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Until recently the Antarctic continent and Peninsula have been little impacted by non-native species, compared to other regions of the Earth. However, reports of species introductions are increasing as awareness of biological invasions as a major conservation threat, within the context of increased human activities and climate change scenarios, has grown within the Antarctic community. Given the recent increase in documented reports, here we provide an up-to-date inventory of known terrestrial non-native species introductions, including those subsequently removed since the 1990s, within the Antarctic Treaty area. This builds on earlier syntheses of records published in the mid-2000s, which focused largely on the sub-Antarctic islands, given the dearth of literature available at that time from the continental and maritime Antarctic regions. Reports of non-native species established in the natural environment (i.e. non-synanthropic) are mainly located within the Antarctic Peninsula region and Scotia Arc, with Deception Island, South Shetland Islands, the most impacted area. Non-native plants have generally been removed from sites of introduction, but no established invertebrates have yet been subject to any eradication attempt, despite a recent increase in reports. Legislation within the Protocol on Environmental Protection to the Antarctic Treaty has not kept pace with environmental best practice, potentially presenting difficulties for the practical aspects of non-native species control and eradication. The success of any eradication attempt may be affected by management practices and the biology of the target species under polar conditions. Practical management action is only likely to succeed with greater co-operation and improved communication and engagement by nations and industries operating in the region.
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REVIEW PAPER
Biological invasions in terrestrial Antarctica: what
is the current status and can we respond?
Kevin A. Hughes Luis R. Pertierra Marco A. Molina-Montenegro
Peter Convey
Received: 5 January 2015 / Revised: 13 February 2015 / Accepted: 26 February 2015
The Author(s) 2015. This article is published with open access at Springerlink.com
Abstract Until recently the Antarctic continent and Peninsula have been little impacted
by non-native species, compared to other regions of the Earth. However, reports of species
introductions are increasing as awareness of biological invasions as a major conservation
threat, within the context of increased human activities and climate change scenarios, has
grown within the Antarctic community. Given the recent increase in documented reports,
here we provide an up-to-date inventory of known terrestrial non-native species intro-
ductions, including those subsequently removed since the 1990s, within the Antarctic
Treaty area. This builds on earlier syntheses of records published in the mid-2000s, which
focused largely on the sub-Antarctic islands, given the dearth of literature available at that
time from the continental and maritime Antarctic regions. Reports of non-native species
established in the natural environment (i.e. non-synanthropic) are mainly located within the
Antarctic Peninsula region and Scotia Arc, with Deception Island, South Shetland Islands,
the most impacted area. Non-native plants have generally been removed from sites of
introduction, but no established invertebrates have yet been subject to any eradication
attempt, despite a recent increase in reports. Legislation within the Protocol on
Communicated by Karen E. Hodges.
Electronic supplementary material The online version of this article (doi:10.1007/s10531-015-0896-6)
contains supplementary material, which is available to authorized users.
K. A. Hughes (&)P. Convey
British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road,
Cambridge CB30ET, UK
e-mail: kehu@bas.ac.uk
L. R. Pertierra
A
´rea de Biodiversidad y Conservacio
´n, Departamento de Biologı
´a y Geologı
´a, Universidad Rey Juan
Carlos, c/ Tulipa
´n, s/n., 28933 Mostoles, Madrid, Spain
M. A. Molina-Montenegro
Centro de Estudios Avanzados en Zonas A
´ridas (CEAZA), Facultad de Ciencias del Mar, Universidad
Cato
´lica del Norte, Coquimbo, Chile
123
Biodivers Conserv
DOI 10.1007/s10531-015-0896-6
Environmental Protection to the Antarctic Treaty has not kept pace with environmental
best practice, potentially presenting difficulties for the practical aspects of non-native
species control and eradication. The success of any eradication attempt may be affected by
management practices and the biology of the target species under polar conditions.
Practical management action is only likely to succeed with greater co-operation and im-
proved communication and engagement by nations and industries operating in the region.
Keywords Antarctic Treaty area Environmental Protocol Alien species Biosecurity
Invasion Eradication
Introduction
The vulnerability of Antarctic terrestrial ecosystems to human-induced environmental
impacts and changes has been a focus of recent research attention (e.g. Bargagli 2005;
Frenot et al. 2005; Kerry and Riddle 2009; Tin et al. 2009; Hughes and Convey 2010,
2012; Cowan et al. 2011; Chown et al. 2012a) and human-assisted establishment of non-
native species, both in the context of those introduced from beyond the Antarctic region
and in the movement of Antarctic biota between different eco-regions within the continent,
has been identified as one of the most pervasive threats to indigenous ecosystems (Hughes
and Convey 2010; Chown et al. 2012b; Terauds et al. 2012).
Antarctica’s isolation and recent colonization by humans, compared to the other con-
tinents, means it has relatively few known non-native species (Frenot et al. 2005; Hughes
and Convey 2010). Globally, intentional non-native species introductions have occurred
for economic, scientific and social reasons, frequently with unforeseen consequences for
local environments (Mack et al. 2000). However, no terrestrial species introduced to the
Antarctic continent intentionally (mainly for scientific research reasons) have been for-
mally identified as invasive (Smith 1996), but intentional introductions to most sub-
Antarctic islands, before the adoption of legislation prohibiting or controlling this activity,
have resulted in substantial impacts (see Convey and Lebouvier 2009). Under current
legislative systems, unintentional introductions present the greatest threats to sub-Antarctic
and Antarctic ecosystems (Frenot et al. 2005), with non-native species potentially being
introduced associated with visitors’ clothing and personal effects (Whinam et al. 2005;
Chown et al. 2012b; Huiskes et al. 2014), cargo (Hughes et al. 2010; Tsujimoto and Imura
2012), building material (Lee and Chown 2009) and fresh foods (Hughes et al. 2011).
