild Geese, Duck and
Snipe, etc.’ (FIA:CS, 1871). The tenure of these guanacos was
short and most likely they were shot out before becoming
established. No further records are available and none survive
today. The second and lasting introduction of guanacos to the
Falkland Islands was by John Hamilton (Strange, 1972) and his
associates in the late 1930s.
The legacy of John Hamilton
John Hamilton was a young man in his early 20s when he
immigrated to the Falkland Islands from Wick, Scotland, in the
early 1880s. Frustrated with the unavailability of land to pursue
his dreams of owning his own sheep farm, he turned to the
mainland in 1885 and pioneered two large grazing ranches
(estancias) at Punta Loyola east of Rio Gallegos, then a part of
Chile. A man of determination, over a 2-year period, he and
four Scottish countryman herded several thousand sheep and
horses 2400 km from the province of Buenos Aires to Rio
Gallegos (Mainwaring, 1983).
Eventually he and two fellow Falkland’s emigrants, brothers
Thomas and William Saunders (J. Cameron, pers. comm.),
formed a partnership and amassed a successful company in
southern Chile based upon agriculture, mining, and the
ranching of four Patagonian estancias. Concurrent to these
large Patagonian holdings and activities, Hamilton kept a close
eye on the Falklands, with the possibility of acquiring land and
returning. As the opportunities arose from the early 1920s to
the mid-1930s, Hamilton purchased Beaver Island and adja-
cent islets (including Staats Island in March 1922), Weddell
Island, Passage Islands, and Saunders Island, all totalling over
41,700 ha (FIA:CS, 1921, 1923, 1928a, 1931). Hamilton,
however, was not a land speculator, but had a forward vision
of land stewardship through rational stocking rates and
potential economic diversification of the Falklands through
the introduction of naturalized wildlife from Patagonia
(Bernhardson, 1988).
Soon after acquiring land in the West Falklands, Hamilton
undertook a number of projects, including the drastic reduc-
tion of sheep stocking rates on Weddell Island to 15% of
previous levels that had caused serious and destructive
overgrazing. It is probable that he also initiated the uncommon
practice of fencing the land to allow pasture resting and
rotational grazing (Bernhardson, 1988). He began restoration
replanting of native tussac grass, which had formerly fringed
W. L. Franklin and M. M. Grigione
664 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
the littoral zones of larger islands and nearly covered many
small ones (FIA:CS, 1934a). Tussac grass is an important
nesting habitat for penguins and other birds and birthing
grounds for some seal species (Hoppe & McAdam, 1992;
Strange, 1992), but uncontrolled grazing and fires set by
sealers, hog and penguin hunters had eliminated it from much
of its former range by the 1920s.
Early upon his return to the Falklands, Hamilton contem-
plated afforestation of the Falklands. He introduced conifers
and a variety of cypress species, of which the exotic Cupressus
macrocarpa is one of the most widely planted windbreaks and
ornamentals in the Falklands today (FIA:CS, 1922). In an
attempt towards economic diversification, Hamilton also
experimented with harvesting sea lions (Otaria flavescens) for
their oil (to be used by industry in the United Kingdom), but
discontinued when the yield was too low. In 1923 Hamilton
wrote… ‘the male seals were so low in condition that it seemed
a useless destruction to continue any longer’ (FIA:CS, 1926a;
Bernhardson, 1988).
Hamilton crusaded for the introduction of exotic animals
from the South American Patagonia to diversify the Falkland’s
economy. He oversaw the stocking of small islands, such as
Staats and Sedge, with naturalized species that in some cases
filled unoccupied niches. For example, in 1926 Hamilton and
Reginald Pole-Evans, manager of the large J. L. Waldron farm
at Port Howard, secured permission from the Falkland
Figure 4 Low elevation, composite-aerial photo of the 500 ha Staats Island looking north-west, where a population of guanacos has
survived 65 years after their introduction in the late 1930s.
Figure 3 The Falkland Island archipelago in the South Atlantic Ocean, dominated by East and West Falkland Islands, contains 750 islands
and covers 12,000 km2. Locations associated with the importation of guanacos in the 1860s and 1930s are identified. Especially note Sedge
and Staats Islands.
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 665
Government to import Patagonian or grey foxes (Dusicyon
griseus) onto Staats and Tea Islands, owned by Hamilton, and
onto River Island, owned by Waldron; the programme first
released five foxes onto Staats Island in 1928 (FIA:CS, 1928b).
Grey foxes were not considered a threat at the time to the
established sheep industry, but a prospective control for the
grazing upland goose (Cholephaga picta) competing for forage
with sheep and destroying newly planted tussac grass seedlings.
Foxes were especially seen as a potential minor fur industry
(FIA:CS, 1924, 1926b), but shipments to London proved to be
unprofitable (R. McGill, pers. comm.). While the grey fox is an
opportunistic feeder, especially on insects (Johnson &
Franklin, 1994), they have had a serious impact on ground
nesting birds in the Falklands (Strange, 1972; S. Poncet, pers.
comm.) and have taken up to 30% of newborn lambs on
Beaver and Weddell Islands (Strange, 1992). The government
naturalist wrote in 1940 to the Colonial Secretary: ‘Foxes and
skunks are both carnivores and I am deeply concerned at the
destruction of native fauna which most inevitably result from
the presence of these creatures. I am informed that the
liberation of foxes on Weddell Id. has been disastrous not only
as regards the birds but also as regards the sheep, lambs in
particular’ (FIA:CS, 1940). Today grey foxes are found on
Staats, Tea, Weddell, Beaver, Split and River Islands and are
considered a pest by most (Strange, 1972, 1992).
