the enigma of guanacos in the falkland islands: the legacy of john hamilton

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: [email protected]

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



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.



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



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



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.



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.



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)



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.



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



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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29, 1929–30, 1930–31, 1931–32, 1932–33.

FIA:CS (1928a) 91/28, 224/27 Crown Grant No. 411 regarding

acreage of Passage Island.

FIA:CS (1928b) 278/26 Letter from John Hamilton to Colonial

Secretary regarding foxes, March 22.

FIA:CS (1930) OD, Series G, Vol 26 Memorandum from

Hodson to Passfield regarding Hamilton, 4 November.

FIA:CS (1931) 557/27 Letter from John Hamilton to Colonial

Secretary, 2 December.

FIA:CS (1932–47) Shipping Register/Stanley, Falkland Islands

1932–47.

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1934–35, 1935–36, 1936–37, 1937–38, 1938–39.

FIA:CS (1934a) 40/26 Letter from John Hamilton to Ellis

regarding Tussac, 14 March.

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Halkett to J. Miller regarding importation, 21 June.

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April.

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naturalist from Montagu Craigie-Halkett, Colonial Secre-

tary, 5 March.

FIA:CS (1940) 136/40 Livestock File J.E. Hamilton [not related

to John Hamilton], Government Naturalist to Colonial

Secretary, 22 July.

FIA:CS (1945) Falkland Islands Weekly News, 27 September.

FIA:CS (1959) Guanaco hunting in the Falklands. Falkland

Islands Monthly Review, February.

Franklin, W.L. (1973) Conservation of the vicuna in Peru.

World Wildlife Yearbook 1972–73 (ed. by P. Jackson), pp.

209–212. World Wildlife Fund, Morges, Switzerland.

Franklin, W.L. (1974) The social behavior of the vicuna. The

behaviour of ungulates and its relation to management (ed. by

V. Geist and F. Walther), pp. 477–487. IUCN, Morges,

Switzerland.

Franklin, W.L. (1975) Guanacos in Peru. Oryx, 13, 191–202.



Franklin, W.L. (1981) Living with guanacos: wild camels of

South America. National Geographic, 160, 62–75.

Franklin, W.L. (1982) Biology, ecology, and relationship of

man to the South American camelids. Mammalian biology in

South America (ed. by M.A. Mares and H.H. Genoways), pp.

457–487. Special Publication Series, 6. Pymatuning Labor-

atory of Ecology, University of Pittsburgh, Pittsburgh.

Franklin, W.L. (1983) Contrasting socioecologies of South

American wild camelids: the vicuna and the guanaco.

Advances in the study of mammalian behavior (ed. by J.F.

Eisenberg and D.G. Kleiman), pp. 573–629. Special Publi-

cation 7, American Society of Mammalogists. Allen Press,

Lawrence, Kansas.

Franklin, W.L. (2003) Guanaco research expedition to Staats

Island in the western Falkland Islands: preliminary results

and proposed plans for the future (December 2002). 6 pp.

Department of Agriculture, Stanley, Falkland Islands.

Franklin, W.L. (2004) Report on guanaco research expedition:

Staats Island–Falkland Islands (December 2003), 11 pp. + 1

map. Department of Agriculture, Stanley, Falkland Islands.

Franklin, W.L. & Fritz, M.A. (1991) Sustained harvesting of the

Patagonia guanaco: is it possible or too late? Neotropical

wildlife use and conservation (ed. by K. Redford and

J. Robinson), pp. 573–629. University of Chicago Press,

Chicago, IL.

Franklin, W.L. & Johnson, W. E. (1994) Hand capture of

newborn open-habitat ungulates: the South American gua-

naco. Wildlife Society Bulletin, 22, 253–259.

Franklin, W.L., Bas, F.M., Bonacic, C.F., Cunazza, C.P. & Soto,

V.N. (1997) Striving to manage Patagonia guanacos for

sustained use in the grazing agroecosystems of southern

Chile. Wildlife Society Bulletin, 25, 65–73.

Franklin W.L., Johnson W.E., Iriarte, J.A. & Sarno, R.J. (1999)

Ecology of the Patagonia mountain lion, Felis concolor

patagonica, in southern Chile. Biological Conservation, 90,

33–40.

Garay, G., Franklin, W. L., Sarno, R.J. & Johnson, W.E. (1995)

Development of juvenile guanaco social behavior: first study

on a wild population. Revista Chilena de Historia Natural,

68, 1–10.

Gustafson, L.L., Franklin, W.L., Sarno, R.S., Hunter, R.H.,

Johnson, W.E., Young, K.M. & Behl, M.J. (1998) Predicting

early mortality of newborn guanacos by birthweight and

hematological parameters. Journal of Wildlife Management,

62, 24–35.