It is well recognised that the implementation of effective biosecurity practices is the
most cost-effective method of reducing non-native species establishment and subsequent
impacts (Chown et al. 2012b) (for examples see the Council of Managers of National
Antarctic Programs (COMNAP) and Scientific Committee on Antarctic Research (SCAR)
‘Checklists for supply chain managers of National Antarctic Programmes for the reduction
in risk of transfer of non-native species’, available at: https://www.comnap.aq/SitePages/
checklists.aspx). This is particularly true for Antarctica, where there are relatively few
standard access points and routes, and human activities have the potential to be highly
regulated by individual national operators and the tourism and fishing industries. However,
with increasing numbers of people visiting Antarctica, an increase in the diversity of
activities undertaken and the complexity of coordinating management measures between
the parties involved, activities that may lead to the introduction of non-native species have
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proven difficult to regulate effectively. Consequently, the introduction and establishment
of some new non-native species (as well as the repeated introductions of existing non-
native species in multiple locations) may be almost inevitable (see Fig. 1). Effective
management response—including non-native species eradications—is therefore of para-
mount importance for the protection of Antarctic ecosystems, even accepting that the most
effective actions would be to prevent introductions occurring in the first place.
Fig. 1 a Nassauvia magellanica eradicated from Deception Island in January 2010 (Photo: K. A. Hughes).
bThe flightless chironomid midge Eretmoptera murphyi, introduced to Signy Island, South Orkney Island
from South Georgia (Photo: P. Bucktrout). cPoa annua on Deception Island and subsequently eradicated
(Photo: M. Molina-Montenegro).]. dPoa pratensis on Cierva Point, Antarctic Peninsula, where it was first
introduced during transplantation experiments in 1954/1955 (Photo: L. R. Pertierra). eTrichocera
maculipennis found in Artigas Base (King George Island, South Shetland Islands) sewage system in
2006/2007 and in surrounding terrestrial habitats (Photo: O. Volonterio). fNon-native potted plant in the
window of Bellingshausen Station in 2010 (King George Island) (Photograph: K. A. Hughes). gRemoval of
an alien grass species from the vicinity of Great Wall Station, Fildes Peninsula, King George Island in 2006
(Photo: S. Pfeiffer)
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Several previous studies have provided syntheses of the then-existing state of knowl-
edge relating to the presence of non-native species within Antarctica, some including the
sub-Antarctic islands within their ‘area of interest’ (Smith 1996; Frenot et al. 2005,2008;
Convey 2008). Smith’s (1996) study focused on records of non-native higher plants, while
Frenot et al. (2005) collated information across all terrestrial vertebrate, invertebrate and
plant groups, further recognising the paucity of information available relating to lower
plant and microbial groups. Over 95 % of the c. 200 established non-native species in-
cluded in Frenot et al.’s (2005) synthesis were present in the sub-Antarctic, with almost all
of the remaining species being recorded from the maritime Antarctic. These reviews,
together with related specific studies, have stimulated increasing interest in the risks of
biological invasions in the Antarctic, both within the scientific research community and in
the governance community of the Antarctic Treaty System, and led to an increase in
studies recording the presence and, in some cases, impacts of non-native species within the
Antarctic Treaty area (defined as the area south of 60S latitude).
The purpose of this study was therefore to bring the work of Frenot et al. (2005)upto
date with regard to the Antarctic Treaty area (excluding the sub-Antarctic Islands), and to
examine the number and distribution of macroscopic non-native species introduced or
identified since c. 1990 and of any earlier introductions still persisting in the Antarctic
terrestrial environment. We also review the current legislation that applies within the
Antarctic Treaty area regarding the eradication and control of non-native species, and
recommend appropriate management action.
Antarctic terrestrial habitats and biodiversity
The unique characteristics of habitats and communities with the Antarctic terrestrial en-
vironment may make them particularly vulnerable to invasive species impacts. Terrestrial
ice- or snow-free habitats in Antarctica comprise only c. 0.34 % of the continental area
(Convey et al. 2009), and the majority of this area is devoid of visible or macroscopic
biota. Terrestrial biodiversity is low, both in terms of species and functional diversity
(Hogg et al. 2006; Convey 2013). Recent biodiversity analyses have identified 15 distinct
Antarctic Conservation Biogeographic Regions (ACBRs) within the continent (Terauds
et al. 2012). The most developed terrestrial ecosystems are present close to the coast,
particularly along the western Antarctic Peninsula and islands along the Scotia Arc, and in
oases along the coast of East Antarctica—areas that are also favoured for the siting of
research stations, and often coincide with concentrations of wildlife and spectacular
scenery that attract tourist activity. However, specialized biological communities are also
present even in the most extreme terrestrial habitats within the continent (e.g. Broady and
Weinstein 1998; Hodgson et al. 2010). The majority of areas of exposed terrestrial ground
are isolated, small and island-like (Bergstrom and Chown 1999; Arnold et al. 2003; Hughes
et al. 2006), factors that are important in driving the evolutionary isolation, divergence and
high levels of regional endemism that appear to characterise Antarctic biota (Chown and
Convey 2007; Convey 2008; Pugh and Convey 2008). Even within the McMurdo Dry
Valleys of southern Victoria Land, by far the most extensive area of ice-free ground within
the continent, studies of microbial and arthropod communities report signals of isolation
and divergence within valleys and catchments (McGaughran et al. 2008,2010; Chan et al.
2013). Isolation, high levels of endemism and a general lack of inter-species competition
within many native terrestrial Antarctic communities may make them particularly vul-
nerable to the impacts of invasive species (Chown and Convey 2007; Convey 2008).
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The distribution of non-native species within the Antarctic continent
The known distribution of non-native species within Antarctica is shown in Fig. 2and the
dataset upon which the figure is based is provided in the supplementary data (Table A1).
All current introductions are found within the Antarctica Peninsula and Scotia Arc, and all
within ACBR 2 ‘South Orkney Islands’ and (by far the most invaded) ACBR 3 ‘Northwest
Antarctic Peninsula’. Although no species appear to be extant, non-native plants have been
previously found and removed from continental Antarctica (Japan 1996; Russia 1999). To
date, there are no confirmed reports of existing non-native species being transported to
further locations within Antarctica, although this possibility presents a significant risk. It
should be noted that many Antarctic non-natives are cryptic or non-charismatic, which may
make their detection and assured eradication more difficult. Nevertheless, multiple intro-
ductions to Arctowski Station (Admiralty Bay, King George Island, South Shetland Is-
lands) of Poa annua from both European and South American origins have been reported
(Chwedorzewska 2008). Our analyses show introductions to date have been dominated by
Collembola and Poaceae, species of both having commonly become established and, in
many cases, becoming invasive, on the sub-Antarctic islands (Frenot et al. 2005).