In addition to foxes, Hamilton introduced into the Falkland
Islands a myriad of Patagonian wildlife species: Austral
parakeets or parrots (Enicgnathus ferrugineus), buff-necked
Ibis (Theristicus melanopis), lesser rheas (Pterocnemia pennata)
(incorrectly called ‘ostriches’ by local residents), hog-nosed
skunks (Conepatus humboldtii), sea otters (Lutra felina), and
guanacos. Rheas (W. Davis, pers. comm.; R. McGill, pers.
comm.), a male and female, and several skunks were released
on Staats Island (W. Davis, pers. comm.). Davis was told of the
rheas and skunks introduced to Staats by Auto Ripp, a German
who spent most of his life in the Falklands after being ship-
wrecked there as a young man before World War II, and who
was a crew member on Hamilton’s boat the Penelope when the
rheas and skunks were dropped off. Bill Davis (pers. comm.)
remembers that during a 1956 visit to Staats, distinctly
smelling but not seeing skunks, and Rodeny Napier (pers.
comm.) recalls that ‘Rheas were placed on Staats Is. and did
well for a few years, but the foxes molested their nests’.
From his native Scotland, Hamilton also imported Shetland
ponies and highland cattle. In addition to guanacos, during the
late-1930s the Falkland Islands Stock Reports listed as imports
three skunks, 23 ‘ostriches’, 28 ibises and 16 foxes (FIA:CS,
1934–39), but none from 1928 to 1933 (FIA:CS, 1928–33).
From all these attempted introductions, only foxes, guanacos
and rare sightings of otters remain today (Strange, 1972, 1992;
Bernhardson, 1988). The Falkland Government acknowledged
Hamilton’s experience with and concern for wild fauna when
he was described by the Secretary of State as ‘a public-spirited
gentleman who has the welfare of the Colony very much at
heart. He is moreover keenly interested in natural life…’
(FIA:CS, 1930).
Unravelling the Falkland guanaco puzzle
Hamilton’s acting agent and manager of his Weddell Island
farm, James (Jimmy) W. Miller, frequently travelled between
Punta Arenas, Rio Gallegos and the Falklands. He handled a
great deal of the livestock importation for Hamilton. On
21 June 1934, Montagu Craigie-Halkett, Acting Colonial
Secretary, wrote to J. W. Miller (FIA:CS, 1934b): ‘Sir, With
reference to your letter dated 18th June, 1934, I have the
honour to inform you that permission is granted to land on
Sedge and Wreck Islands a number of Guanacos, Skunks
and Ostriches’ (see Fig. 3). What ensued can be pieced
together based upon surviving documents and what we
know today.
In the Government’s annual survey of livestock and animals
in the late 1930s, Stock Reports stated that four imports of
‘guanacho’ and guanaco totalling 26 animals were made into
the Falklands (FIA:CS, 1934–39; Table 1). Unfortunately,
specific dates, landing sites, and names of the importers are
not given. Ships from the mainland carrying cargo for camps
(farms) in the West Falkland stopped at Port Stephens, Port
Howard and Fox Bay. Ships carrying livestock typically
stopped at Fox Bay first (J. Cameron, pers. comm.). A revealing
shipping register for Fox Bay (FIA:CS, 1914) recorded that the
Chilean S. S. Patagonia was ‘Whether Bound’ for Port Howard,
Sedge Island and Weddell Island on 30 March 1936. The
175-ton ship with a crew of 22 was captained by J. Gomez and
had originated in Magallanes (Punta Arenas) and Rio Gallegos.
It carried general cargo, 35 mares and six guanacos. This
matches the imported six guanacos reported in the 1935/36
Government Stock Report. Apparently after leaving Fox Bay,
the S.S. Patagonia went north to Port Howard and Sedge first.
Two weeks after leaving Fox Bay a short note appeared in the
Stanley ‘Penguin’ news (FIA:CS, 1936a): ‘The s.s. ‘‘Patagonia’’
left Fox Bay for Weddell Island at 6 a.m. on Sunday, having
run into Fox Bay on Saturday owing to the terrific sea outside
caused by a southerly gale’.
Unfortunately the detailed cargo record for the S. S.
Patagonia was an exception. Earlier records often gave remarks
concerning noteworthy passengers, circumstances, or cargo,
but by the 1930s the normal comment was only ‘With Cargo’
Table 1 Ships confirmed* (FIA:CS, 1914) and suspected to have
transported guanacos from Rio Gallegos in southern Patagonia
to Fox Bay and beyond in the West Falklands in the 1930s.
Guanacos are documented numbers from Falkland Government
Stock Returns reporting imported animals (FIA:CS, 1934–39 and
FIA:CS, 1932–47). See text for references and documentation
explaining islands where guanacos were taken
Arrival Fox Bay Ship/Registered/Tons/Crew/Captain Guanacos
19 October 1934 Lovart/Chilean/268/24/J. Gomez 5 to Sedge
30 March 1936 *Patagonia/Chilean/175/22/J. Gomez 6 to Sedge
24 April 1938 Pilar/Chilean/354/27/J. Gomez 5 to Staats
19 May 1939 Pilar/Chilean/354/27/J. Gomez 10 to Staats
W. L. Franklin and M. M. Grigione
666 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
or ‘In Ballast’ (J. Cameron, pers. comm.). The best clues for
reconstructing the importation of guanacos into the Falklands
is by searching for ships that stopped or originated in Rio
Gallegos because of its close proximity to Hamilton’s estancias
in Punta Loyola, and that were destined for Fox Bay.