Hoppe, J. & McAdam, A. (1992) Management of tussac grass

communities for wildlife, tourism and agriculture. Aspects of

Applied Biology, 29, 377–381.

Johnson, W.E. & Franklin, W.L. (1994) Role of body size in the

diets of sympatric gray and culpeo fox. Journal of Mam-

malogy, 75, 163–174.

Johnson, W.E., Franklin, W.L. & Iriarte, J.A. (1990) The

mammalian fauna of the northern Chilean Patagonia: a

biogeographical dilemma. Mammalia, 54, 457–469.

Kerr J. A. & McAdam, J.H. (2003) Characterization of patches

of short grass and herbs in the Falkland Islands and their role

in sheep grazing, 27 pp. Report for the Department of

Agriculture, Stanley, Falkland Islands.

Lawrence, R.K. (1990) Factors influencing guanaco habitat use

and group size in Torres del Paine National Park. MS Thesis,

Iowa State University, Ames.

Lever, C. (1985) Naturalized mammals of the world. Longman,

New York.

Lurcock, S. (1990) Report on visit to Staats Island 20–22 October

1990. 11 pp. + 2 maps. Department of Agriculture, Stanley,

Falkland Islands.

Lurcock, S. (1991) Handwritten notes 11pp including results of

her guanaco survey on Staats Island in 1991 and personal

interview with Ian J. Strange. Falkland Islands Department of

Agriculture, Stanley.

Mainwaring, M. (1983) From the Falklands to Patagonia. All-

ison and Busby, London.

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lization and conservation in South America. BioScience, 34,

580–584.

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natural history, prehistory and ethnography at the uttermost

part of the earth. Princeton University Press, Princeton.

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endangered ungulates of Chile: vicugna, lama, hippocame-

lus, and pudu. North American Wildlife Natural Resources

Conference, 38, 55–68.

Moore, D.M. (1968) The vascular flora of the Falkland Islands.

British Antarctic Survey Scientific Reports, Number 60.

NERC, London.

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London.

Nowak, R.M. (1999) Walker’s mammals of the world, Vol. II.

Johns Hopkins University Press, Baltimore, MD.

Ortega, I.M., & Franklin, W.L. (1995) Social organization,

distribution and movements of a migratory guanaco

population in the Chilean Patagonia. Revista Chilena de

Historia Natural, 68, 489–500.

Prichard, H. (1902) Field-notes upon some of the larger

mammals of Patagonia, made between September 1900 and

June 1901. Proceedings of the Zoological Society of London, I,

272–277.

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flippers: the evolution of hoofed mammals. John Hopkins

University Press, Baltimore, MD.

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the guanaco (Lama guanicoe) of Magallanes, Chile. PhD

Dissertation, University of Washington, Seattle, WA.

Sarno, R.J. & Franklin, W.L. (1999a) Maternal expenditure in

the polygynous and monomorphic guanaco: suckling

behavior, reproductive effort, yearly variation, and

influence on juvenile survival. Behavioral Ecology, 10, 41–47.

Sarno, R.J. & Franklin, W.L. (1999b) Population density and

annual variation in birth mass of guanacos in southern

Chile. Journal of Mammalogy, 80, 1158–1162.

Sarno, R.J., Hunter, R.L. & Franklin, W.L. (1996)

Immobilization of guanacos by use of Tiletamine/



Zolazepam. Journal of the American Veterinarian Medical

Association, 208, 408–409.

Sarno, R.J., Franklin W.L., O’Brien, S.J. & Johnson, W.E.

(2001) Assessing genetic differentiation between an island

and mainland population of guanacos in southern Chile

utilizing mtDNA and microsatellite markers. Animal Con-

servation, 4, 93–101.

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the Falkland Islands. The Falklands Islands Journal, 2, 32–34.

Strange, I.J. (1972) The Falkland Islands. Stackpole Books,

Harrisburg.

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Falkland Department of Agriculture, Stanley, Falkland

Islands.

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report (Falkland Islands). Typed. Department of Agriculture,

Stanley, Falkland Islands.

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Islands and South Georgia. Harper Collins, London.

Summers, D. (2001) A visitor’s guide to the Falkland Islands.

Hanway Press, Hong Kong.

Woods, R.W. (2002) Corrections to a survey of the number,

size and distribution of islands in the Falklands archipelago.

The Falklands Islands Journal, 8, 120.

Young, J.K. & Franklin, W.L. (2004) Territorial fidelity of male

guanacos in the Patagonia of Southern Chile. Journal of

Mammalogy, 85, 72–78.

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



the enigma of guanacos in the falkland islands: the legacy of john hamilton

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