The first report of a non-native plant to become established in Antarctica was of a now
absent ‘flowering grass’ found near houses on Deception Island in January 1936 by the
British Graham Land Expedition (Smith 1996); however, the plant known to have persisted
longest in Antarctica is Poa pratensis, which was introduced to Cierva Point over 60 years
ago with little range expansion before its removal in January 2015 (Pertierra et al. 2013;L.
R. Pertierra, pers. obs., 2015). The first non-native invertebrate to be reported was the
Collembolan Hypogastrura viatica found on Deception Island in the 1940s (Hack 1949).
These findings suggest that the introduction and at least transient establishment of non-
native species is likely to have been occurring for as long as humans have been inhabiting
suitable Antarctic locations, and possibly from the early nineteenth century when sealers
first visited the northern Antarctic Peninsula and South Shetland Islands. Prior to the
implementation of the Protocol on Environmental Protection to the Antarctic Treaty (also
known as the Madrid Protocol or Environmental Protocol) in 1998, which prohibited the
introduction of non-Antarctic soil, cultivation of non-native plants in imported soil oc-
curred at several Antarctic stations [e.g. Maitri and Novolazarevskaya Stations (Arif and
Joshi 1995)] and non-native invertebrate species were reported from within the imported
soils (Arif 1995). Furthermore, at some stations considerable quantities of fodder was
imported to feed domesticated animals [e.g. c. 30 sheep and 100 fowl were kept at Arturo
Prat Station on Greenwich Island, South Shetland Islands (Anonymous 1960)], which was
likely to contain viable plant propagules and other non-native species. It would seem
appropriate that monitoring for the presence of non-native species in the vicinity of these
locations should be a priority (see Smith 1996 for an overview). Contrary to the Envi-
ronmental Protocol, non-native potted plants are still found in some Antarctic stations in
both private quarters and public spaces (for example, potted rose and cycad plants were
found to be present at the entrance to Bellingshausen Station, King George Island, South
Shetland Islands (P. Convey, pers. obs., 1 Feb 2015) (see also Fig. 1f). At a minimum,
Treaty Parties should conform to the legislation contained within the Environmental
Protocol and remove non-native species introduced for decorative purposes.
Figure 3shows the number of discrete locations currently colonised by each non-native
species known within Antarctica. The most widely dispersed species are micro-inverte-
brates, with knowledge of their distributions resulting from recent soil surveys at popular
visitor sites (Russell et al. 2013). The Actinedida Speleorchestes sp., Coccotydaolus cf.
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Fig. 2 Map of the Antarctic Peninsula region showing the distribution of known non-native species
(Cryptopygus caecus has not been included on this and subsequent maps due to uncertainty over its native/
non-native status (see Russell et al. 2013), while Alicorhagia sp. has not been included as the single report
recorded only a single individual and it was unclear whether or not this species had established)
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krantzii,Terpnacarus gibbosus and Collembolan Hypogastrura viatica were all found at
multiple locations, with Speleorchestes sp. the most widely dispersed across six locations
within the northern Antarctic Peninsula and South Shetland Islands area. P. annua is now
present at only one location (Admiralty Bay, South Shetland Islands), with small numbers
of plants being removed from four other locations during the 2009/2010 summer (De-
ception Island and three sites on the Antarctic Peninsula; see Table 1). Nevertheless,
removal is no guarantee of eradication as propagules may remain in the seed bank resulting
in the potential reappearance of plants at the site of initial removal. Figure 4shows the
locations currently colonised by at least one non-native species. On the basis of rather
limited surveys, Deception Island, with nine non-native species, is the most invaded,
followed by Fildes Peninsula and Neko Harbour with four each. All of these locations are
subject to high levels of national operator and tourist activity. The majority of other
locations containing one or two non-native colonists are close to, or located within,
established research stations or popular tourist visitor sites (see Fig. 1 in Molina-Mon-
tenegro et al. 2014). These findings are also closely consistent with areas predicted to be at
highest risk of non-native species introductions, based on propagule pressure and climate
suitability (Chown et al. 2012b).
Figure 5shows the number of non-native plants (upper panel) and invertebrates (lower
panel) discovered within Antarctica over time (pre-1990s, 1990s, 2000s, 2010s). Species
Fig. 3 The number of locations
colonised by each non-native
species within Antarctica
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Table 1 Non-native species removal and eradication attempts within the Antarctic continent and off-shore islands
No. Species Location Colonisation date and extent Eradication date and
outcome
Reference
1Poa trivialis
Reclassified as Puccinellia
svalbardensis
Near Syowa Station, Enderby Land,
East Antarctica
1993? Single plant Removed in 2007 (S.
Imura, pers. comm.)
Japan (1996) and
Tsujimoto et al. (2010)
2Poa annua Faraday Research Station (now
Vernadsky Station) Galindez
Island, Argentine Islands
Introduced pre-1981 Destroyed before 1985 Smith (1984,1996)
3Poa annua General Bernardo O’Higgins
Station, Trinity Peninsula, northern
Antarctic Peninsula
2007/2008: single plant
2009/2010: two plants
Removed 2009/2010 Molina-Montenegro et al.
(2012)
4Poa annua Gabriel Gonzalez
Videla Station, Paradise Bay,
northern Antarctic Peninsula
2007/08: single plant
2009/2010: four plants
Removed 2009/2010 Molina-Montenegro et al.
(2012)
5Poa annua Almirante Brown Station, Paradise
Bay, northern Antarctic Peninsula
Introduced pre-2009/2010:
two plants
Removed 2009/2010 Molina-Montenegro et al.
(2012)
6Poa annua Whalers Bay, Deception Island,
South Shetland Islands
Introduced pre-2009/2010:
one plant
Removed 2009/2010 Molina-Montenegro et al.