Wild adult guanacos are very difficult to live capture and
transport, even with modern immobilization drugs (Sarno
et al., 1996). Hand-raised guanaco juveniles, however, are
manageable. Newborn guanacos (chulengos) can be hand-
captured (Franklin & Johnson, 1994) and bottle-fed until
weaning at 4–6 months of age (Franklin, 1981; Garay et al.,
1995; Sarno & Franklin, 1999a). At this southerly latitude on
the South American mainland, the guanaco birthing season is
in early summer from late-November to early December
(Franklin & Johnson, 1994; Gustafson et al., 1998), making the
favourable window for transporting weaned guanacos in the
austral autumn from March to May. Such was the timing of
the six guanacos shipped in March 1936, 4 months after the
guanaco birth season. Following this scenario, chulengos hand-
captured, bottle-fed and weaned at Hamilton’s Punta Loyola
estancia, would have been loaded in Rio Gallegos and fed cut
pasture while en route to the West Falklands.
By examining the 25 ships (FIA:CS, 1932–47) that crossed
from Rio Gallegos to Fox Bay in the late 1930s and matching
for autumn shipping, two other cargo vessels emerged as the
likely ships that made the subsequent importations (Table 1).
The first import of guanacos in 1934, however, is believed to
have been an exception to the autumn pattern since author-
ization was not granted until 21 June, making a winter
introduction too risky, forcing postponement until the
following spring in October. Note that all four ships were
captained by the same master (Table 1). During this period, it
was not uncommon in this region for sea captains to sail on
more than one ship. This would have enabled the one captain
to oversee the shipments of such unusual ‘livestock’ and be
experienced in piloting the precarious seas and channels of the
West Falklands.
Hamilton and Miller apparently started as early as the 1920s
in preparation for importing Patagonian wildlife to the
Falklands. Trevor Halliday (pers. comm.), who was born in
Port Stephens in 1939 and lived in Stanley until 1961, recalls:
‘My father, Andrew John Halliday, a contract sheep shearer
from the Falklands, worked in the Patagonia from 1919 to
1929 during the off season for large company estancias, such as
Pali-Ike, in southern Chile and Argentina near Rio Gallegos.
This was the area that the guanacos were caught for the
Falkland Island breed stock. It is my understanding that the
young guanacos were the easiest to catch. I cannot recall him
saying that he ever worked for or was employed by Hamilton’s
estancias. But he, along with family cousins and friends shot
foxes, guanacos, and the big prize, pumas. There was a fair
winter trade going on at the time in pelts and feathers, always
done in the off shearing times to supplement their income.
Hamilton’s agent, James (Jimmy) Miller, a Falkland Islander
working out of Punta Arenas, and any trade that was to the
Falkland Islands from Argentina had to go through Chile, due
to Argentine political problems. In the off season of 1923 and
1924, he and fellow islander Mr William ‘Chico’ Frazer
collected Guanacos, foxes and ostriches to take to the Falklands
for shipment by Jimmy Miller. The Guanacos were caught at
the place where they worked and as far I can recall him, he told
me there were 28 in all. These were on consignment to
Mr John Hamilton and Mr George Scott’.
Andrew John Halliday was born on Dyke Island (south of
Weddell Island) in 1899 and resided on West Falkland until
1963. He would have known John Hamilton (J. Halliday, pers.
comm.). George Scott was born in 1886, the son of a shepherd
at Chartres, a large farm on West Falkland. He went to work
on New Island just north of Staats Island and in 1909 married
Fanny Cull, a widow who lived there and bought the island
that same year. Presumably, George Scott would have had a
good deal of contact with John Hamilton as they were such
close island neighbours (J. Cameron, pers. comm.), but just
how he was involved in the guanaco importations is unknown.
While the number of guanacos (28) recalled by Mr Trevor
Halliday is close to the number ultimately exported to the
Falklands (26), it is improbable that these were the same
animals taken to Sedge and Staats Islands 12–13 years later, yet
not impossible, since wild guanacos reproduce into their 20s
(Franklin, unpubl. data).
Sedge Island guanacos: authorized and disappeared
Today there are no guanacos on either Sedge or Wreck Islands
(Strange, 1992), where they were authorized to be introduced.
But were they ever there? There is no indication that guanacos
were put on Wreck, a small rocky island with little to no forage
(Pole-Evans, pers. comm.). The evidence, however, indicates
that they were on Sedge Island. In 1936, after the first two
importations had placed 11 guanacos on Sedge, an internal
government memo from the Colonial Secretary (Montagu
Craigie-Halkett), acknowledges the introductions: ‘I see no
objections to permission being granted In 436/30 attached sheet
8, Mr Miller was given permission to land on Sedge Is.
Guanacos and Skunks – about 15 of the former were put on this
Island also a number of the latter. The Island has not since been
visited so no information is available as to how the animals are
progressing.’ (FIA:CS, 1936b). Then soon after the fourth and
final introduction was made to the Falklands, the government
naturalist wrote in 1940 to the Colonial Secretary: ‘I would
bring to your notice the matter of the introduction of assorted
wild animals by Mr James Miller. So far as I am aware he has
introduced, on Sedge Island guanacos, rheas, foxes and skunks
and Weddell Island foxes’ (FIA:CS, 1940).