(2014)
7Alopecurus geniculatus,Puccinellia
distans,Rumex pulcher,Stellaria
media and Chenopodium rubrum
Progress II Station, Larsemann Hills,
Ingrid Christensen Coast, Princess
Elizabeth Land
Seventeen plants found in
1995 near the staircase of
the medical block
No further reports,
presumably removed (c.
1999?)
Russia (1999)
8Cerastium sp. and non-native
graminoids
Great Wall Station, Fildes Peninsula,
King George Island, South
Shetland Islands
1997? Removed 2005 Smith (2003), Peter, H.-
U., (2005) (quoted in
Hughes and Convey
2010)
9 Unidentified Poaceae
Poa annua?
Great Wall and Bellingshausen
Stations, Fildes Peninsula, King
George Island, South Shetland
Islands
Introduced c. 2004 (possibly
as early as 1996). Several
individual plants
established in the environs
of the stations
Removed 10 February
2006
Plants had flowered. Seed
bank possible
Peter et al. (2008,2013)
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Table 1 continued
No. Species Location Colonisation date and extent Eradication date and
outcome
Reference
10 Unidentified
Poaceae
Poa annua?
Bellingshausen Station, Fildes Peninsula Single plant, 8 cm diameter Removed 30 December
2008
Peter et al. (2013)
11 Nassauvia
magellanica
Whalers Bay, Deception Island, South
Shetland Islands
Introduced pre-Jan 2009.
Single plant
Removed 23 Jan 2010.
Four plants were found
originally, but three had
been washed away,
together with another
non-native species
Gamochaeta nivalis,by
the time of the
eradication
Smith and Richardson (2011), United
Kingdom and Spain (2010) and Hughes and
Convey (2012)
12 Poa pratensis Primavera Station, Cierva Point, Danco
Coast, Palmer Archipelago, Antarctic
Peninsula
Introduced in soil imported
from Ushuaia during
transplantation experiments
in 1954/1955. Between
1995 and 2015 it expanded
from a patch c. 40 cm
across to c. 1 m across
Removed January 2015 Corte (1961), Smith (1996), Pertierra et al.
(2013) and Pertierra, pers. comm. (2015)
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which were introduced deliberately for scientific investigations and were then either re-
moved or did not survive are not included in the analysis (see Smith (1996) for further
details). It is clear that removal of single or small numbers of plants at different Antarctic
locations has been effectively achieved over recent decades (Argentina et al. 2013), with
the notable exception of P. annua in Admiralty Bay. In contrast, no eradication or control
of non-synanthropic invertebrates has been attempted in the region, while recent surveys
have resulted in a substantial increase in the number of species and locations known to be
invaded by this biological group.
It is not entirely clear which, if any, of the non-native species that have established in
Antarctica have become invasive according to the definition contained within the CEP Non-
native Species Manual (2011) i.e. ‘are extending their range in the colonised Antarctic region,
displacing native species and causing significant harm to biological diversity or ecosystem
functioning’. However, the non-native grass, P. annua has spread into the local terrestrial
communities near Arctowski Station and in laboratory experiments has been shown to have
negative impacts on photosynthetic performance and biomass of the two native Antarctic
vascular plants Colobanthus quitensis and Deschampsia antarctica (Molina-Montenegro
et al. 2012). Furthermore, it has been estimated that larvae of the introduced chironomid
midge Eretmoptera murphyi, that now occupies an area of over 35,000 m
2
on Signy Island,
South Orkney Island, may be able to cycle soil nutrients up to nine times faster than the entire
native soil invertebrate community and, therefore, could have a major effect on terrestrial
habitats across colonised areas on Signy Island (Hughes et al. 2013).
Fig. 4 Number of non-native
species present at each invaded
location
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Fig. 5 Location of known non-native plants (upper panel) and invertebrates (lower panel) within
Antarctica up until 1990 and during the 1990s, 2000s and 2010s. Plants eradicated or removed are not shown
in subsequent figures. No records of non-native invertebrates exist outside the Antarctic Peninsula region
and no eradications have been attempted
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Antarctic legislation concerning non-native species and their control and eradication
General legislation concerning non-native species within the Antarctic Treaty area is re-
viewed elsewhere (Hughes and Convey 2010,2014). A brief summary is provided here to
give context to the subsequent discussion, with legislation of specific relevance to
eradication activities described in more detail.
Legislation relating to non-native species within the Antarctic Treaty area
Legislation relating to non-native species is contained within the Protocol on Environ-
mental Protection to the Antarctic Treaty, predominantly in Annex II Conservation of
Fauna and Flora. Article 4 of this Annex prohibits specifically the intentional introduction
of non-native plants and animals to land, ice shelves or into water within the Antarctic
Treaty area, unless for a defined scientific purpose and in accordance with a permit. Non-
native species can be introduced in accordance with a permit for laboratory studies, but
only on the condition that they are destroyed or removed from the Treaty area at the end of
the permitted period, and measures are taken to eliminate any potential risks to native
plants and animals. The prohibition of non-native species importation does not apply to
food items, but no live animal may be imported for human consumption. Furthermore, all
imported plant and animal produce (fresh fruit, vegetables, eggs and meat) must be stored
under controlled conditions, although there is no general requirement, for instance, for
quarantine or inspection facilities, or fumigation of containers used for food transport.
Imported animal carcasses must be disposed of by incineration, autoclaving or made sterile
before disposal, or be removed from the Antarctic Treaty area. Precautions must be taken
to prevent the introduction of non-native microorganisms: diseased meat is not to be
imported to the Treaty area and the importation of non-sterile soil originating outside the
Treaty area is to be avoided as far as possible. However, the Environmental Protocol makes
little reference specifically to the unintentional introduction of non-native species, or the
transportation of species (including native species) between different biogeographic re-
gions within the Antarctic Treaty area.
The Protocol does not differentiate between different categories of non-native species
(for instance invasive versus persistent; for further discussion of non-native species clas-
sifications as have been applied in Antarctica see Frenot et al. (2005) and Greenslade et al.
(2012)), rather prohibiting the introduction of all non-native plants, animals and mi-
croorganisms, irrespective of likely colonisation status. Consequently, many of the prob-
lematic issues faced by policymakers in other regions of the world concerning which non-
native species to allow or tolerate and which to exclude, control or eradicate are not
relevant within the Antarctic Treaty area. However, the unusual nature of Antarctica’s
legislative framework may present disadvantages for conservation, as some legislation
applicable elsewhere in the world may not apply in Antarctica (Baker et al. 2005). For
example, the Convention on Biological Diversity (CBD) Article 8(h) ‘Alien species that
threaten ecosystems, habitats or species’ does not apply in the Antarctic Treaty area as the
CBD itself applies explicitly to sovereign territory. This is despite the fact that many of the
Antarctic Treaty signatory nations have themselves signed up to the CBD.
Annex VI to the Environmental Protocol, which has yet to enter into force, concerns
liability arising from environmental emergencies. It is unclear if non-native species in-
troductions can be included under the definition of an ‘environmental emergency’, or if it
will be possible to use this legislation to reclaim costs for the control or eradication of a
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non-native species introduced by the actions of another operator (see Hughes and Convey
(2014) for further discussion).
Legislation pertinent to non-native species eradication and control
All activities undertaken by Consultative Parties within the Antarctic Treaty area must
conform with the requirements of the Environmental Protocol, as must those of non-
Consultative Parties who are signatories to the Protocol. Various issues addressed in the
Protocol may have practical implications for the planning and achievement of eradications.
The need for evidence to support eradication or control action
Annex II, Article 4(4) states that ‘Any other plant or animal introduced into the Antarctic
Treaty area not native to that area, including any progeny, shall be removed or disposed of,
by incineration or by equally effective means, so as to be rendered sterile, unless it is
determined that they pose no risk to native flora or fauna’. While this Article is apparently
clear that non-native species shall be eradicated, any management action may be directed
by two criteria which may be difficult or impossible to prove conclusively, specifically (1)
that the species concerned is actually non-native to Antarctica (also noting that the wording
does not explicitly recognise the need for removal of species transferred between what are
now recognised as biogeographically distinct areas within Antarctica) and (2) that it may
pose a risk to native biota (Hughes and Convey 2012). While the rapidity of response to
non-native species introductions is a major factor in determining eradication success [and
is a key guiding principle in the Committee for Environmental Protection Non-native
Species Manual (CEP 2011)], obtaining this information may delay eradication action. A
more appropriate policy for Antarctica might be the approach advocated by Simberloff
(2003), which is characterized by the principle ‘shoot first, ask questions later’. Here, the
need for a detailed risk assessment and characterisation of the introduced species followed
by investigations into its interactions with other native species (all of which take up
valuable time during which the species may be expanding and becoming increasingly
difficult to eradicate) becomes of secondary importance. One drawback here, however, is
the potential for eradication of species that have colonized Antarctica by natural means,
which may impact upon the already very low natural re-colonization rates for the continent
(see Hughes and Convey 2012). For the second criterion, one option is that the burden of
proof should be placed on managers to show that the non-native species poses ‘no risk to
native flora or fauna’, with the meaning of ‘risk’ defined by scientific experts. Where any
doubt exists, eradication should proceed at the earliest opportunity.
The environmental impact assessment process
All activities planned within the Antarctic Treaty area must undergo some level of envi-
ronmental impact assessment (EIA) as mandated by Annex I to the Environmental Pro-
tocol. Activities assessed as having an impact less than ‘minor or transitory’ can proceed
forthwith. However, in some cases the removal of a non-native species may have an
environmental impact that is ‘minor or transitory’, triggering the need for an Initial En-
vironmental Evaluation (IEE), or impact greater than ‘minor or transitory’, which would
necessitate a Comprehensive Environmental Evaluation (CEE). CEEs must be presented
by the proponent Party at an annual meeting of the Committee for Environmental
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Protection, which could cause delays in the commencement of any eradication attempt.
Annex I does allow for activities to be undertaken in cases of emergency which would
otherwise have required preparation of a CEE. Under such circumstances, notice of the
activities ‘shall be circulated immediately to all Parties and to the Committee [for Envi-
ronmental Protection] and a full explanation of the activities carried out shall be provided
within 90 days of those activities’ [Article 7(2)]. How this would operate in practice is
untested as no precedents exist, but at present it seems that individual Parties would decide,
on a case by case basis, whether or not the discovery and need to eradicate a non-native
species constituted an emergency.
Impacts of non-native species eradication and control activities on non-target native
species
Methodologies used to eradicate or control non-native species may have unavoidable
negative effects upon invaded habitats and native species. However, Annex II Article 3(3c)
states that ‘the diversity of species, as well as the habitats essential to their existence, and
the balance of the ecological systems existing within the Antarctic Treaty area be main-
tained’. While eradication of non-native species may help maintain indigenous species
diversity and Antarctic habitats, it is less clear how to assess objectively whether or not the
benefits of undertaking the eradication of a non-native species outweigh any potential
negative impacts upon indigenous biota. Furthermore, no indication is given as to the
spatial scale that should be taken into consideration. For example, eradication of the
flightless midge Eretmoptera murphyi and the enchytraeid worm Christensenidrilus blocki
on Signy Island, South Orkney Islands, or P. annua on King George Island may entail
destruction of a substantial area of habitat and associated non-target indigenous species
(e.g. an area of at least c. 35,000 m
2
in the vicinity of the UK research station in the case of
E. murphyi, based on a survey carried out in 2007, and recognising that the species
appeared to have entered a phase of rapid distribution expansion at that time), and Parties
may be reluctant to support such an activity (Hughes and Worland 2010; Olech and
Chwedorzewska 2011; Hughes et al. 2013; Olech 1996).
Allocation of permits to undertake eradication or control of non-native species
Permits issued by an appropriate national authority to take or interfere in a harmful manner
with native biota are required and can only be granted to provide specimens for scientific
study, to supply specimens for museums and other educational establishments, or to
‘...provide for the unavoidable consequence of scientific activities………or of the con-
struction and operation of scientific support facilities’. It is not clear how the destruction of
species and habitat as a side-effect of non-native species eradication aligns with this
legislation [Annex II, Article 3(2)].