In addition to government correspondence, other evidence
that guanacos were imported to Sedge Island comes from
personal interviews. While Roderick Napier (pers. comm.) of
West Point Island is ‘certain guanacos were never put there’ on
Sedge, other personal experiences support the idea that they
were. Wally (now deceased) and Phyllis MacBeth (pers.
comm.) and their family farmed sheep and lived on Sedge
Island from 1966 to 1980 and worked the island seasonally for
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 667
the seven previous years from nearby Carcass Island. Phyllis
and her two children (Raymond ‘Dugal’ MacBeth and Rowena
MacBeth Summers) all remember being told that there had
been guanacos, skunks and ‘ostriches’ on Sedge Island. ‘They
were put there by Jimmy Miller’ Phyllis recalls being told by
Cecil Bertrand (born about 1910, who was alive when
guanacos were put on Sedge; resident and farmer on Carcass
Island 1953 to 1982, J. Cameron, pers. comm.). The MacBeths
also found long bones and part of a skull that they believed to
be from guanacos. When they first arrived on Sedge in the late
1950s the only inhabits were foxes and wild cattle. One of the
first things Wally (assisted by Ted Robson) did was to shoot all
the 72–74 cattle. Three years later after giving the tussock grass
some time to recover, the first sheep (300) were introduced to
the island. It took them nearly 15 years to shoot and trap
more than 300 foxes before eliminating them from Sedge
(R. MacBeth, pers. comm.).
Tony Pole-Evans (pers. comm.), resident of nearby Saunders
Island for 55 years, also remembers when he was 10–11 years-
old being told by his father that ‘Jimmy Miller from Punta
Arenas brought guanacos, foxes, ostriches and skunks to Sedge
Island’ and that the boat Pilar was still bringing cargo to West
Falkland when he was a boy. Tony Pole-Evans’ father was
Reginald Pole-Evans, close business associate of John Hamil-
ton with whom foxes were introduced into the Falklands.
Trevor Halliday (pers. comm.) also recalls of talk by people in
Stanley that guanacos had been authorized to be put on Sedge
and Wreck Islands.
Of the 26 guanacos introduced in the Falklands, it appears
that the first two shipments totalling 11 animals were taken to
Sedge. What happened to them? Tony Pole-Evans (pers.
comm.) recalled that ‘The guanacos were put on Sedge and left
there. People rarely visited those outer islands in those days.
I have no idea of what happened to them’. It is probable that
the guanacos died off naturally before becoming established.
Or perhaps someone shot them for sport or did not want
guanacos in the Falkland Islands. As for the other introduced
wildlife, Napier (pers. comm.) recounts that ‘the skunks were
put on and never seen again, and of the 4–5 ostriches the last
one was seen in 1948’.
Staats Island guanacos: unauthorized? and flourished
There is a long oral history that Hamilton also put guanacos
on Staats Island (Strange, 1972), but the details of that
introduction are an ambiguous part of the Falkland guanaco
story. The presence of guanacos on Staats is historically
unexpected, as there is yet no known documentary evidence
authorizing their introduction. Permission granted to land
animals on one island, but placed on another, would likely not
occur without considerable ensuing correspondence with the
Falkland Government (J. Cameron, pers. comm.). In the 1930s
and 1940s there was a serious debate going on regarding the
consequences of introducing new species, with the Govern-
ment taking a cautious stance and exercising as much control
as possible (Bernhardson, 1988). Unfortunately, much of the
pre-1959 historical documentation and material were lost or
damaged in the March 1959 Falkland Government Secretariat
fire (J. Cameron, pers. comm.).
Yet, in spite of the lack of known historical documentation
addressing the question, we believe that the last two shipments
of guanacos in 1938 and 1939 totalling 15 animals were left on
Staats Island. The first two shipments of 11 animals firmly
appear to have been taken to Sedge based upon the correspon-
dence of the Colonial Secretary, Montagu Craigie-Halkett, when
he stated ‘about 15 of the former [guanacos] were put on this
Island’, as well as the testimonies of people who lived on Sedge
and with living individuals whose family had close connections
to Hamilton. Following their successful introduction, numbers
of guanacos on Staats Island fluctuated widely over the years
primarily due to reproduction and hunting, i.e. culling (shot
with permission) and poaching (without permission).
Why guanacos were originally taken to Staats Island is not
clear. Hamilton and Miller had been established on Staats
Island since the mid-1920s where they had set up a holding
facility for importing foxes (and probably other Patagonian
wildlife) before transferring the animals to other islands
(FIA:CS, 1926b). There was housing on Staats in the form of
a shanty built by Henry and James Waldron in the 1870s
(today called ‘Waldron’s Shanty’ at Hamilton Bay (Fig. 5),
where Hamilton and Miller could have stayed. Perhaps stormy
weather or ship mechanical problems forced them to unload
the guanacos temporarily at Staats, but the animals escaped
before they could be reloaded and taken to Sedge. Perhaps they
realized that because Staats had been without sheep for
25 years since 1917 (FIA:CS, 1926b) and offered favourable
grazing greens (i.e. vegas), they wanted to divide their guanaco
importation between two islands and would later explain the
matter to the government. Or perhaps they wanted to provide
a permanent source of carrion for their imported foxes. Rob
McGill (pers. comm.) recalls as a child hearing of accounts that
before guanacos had been brought to Staats, the Penelope made
several trips a month from Weddell carrying a quantity of live
sheep that were killed for fox food. Or finally, perhaps the
government granted permission to Hamilton and Miller, but
the documentation has been lost or is yet to be uncovered.
Whichever the case, guanacos were introduced to Staats Island
and rapidly began to increase, although they also came very
close to disappearing 20 years later. It is interesting to note the
same combination of naturalized species from Patagonia was
introduced both to Sedge and Staats Islands: skunks, foxes,
rheas and guanacos.
Nearly exterminated
In December 1940, ownership of Hamilton’s islands was
transferred to the John Hamilton Estates Ltd., known locally as
the ‘Company’. Staats Island was overseen and managed out of
the headquarters on Weddell Island. Beginning in the mid-
1950s, an organized effort was made to exterminate the
guanacos of Staats Island, because of the poor state of
the animals and overgrazing that had occurred on the island.
W. L. Franklin and M. M. Grigione
668 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
The Company put out a request for shooters to go to Staats.
William Davis, at the time a shepherd on Weddell Island, and
Dan Cronin, a travelling-school teacher in the West Falklands,
volunteered (W. Davis, pers. comm.). Over a 1-week period in
the winter of 1956 they culled 103 guanacos. ‘The animals had
eaten all their food. From close range we shot animals that
were so weak they couldn’t get up the hills. There was a bunch
still left, 150–170’ (W. Davis, pers. comm.). Months before
talking to Mr Davis, based upon estimated rates of increase, we
had calculated that there were 150 guanacos remaining after
this first major hunt (Table 2). Davis and Cronin, following
the long tradition of inscribing your name in Waldron’s
Shanty, carved and burned on the cupboard door: ‘W.J.D.
D.R.C. 14-VII-1956 103 guanacoes’ (Franklin, unpubl. Shanty
data).
Three years later Dan Cronin, well known marksman
(R. McGill, pers. comm.), again returned to Staats, this time
with John Bound, a government man from Stanley (W. Davis,
pers. comm.). The outcome was reported in the Falkland
Islands Monthly Review (FIA:CS, 1959): ‘Some 20 odd years
ago the late John Hamilton imported various animals and
birds to the Colony from Chile and placed them on islands in
the Weddell group. Today the main survivors are the guanacos
and foxes which have become a menace to the sheep farming
industry – the guanaco is a natural forage grazer and the fox a
killer. Recently John Bound and Dan Cronin visited Staats
Island, by arrangement with the Manager of Weddell (Mr. M.
W. McGill) and shot off 111 Guanaco in a week. The Guanaco
population of the island is now estimated to be in the region of
150…, unless they are completely exterminated before too
long, soil erosion (already in evidence) is likely to become a
major problem’. This time with Bound, Danny Cronin again
carved and burned on the same cupboard door in Waldron’s
Shanty the results of his second hunt: ‘J.B. D.R.C. 10-I-1959
111 Guanacoes’ (Franklin, unpubl. Shanty data).
The Cronin-Bound estimate of 150 guanacos surviving their
hunt was low, for 2 months later, brothers Rob and Lyall
McGill from Weddell Island modestly recorded on the
cupboard the results of a 1-week cull: ‘R.M.G. L.M.G.
8-3-1959 175 Guanacoes & 7 Foxes’ (Franklin, unpubl. Shanty
data). Rob McGill recounts, ‘There was sort of a competition
going on with John Cronin, but still there was no pleasure in
the shooting. We were there as employees of the Company’. At
the end of their hunt the McGills counted 46 guanacos
Figure 5 High elevation aerial photo (taken
at 3,700 m in 1956) of Staats Island, West
Falklands indicating names of key locations
and geographical features.
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 669
scattered across the island (R. McGill, pers. comm.). These
early culls were amazingly successful, in part because the
population had been unhunted since its introduction. Yet as
the years of hunting passed, the animals became more leery
and difficult to shoot. On 2 November 1991, P. Berntsen wrote
on a ceiling beam in Waldron’s Shanty: ‘1 quanaco (sic) 1 fox
60 Bullets’ (Franklin, unpubl. Shanty data).
It was the Hamilton Estate’s intention during the late-1950s
to kill off all the guanacos on Staats Island and replace them
with sheep (R. McGill, pers. comm.). Ian Strange, now a well-
known Falkland author and naturalist, tried to convince the
Hamilton consortium not to destroy the whole population
because of its historical and biological value. Fortunately, other
farm duties and perhaps logistical problems prevented them
from commencing the final cull (Strange, pers. comm.).
Company policy during this era, plus the general impression
that ‘guanacos… have become a menace to the sheep farming
industry’ that should be ‘completely exterminated’ (FIA:CS,
1959), created an ‘outsider-pest’ attitude, that resulted in open
hunting of guanacos on Staats for adventurers from Weddell
Island, Port Stephens, and beyond. Sometimes it was carried
out with authorization, other times not. The McGill hunt in
1959 was the last major, organized culling of the population,
yet individuals continued to visit the island and shoot animals
(R. McGill, pers. comm.). Claims have since been made that
‘we used to go over and cull them ever year’. As a consequence,
we believe that in the early 1960s the population was reduced
to as few as 10–20 guanacos and came dangerously close to
disappearing (Table 2, Fig. 6). Ten years later in 1970 it had
recuperated to only some 60 animals (Strange, 1972).
How reasonable are the estimated numbers of guanacos on
Staats over the past half-century (Table 2)? The numbers in
fact support the assertion that 15 animals were originally
introduced onto Staats Island. Growing South American wild
camelid populations typically increase at a rate of 11–16% per
year (Franklin, 1973, 1974, unpubl. data). It is not unreason-
able to assume a 18–19% annual rate of growth for a new
population filling an unoccupied niche with favourable
habitat. In this case, at a 18–19% annual increment,
a population of 15 guanacos in 1939 would have increased
to the estimated 275 animals just prior to first major culling in
1956. As carrying capacity and habitat quality decreased on
Staats, the annual rate of increase would have declined to a
more typical rate of 12–14%, as it did between 1982–89,
Table 2 Numbers of guanacos imported, counted, estimated by calculation (est.), and culled (shot) on Staats Island, Falkland Islands
1938–2003
Date Number Remarks Source
April 1938 5 Imported FIA:CS (1934–39)
May 1939 10 Imported FIA:CS (1934–39)
May 1939 15 Total after two imports
June 1956 275 Pre-cull population est. at 18–19% growth since 1939
14 July 1956 103 Culled by William Davis (WJD) & Dan Cronin (DRC) Davis (pers. comm.), Franklin
(unpubl. Shanty data)
August 1956 175 Post-cull est. numbers
August 1956 150–175 Visually assessed Davis (pers. comm.)