Eradication of non-native species using pesticides
Practical methods of non-native species eradication may be restricted by Annex III Waste
Disposal and Management, as Article 7 states that ‘No……pesticides (other than those
required for scientific, medical or hygiene purposes) shall be introduced onto land or ice
shelves or into water in the Antarctic Treaty area’. Environmental management and con-
servation purposes are not listed in the Annex, and this issue may therefore require further
Biodivers Conserv
123
consideration by the CEP and ATCM before use of such chemicals is authorised. Parties’
domestic sensitivities regarding the use of these chemicals may further complicate any
decision making.
Eradication of non-native species in protected areas
Further delays in the initiation of any eradication attempt may be encountered if the non-
native species is found within an Antarctic Specially Protected Area (ASPA) or Antarctic
Specially Managed Area (ASMA). The accompanying management plans may need to be
revised to allow eradication activities to occur, as the disruption of habitat and use of
pesticides and herbicides (which are technically a sub-set of pesticides) are generally not
permitted. Seventy five percent of all ASPAs, including almost all those that protect
terrestrial habitats, prohibit specifically the use of herbicides and pesticides within the
Area. The one notable exception to this is contained within the management plan for ASPA
136 Clark Peninsula, Budd Coast, Wilkes Land, East Antarctica, which prohibits herbi-
cides from being taken into the Area ‘...unless needed to mitigate any non-native species
incursions. Such chemicals must only be used as a last resort and controlled by permit
conditions.’ Within almost all ASPAs the current management plans may limit Parties’
choice of methodologies to respond rapidly to a non-native species introduction (Table 2).
Considerations for management of non-native species eradications in Antarctica
Eradication attempts within the Antarctic Treaty area
Eradication attempts that have taken place within the Antarctic Treaty area are listed in
Table 1. To date, management responses to non-native species introductions to the natural
habitats within the Treaty area have either not occurred or have been carried out in an
opportunistic or ad hoc manner. Of these, successful eradications have been limited to
removal of small numbers of individual vascular plants located near research stations or
frequently visited sites. In some cases Parties have been slow to respond to non-native
species introductions, despite requests from other organisations. For instance, SCAR
recommended that P. annua be eradicated from around the Polish Henryk Arctowski
Station, Admiralty Bay, King George Island in the early 1990s (Smith 2011). However,
although considerable efforts have been made to monitor and study the plants (Olech 1996,
1998,2003; Chwedorzewska 2008,2009), no attempt at eradication has been reported
(Olech and Chwedorzewska 2011).
Non-native species have also been found living synanthropically within Antarctic
buildings, sewage treatment facilities and hydroponic facilities (Hughes and Convey 2010;
Volonterio et al. 2013). While successful eradications have occurred within station
buildings and hydroponic facilities, no successful eradications of non-native invertebrates
have occurred within Antarctic sewage treatment systems (Table 3; Hughes et al. 2005). In
the case of the boreal trichocerid fly Trichocera maculipennis, first discovered in the
sewage system of the Uruguayan Artigas Base (Volonterio et al. 2013), the history of
colonisation, eradication and apparent subsequent recolonisation of the tanks after several
years’ absence could also be interpreted as being consistent with this pre-adapted cold
environment fly having currently undetected source population(s) in the natural environ-
ment of King George Island rather than it being restricted to the confines of the station
itself.
Biodivers Conserv
123
Table 2 Measures within Antarctic Specially Protected Area (ASPA) management plans detailing use and
storage of herbicides and pesticides within the protected area
No. Management plan measures ASPA (primary value being protected) Percentage
of all
ASPAs %
1 Herbicides and pesticides are prohibited
within the Area
54 ASPAs, including all ASPAs primarily
protecting terrestrial habitat
75.0
2 No chemicals to be used in the Area except in
accordance with a permit
ASPA 120 (birds and mammals), ASPA 125
(geological values), ASPA 127
(penguins), ASPA 166 (historic values)
5.6
3 Chemicals may only be introduced for
permitted scientific or conservation
purposes
ASPAs 155, 157, 158, 159, 162 (all
protecting historic values)
7.0
4 Use of chemicals within the Area is not
prohibited, but storage within the ASPA not
permitted
ASPA 174 (geological values) 1.4
5 ‘Chemicals……which may be brought into
the Area for scientific or management
purposes specified in the Permit, shall not
be released into the environment
ASPA 175 (geothermal habitat) 1.4
6 ‘No herbicides are to be taken into the Area
unless needed to mitigate any non-native
species incursions. Such chemicals must
only be used as a last resort and controlled
by permit conditions’
ASPA 136 Clark Peninsula, Budd Coast,
Wilkes Land, East Antarctica
1.4
7 No prohibition of chemicals ASPA 122, 144, 145, 146, 156, 168. None
protect terrestrial habitat primarily
8.3
Most management plans specifically prohibit use of both herbicides and pesticides, which suggests some
confusion over definitions of these terms, as herbicides are a sub-category of pesticides
Table 3 Invertebrates that have colonised station buildings and sewage treatment plants on Antarctic
research stations
Species Station Date
introduced
Notes References
Lycoriella
inge´nue
Casey Station, Budd
Coast, Wilkes Land
1998 Extensive eradication
attempt in 2005 proved
unsuccessful
Hughes et al.
(2005) and Smith
(2005)
Lycoriella sp. Rothera Research
Station, Marguerite
Bay, Antarctic
Peninsula
2005 Successful eradication of
flies from alcohol store in
2005
Hughes et al.
(2005)
Trichocera
maculipennis
Artigas Station, Fildes
Peninsula, King
George Island, South
Shetland Islands
2006? Early eradication attempt
unsuccessful. Species is
now found in the
surrounding environment
Volonterio et al.