31 December 1958 300 Pre-cull population est. at 14–15% growth since August 1956
10 January 1959 111 Culled by John Bound (JB) & Dan Cronin (DRC) FIA:CS (1959), Franklin (unpubl. Shanty data)
10 January 1959 150 Incorrectly assessed by JB & DRC FIA:CS (1959)
8 March 1959 175 Culled by Rob McGill (RMG) & Lyall McGill (LMG) Franklin (unpubl. Shanty data),
R. McGill (pers. comm.)
8 March 1959 46 Post-culling count R. McGill (pers. comm.)
Early 1960s 10–20 Est. low numbers
1970 60 Counted on ground (12% growth since early 1960s) Strange (1972)
1982 136 Counted by helicopter Strange (1992) and see Lurcock (1991)
1983 150–200 Estimated population By Ian Strange (see Lurcock, 1991)
May–September 1985 21 Culled by Tiny Bennett, Jerry Jones,
Steve Forky, and Terry Harding
Franklin (unpubl. Shanty data)
October 1989 306 Counted by motorbike (13% incr. since 1982) S. Poncet (pers. comm.)
July 1990 15 Culled by owners S. Poncet (pers. comm.)
21 October 1990 223 Counted on foot Lurcock (1990)
April 1991 252 Counted on foot (13–14% incr. since 1990) Lurcock (1991)
1994–95 23 Culled by owners S. Poncet (pers. comm.)
1997 20 Culled by owners S. Poncet (pers. comm.)
17 November 1999 338 Counted on foot Franklin (2003)
2001–02 23 Poached by ?? Franklin (2003)
11 December 2002 303 Counted on foot Franklin (2003)
15 December 2003 392 Counted on foot (13% incr. since 2002) Franklin (2004)
W. L. Franklin and M. M. Grigione
670 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
1990–91 and 2002–03 (Table 2). In face of occasional culling
and poaching, there are now some 400 guanacos on Staats
Island (392 counted part way through the December 2003
birthing season) (Franklin, 2004).
The smallness of the original founding population intro-
duced onto Staats, but especially the greatly reduced numbers
in the early 60s to 10–20 guanacos (or perhaps even less) is
suggested by our recent fieldwork on Staats. Franklin (2003,
2004) and his colleagues have found strong preliminary
evidence of the deleterious consequences of inbreeding in the
form of premature birthing, late-term abortions, newborn
physical deformities, and birthing problems resulting in the
death of newborns.
Territorial males (Solo and Family Group) are especially
vulnerable to hunting because of their strong site fidelity (see
Young & Franklin, 2004). Territorial males are commonly
reluctant to leave their territories, even in the face of danger
when other group members retreat. Consequently, they are
approachable on foot and easier to shoot, as Sarno et al. (1996)
found when immobilizing adult territorial males for popula-
tion studies on the mainland in Torres del Paine National
Park. When the Staats Island guanaco population was severely
reduced by shooting and approached extirpation in 1959, it is
quite possible that, being easier targets, only one to two males
survived, further narrowing the impact of genetic bottleneck-
ing and subsequent genetic defects.
Why did guanacos endure on Staats Island? Its isolation on
the western edge of a 12,000 km2 archipelago, in which the vast
majority of the people live on the eastern edge in East Falkland
(Strange, 1972, 1992), was surely a factor. Staats Island is in a
remote region where few people venture. Its growing guanaco
population probably did not attract much attention until the
mid-1950s when the first major culling was carried out
17 years after their introduction. Another important factor is
the favourable guanaco habitat found on Staats Island.
Abundant greens or vegas occur in the four major valleys
(Fig. 5) of Staats Island (Franklin, 2004). We now know that
vega is a high quality (12% protein), highly preferred habitat
for this species, with guanacos spending as much as 80–95% of
their feeding time in these areas (Franklin, 1983; Lawrence,
1990; Ortega & Franklin, 1995; Bank et al., 1999). While tussac
grass and native boxwood (Hebe eliptica) have nearly disap-
peared from Staats Island due to fire and overgrazing by cattle,
sheep and guanacos (Strange, 1987; Franklin, unpubl. data)
and there are areas of sheet erosion on some steep slopes, the
vegas continue to be in good condition, supporting a guanaco
population in good physical condition (Franklin, 2004).
Conditions will improve even more with the restoration of
tussac grass communities by fencing off selected barren areas
on Staats Island, as currently being considered by the island’s
owners (S. Poncet, pers. comm.).
IMPLICATIONS, QUESTIONS, AND THE FUTURE
It is very surprising, that despite the importance of wool
production over the past century in the Falklands, the high
economic value of guanaco wool has been totally ignored.
Although the first governor of the Falklands suggested as long
ago as the 1840s that raising guanacos would be a ‘worthy
endeavour’ because of the value of its wool and hides (Lever,
1985), the guanaco has been viewed as a pest. Ironically, while
the Falkland Islands Monthly Review (FIA:CS, 1959) was
referring to them as a ‘menace’ that should be exterminated,
guanacos were being harvested for their valuable wool in a
multimillion-dollar industry in Argentina on the mainland
(Mares & Ojeda, 1984; Franklin & Fritz, 1991). Sheep wool
ranges in price from US$0.50 to 1.50 per pound (c. 0Æ5 kg),
while guanaco wool is worth US$50 to 150 per pound and
more. Today, harvesting of guanaco wool from wild and farm
guanacos is being carried out in Chile, Argentina (Franklin
et al., 1997) and Wales (R. Lerwill, pers. comm.). The
guanacos of Staats Island represent a potential source for
introductions to other Falkland islands for guanaco ranching
and wool production. When some 400 animals were shot in
the late 1950s, the hunters would have been shocked had they
realized the guanaco wool left on the rotting carcasses blowing
away in the South Atlantic winds, was worth $20,000–25,000,
based upon today’s value.
Biologically, Staats Island is a unique natural experiment in
progress. It is an outdoor laboratory whose large mammal
population is isolated, relatively small, and completely closed.
No animals can leave except by death, nor enter except by
birth. The powerful component of this ecological drama is that
it is all unfolding on an island within a limited, defined
resource base. What is there, is there, and no more. It is
amazing that the guanacos have not only survived, but are
thriving, a testimony to their adaptiveness and resiliency.
While guanacos have impacted the landscape, they have not
destroyed their own food base (Franklin, 2003, 2004). How
have they done it? How has the habitat been able to sustain
such heavy use? How does this population differ from
mainland populations where puma predation is a major cause
of mortality (Franklin et al., 1999)? Are the significantly lower
birth weights of newborns on Staats compared with the
0
50
100
150
200
250
300
350
400
1935 1945 1955 1965 1975 1985 1995 2005Year
Nu
mb
er o
f G
uan
aco
s
Figure 6 Reconstruction of the numbers of guanacos on Staats
Island in the West Falklands since their introduction in 1938–39,
as primarily influenced by population growth, culling and
poaching. See Table 2 for details.
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 671
mainland (24 lb vs. 30 lb) (Franklin & Johnson, 1994;
Franklin, 2004) influencing juvenile survival (Sarno &
Franklin, 1999b)? In addition, what about the question of
genetic bottlenecking and differences compared with mainland
populations (Sarno et al., 2001)? Has this small gene pool held
sufficient genetic diversity for an expanded, healthy popula-
tion? So far, it seems so. What is the full impact of 65 years of
inbreeding on chromosomal aberrations and birth defects? Can
the limited genetic diversity of Staats Island be a model,
whether positive or negative, for small and insular-mainland
populations targeted for conservation (Franklin et al., 1997)?
Is the Staats’ guanaco population genetically healthy enough to
be used for introductions to other islands for guanaco farming?
Future field studies and genetic analyses will further clarify
these observations and questions.
The vicuna (Vicugna vicugna), the other South American
wild camelid with a similar population and behavioural
ecology to the guanaco, has been shown to self-regulate its
population density through density-dependent and beha-
vioural mechanisms (Franklin, 1983). We do not know if the
guanacos of Staats Island have such a self-controlling mech-
anism or not. It is noteworthy to point out, however, that
inbreeding and/or natural selection have resulted in self-
reduction of the population through late-term abortions,
premature births, birth defects and even dystocia (difficult
birthing) (Franklin, 2004). Could these events be a part of a
natural self-regulating mechanism? Will the population con-
tinue to grow? Will it reach a plateau and level off because of
such genetic and density-dependent mechanisms? Or will it
suddenly decline? Only the future will tell. Staats Island offers a
rare opportunity for ungulate field studies to be taken to an
advanced level by marking and potentially genetically identi-
fying every animal on the island, making it possible to assess
the influence of relatedness, ancestry, and genetic health upon
the behaviour, distribution and survival of a truly closed
population (Franklin, 2003).
There is currently a growing ethic in the Falklands Islands
towards environmental stewardship. With the general col-
lapse of the sheep industry that dominated the economy
over the past century, tourism and fishing have become the
new lifelines of support for this island-nation and its peoples
(Summers, 2001). As Hamilton himself once pioneered and
practiced, the removal and reduction of sheep is now
allowing the regeneration of overgrazed rangelands and lost
native habitats throughout the Falklands. The beauty of the
landscape and abundant sub-Antarctica wildlife is drawing
increasingly more tourism with the promise of a genuine
broader economic base. John Hamilton undertook numerous
new and creative activities that promoted land stewardship
and economic development. Not all his attempts were
successful, nor were all viewed as positive contributions by
fellow Falklanders. Now, as the Falklands move toward a
more diverse economy with tourism becoming increasingly
important, its history and spectacular wildlife are its biggest
attractions (Summers, 2001). Penguins and marine mammals
are the stars. Yet, this unique island population of guanacos
on Staats Island offers a concurrent opportunity for visitors
to see one of the premiere wild mammalian herbivores of
the Americas. Hamilton’s original intention to help diversify
the country’s economy with the introduction of guanacos,
may still become a reality.
What is in store for guanacos of the Falkland Islands? At this
stage it is undecided. However, over the past several years
consideration has been given to everything from complete
eradication to the creation of a National Nature Reserve or
Trust. Preservation and management seem to be the direction,
yet neither the island nor its denizens are under any official
protection. Whatever the course, recognition of the popula-
tion’s historical importance, biological uniqueness, and eco-
nomic value will surely be major considerations in determining
its future.
CONCLUSIONS
There are some gaps in our understanding of all the historical
and biogeographical events that surrounded the introduction of
guanacos into the Falklands Islands. Yet, the enigma of how and
why guanacos are there is resolved. With government approval,
two shipments totalling 11 guanacos were introduced onto
Sedge Island in 1934 and 1935, but subsequently disappeared.