(2013)
Unidentified
mosquito
Frei Station, Fildes
Peninsula, King
George Island, South
Shetland Islands
Pre-2009/
2010
season
Larvae persist in the sewage
treatment plant. No
counter measures are
undertaken
V. Vallejos, pers.
comm., quoted in
Peter et al. (2013,
Sect. 3.1.4)
Hydroponic facilities, operated by Parties including Australia, New Zealand and the US, have been tem-
porarily closed down and cleaned due to infestations by imported non-native invertebrates (COMNAP 2013)
Biodivers Conserv
123
Factors affecting the likely success of an eradication within an Antarctic context
Eradication success can depend upon many factors, with each weighted differently de-
pending upon the particular circumstances of each non-native species introduction (Sim-
berloff 2002). It is unlikely that a standard policy for non-native species eradication within
the Antarctic Treaty area can be formulated at anything other than the most general level;
however, those responsible for controlling non-native species should be wary of delaying
extermination (potentially catastrophically) due to lack of complete information. Decision-
making should benefit from consideration of the following factors:
Economic benefits and use of resources
In the Antarctic continent, the low number of identified non-native species (and fewer, if
any, confirmed invasive species) (Frenot et al. 2005; Hughes and Convey 2012) implies
that priorities and resource allocation can still usefully be focused towards implementing
effective biosecurity precautions (Rout et al. 2011). Economic drivers for eradications are
largely absent as Antarctica has no income generating industries that may be directly
impacted by non-native species, such as terrestrial agriculture, freshwater aquaculture or
nearshore fish farming (Perrings et al. 2000; Pimentel et al. 2005). Nevertheless, bio-
prospecting activities, (including those concerning microbial species) may be impacted by
non-native species, which could be a driver for improved conservation and/or non-native
species control in the future (Hughes et al. 2015).
The Antarctic Treaty Parties have no central reserve to fund environmental initiatives.
Consequently Parties, in general, act either independently or in collaboration with a small
number of like-minded nations. If eradications are undertaken, it is important that Parties
are fully aware of the financial and time (including logistic) commitment necessary from
the outset, so that adequate resources are allocate to complete the task. The position in
Antarctica contrasts with the sub-Antarctic islands, each of which is governed by a single
sovereign nation, and which have experienced a far greater presence and impact of non-
native species (Frenot et al. 2005; Convey and Lebouvier 2009). Here, in recent years,
attention has focused on the eradication of various herbivorous and predatory vertebrates,
funded either by the national government, or by private or charitable donors (Bloomer and
Bester 1991; Frenot et al. 2005; Bergstrom et al. 2009). Some mammalian eradications
have led to unintended consequences, such as the destruction of plant species by increased
rabbit numbers following the eradication of cats on Macquarie Island (Bergstrom et al.
2009). While high profile, vertebrate eradications represent only part of the challenge as
non-native plants and invertebrates represent the majority of known introductions and
include a range of invasive species having considerable impacts on ecosystems and native
biota (Frenot et al. 2005). Remarkably, within the Antarctic the majority of eradications
have removed the vascular plants while no invertebrate eradications have yet been at-
tempted, illustrating the different priorities and capacities for managing different taxa (see
Fig. 5).
Lines of authority
The Antarctic Treaty System operates by consensus, and no mechanism exists by which a
Party can be compelled to engage in an activity or cooperate with any eradication plan.
Due to ambiguity in the text of the Environmental Protocol, Parties may interpret the
legislation in different ways (Joyner 1999), or rank other aspects of their Antarctic
Biodivers Conserv
123
activities as of higher priority. Within the Antarctic Treaty area, the pace of action may be
delayed due to the need for Parties to inform and engage with each other, before under-
taking activities not strictly limited to their own logistics and research. Therefore, if a non-
native species is found in an area where operational footprints overlap, eradication at-
tempts may be delayed while consensus between Parties is reached regarding timing,
methodology and on-going monitoring, despite the CEP Non-native Species Manual
(2011) key guiding principle that any eradication response should be rapid.
An alternative case has been proposed whereby scientists should study the establish-
ment and expansion of non-native species as a research subject itself. This was apparently
put forward in response to requests and advice to eradicate P. annua from the vicinity of
the Polish Arctowski Station in the 1990s (see Smith 2011). Olech (1996) stressed the
importance of evaluating properly the effects of man’s activities on Antarctic ecosystems
which, in the case of P. annua at Arctowski station, may be one reason that scientific
values were prioritised over environmental protection (Smith 2011). However, given the
wealth of information already available about biological invasions, their impacts, and their
management and the challenges therein available from the rest of the world, the case for
any expectation of different principles applying to invasions in Antarctica would seem
weak.
Biology of the target species
A clear understanding of the biology of the non-native species is important when planning
an eradication attempt. This is particularly relevant once a species has increased its dis-
tribution beyond the initial point of establishment, as eradication and monitoring tech-
niques suitable for small sites may be impractical for larger colonised areas. Important
information to guide choice of the most effective eradication methodology can be gleaned
from the species’ distribution, life cycle characteristics (Thompson et al. 1995; Crawley
et al. 1996) and invasiveness in similar environments (e.g. in the sub-Antarctic islands, or
in Arctic or high altitude habitats), but with a recognition that these characteristics may be
different within an Antarctic context (e.g. see the risk assessments described in Greenslade
(2002) and Greenslade and Convey (2012)). Of particular importance is an understanding
of which life cycle stages are most likely to aid species dispersal and establishment (e.g.
possession of a winged stage, or diapause capacity; production of seeds or vegetative
propagules), which are most vulnerable to the eradication methodologies available, the rate
of dispersal in other habitats and the physiological limits or thresholds relevant to different
life cycle stages. For example, larval size class data and growth rate modelling for the non-
native chironomid midge Eretmoptera murphyi, which was introduced to Signy Island,
South Orkney Islands, from South Georgia, has suggested that it persists as larvae in the
soil for 2 years on Signy, while this life cycle stage may last only 1 year on the warmer
South Georgia (Hughes et al. 2013). The physical, biological and chemical characteristics
of colonised ground may affect the rate of spread. For example, disturbed ground may
facilitate expansion of ruderal species (Grime 1977), as has been observed in Antarctica
with P. annua (Olech 1996; Molina-Montenegro et al. 2014), as could the spatial limits of
appropriate microclimatic conditions.
Understanding the phenological strategies and physiological capacities of vascular
plants are crucial to designing appropriate methods of eradication. P. annua has shown a
remarkable flowering capacity even under extreme environmental conditions, therefore any
attempted eradications of fast-growing plants (r-strategists) must address the likelihood of
an extensive seed bank that may persist long after any physical removal of plants (Pertierra
Biodivers Conserv
123
et al. 2013;Wo
´dkiewicz et al. 2013). Under these circumstances pesticide use may be
effective, combined with on-going monitoring for plant re-growth at the site.