Two shipments totalling 15 guanacos were then introduced
either accidentally or intentionally onto Staats Island in 1938
and 1939. These animals, in spite of a tumultuous history of
heavy culling that nearly caused their extirpation in the late-
1950s and early 1960s, have survived over the past 65 years and
today some 400 guanacos are found on Staats Island. Staats
Island is a unique natural experiment in progress. As manifes-
ted in the classical studies by Clutton-Brock et al. (1982) on red
deer (Cervus elaphus) on the Scottish Isle of Rhum, closed
populations offer an unusual opportunity for addressing
advanced questions in ungulate population, behavioural and
genetic ecology. In addition, the guanacos of Staats Island
represent potential economic contributions to Falkland Island
tourism and breeding stock for other islands.
John Hamilton, the spirited and visionary Scottish pioneer
to the Falklands in the early 1880s, was the driving force
behind the importation of guanacos into the Falklands. But as
the guanaco project was gaining momentum in the late-1930s,
it abruptly came to an end with the beginning of War II and
the death of Hamilton in 1945 (FIA:CS, 1945). In the end, his
progressive ideas did not survive the man (Bernhardson,
1988). Yet among his many efforts to practice land stewardship
and promote economic diversity through the introduction of a
myriad of Patagonian wildlife species, a remaining and lasting
legacy of John Hamilton to the Falkland Islands is undeniably
the guanacos of Staats Island.
ACKNOWLEDGEMENTS
We would like to express our appreciation to Jerome and Sally
Poncet for permission to work on Staats Island; their interest
in conservation and research are essential to this work and the
W. L. Franklin and M. M. Grigione
672 Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd
well being of the island’s guanacos. We thank the Turner
Foundation, Hidden Corners Inc., University of South Florida,
and paying research-expedition participants for their financial
support of the field studies on Staats Island, as well as the
stalwart field assistance of Ross Burrus, Liza Carter, Cody
Franklin, Doug Frohling, Charlotte M. Hughes, Joyce H.
McWilliams, Mary Parke ‘Perky’ Manning, Ron Sarno, and
Tim Stotts. Mark Mueller (University of South Florida,
Department of Environmental Science and Policy) assisted
with the preparation of figures. The Falkland Island Govern-
ment Department of Agriculture provided important colla-
boration, including the hardy field assistance of Dr. Cameron
Bell, Suzanne Halfacre, and Dr. Kevin Lawrence from the
Division of Veterinary Services. We thank the many individ-
uals we interviewed who provided invaluable insight and
information, especially Bill Davis of Goose Green (East
Falkland), Trevor Halliday of Tasmania, Dave Hawksworth
of Sedge Island, Sarah Lurcock of South Georgia Islands,
members of the MacBeth family (Phyllis, Raymond, and
Rowena M. Summers) who formerly lived and farmed on
Sedge Island, Rob McGill of Carcass Island, Roderick Napier of
West Point Island, Tony Pole-Evans of Saunders Island, and
Ian Strange of New Island South Wildlife Reserve. Neil
Rowlands of Hebe Tours provided important assistance with
logistics and radio communication. Thanks to Tom Chater for
his flying skills and taking aerial photos in very windy skies,
and to our boat captains who expertly sailed the challenging
Falkland seas: Michael J. Clarke (Penelope), Robin Lee
(deceased) (West Swan), Michael McRae (Laura Jay), and
Jerome Poncet (Golden Fleece). Special thanks to Dr. Wayne
Bernhardson for generous use of his primary research on John
Hamilton and the Falkland Islands. Colour work was gener-
ously covered by the Department of Natural Resource Ecology
& Management at Iowa State University. And finally, this work
could not have been accomplished without the monumental
help of Archivist Jane Cameron, whose expertise and devotion
to her important work at the Falkland Islands Archives in
Stanley are highly appreciated.
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Bernhardson, W. (1988) John Hamilton’s legacy in the Falkland
Islands, 22 pp. Department of Geography, University of
California, Berkeley.
Bernhardson, W. (1989) Land and life in the Falkland Islands
(Islas Malvinas). PhD Dissertation, Department of Geog-
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Bridges, E.L. (1950) Uttermost part of the earth. Dutton, New
York.
Clutton-Brock, T.H., Guinness, F.E. & Albon, S.D. (1982) Red
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BIOSKETCHES
Bill Franklin is Professor Emeritus of Animal Ecology from Iowa State University. As a Mammalian Wildlife Ecologist his research
interests have focused on the behavioural and population ecology of a variety of mammals, but primarily the wild camelids of South
America, for over 35 years. His initial work was with endangered vicuna at the Pampa Galeras National Vicuna Reserve in the
altiplano of southern Peru. He then conducted and directed field research on the guanaco in Tierra del Fuego and Torres del Paine
National Park in southern Chile from the mid-1970s until his retirement in 2000.
Melissa Grigione’s primary research interest is mammalian spatial ecology and understanding how ecological and man-made
elements influence individual location and home range size. She works with a variety of species (including mountain lions, manatees,
burrowing owls and elephants) whose populations have been seriously altered as a consequence of habitat degradation and
fragmentation. In addition to wildlife ecology, her research utilizes knowledge of political and legislative systems, and community-
level human dimension practices in order to conserve species. Her international work includes conservation projects for the guanaco
and vicuna in South America and conservation of neotropical cats along the US–Mexico border.
Editor: Philip Stott
Guanacos of the Falkland Islands
Journal of Biogeography 32, 661–675, ª 2005 Blackwell Publishing Ltd 675
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