Probability of re-introduction
Within Antarctica, and between Antarctica and the other southern continents, there are a
relatively limited number of logistic routes and connections between locations and, hence,
control of species transfer through application of biosecurity measures may be more
practicable and effective than in the rest of the world (Chown et al. 2012b; Molina-
Montenegro et al. 2012; Hughes et al. 2014). Treaty Parties and tour operators, in large
part, are in control of the people and cargo that are transported into Antarctica, and it is
largely within their power to control the risk of re-introduction through investment in
biosecurity measures. Eradication efforts may be largely wasted if adequate biosecurity
precautions that reduce further introductions are not implemented.
Habitat restoration
An eradication attempt will have failed if indigenous species are driven extinct at the
eradication location alongside the introduced species, or are not able to regain earlier
population levels. The consequences of an eradication may be hard to predict, but could
include negative and unforeseen impacts on local indigenous communities, such as a shift
in the dominant native biota at a site (Bergstrom et al. 2009). Given the contemporary
relative scarcity of non-native species in Antarctica, to a large extent habitat restoration in
Antarctica is not an issue as yet, though it is clearly important in the sub-Antarctic. If
Antarctic habitats are damaged by eradication action then, by analogy with the rates of
recovery from other impacts (such as ground disturbance or trampling), recovery may
require decades at least (Tin et al. 2009).
Practical considerations
Eradication of many introductions requires trivial commitment of resources [e.g. a person
with a spade (see Fig. 1g)], and lessons could also be learnt from previous eradications in
the Antarctic and sub-Antarctic (Table 1) (Smith 1984,2003; Japan 1996; Peter et al.
2008; United Kingdom and Spain 2010; Tsujimoto et al. 2010; Smith and Richardson
2011; Hughes and Convey 2012 (see Table 5 therein); Shaw 2013). Those undertaking an
eradication should consider: (1) the most appropriate method (or methods), (2) the im-
plications of there being more than one non-native species at a location and whether it is
feasible to eradicate them simultaneously (see Table A1 and Fig. 4), (3) how to prevent
inadvertent further distribution of the targeted species in the local area during the
eradication work, (4) how best to co-ordinate the proper disposal of any removed organ-
isms and other associated material such as soil or vegetation, and (5) what monitoring
programme should be put in place to ensure the eradication has been effective.
Conclusions and recommendations
The level to which biosecurity measures are implemented by national operators and the
tourism industry in the Antarctic has not been rigorously assessed or tested (but see
Biodivers Conserv
123
COMNAP 2008). However, within Fildes Peninsula, King George Island, which is located
in one of the regions at highest risk of non-native species introductions (Chown et al.
2012b, Fig. 4), Peter et al. (2013) reported that ‘The various stations of the Fildes
Peninsula currently take either no measures or limited measures to prevent the introduction
of non-native speciesOn the contrary, people still commonly keep house plants in a
number of stations. To our knowledge, no measures are implemented to monitor non-
native species’ (see Fig. 1f). It is not known if this level of engagement in non-native
species and biosecurity issues is common throughout Antarctica, but the assessment is a
cause for concern, following as it does the matter of non-native species being attributed the
highest priority on the Committee for Environmental Protection work plan since first used
as a management tool in 2007 [ATCM XXX Final Report (para. 230) http://www.ats.aq/
devAS/ats_meetings_meeting.aspx?lang=e]. Furthermore, recent monitoring activities on
the northern Antarctic Peninsula have revealed a higher number and distribution of non-
native species than was previously known (Greenslade et al. 2012; Molina-Montenegro
et al. 2012; Russell et al. 2013,2014), suggesting that biosecurity practices employed to
date are not adequate.
Successful management of non-native species will require cooperation between Treaty
Parties, national operators, the tourism industry and other stakeholders (Convey et al.
2012). In 2002, Simberloff wrote ‘successful eradication may be as much a function of
political skill and public education as of technology’. This is likely to be particularly true
for Antarctica as consensus is required before any change in legislation can occur or major
activity be undertaken. Therefore, the objections of even a single Party could put in
jeopardy plans for any sizeable eradication attempt. Furthermore, lack of communication
between scientists, policy makers and those putting policy into practice, both within in-
dividual nations and across the Antarctic Treaty System, may limit the success of efforts by
any one group alone. Initiatives such as the Environments Portal (www.environments.aq),
which is supported by several Treaty Parties and SCAR, may go some way in bridging
these knowledge gaps. In an attempt to produce a more integrated, comprehensive and
dynamic approach to conservation in the region and to inform conservation decision-
making and policy, including those concerning non-native species, SCAR is developing an
Antarctic Conservation Strategy (SCAR et al. 2012).
To reduce the impact of non-native species within the Antarctic Treaty area, almost all
areas of management action would benefit from improvement, particularly:
Better education of all Antarctic visitors.
Application of consistent and effective biosecurity practices in all Antarctic operations,
recognising that ‘prevention is better than cure’.
Monitoring of high activity sites for non-native introductions and suitable reporting,
including support for appropriate expertise with which to identify potential
introductions.
The development of contingency plans following the discovery of a non-native species,
including the provision of eradication protocols and appropriate equipment.
Inclusion of information on how to respond to a non-native species introduction within
protected area management plans.
Better use and resourcing of scientific expertise to inform our understanding of non-
native species issues and advise on response action.
Assuming there is the political will to uphold and apply the legislation agreed in the
Environmental Protocol, undertaking these activities will make financial sense. The most
cost-effective option is to implement measures to prevent introductions in the first instance.
Biodivers Conserv
123
However, should a non-native species establish, eradication as soon as possible after
introduction may still prove relatively inexpensive. Failure to take action, resulting in
species expansion, may render eradication practically impossible, leaving control of spe-
cies spread as the only form of mitigation—a long-term and potentially expensive
commitment.
Acknowledgments This review paper is a contribution to the SCAR AntEco (State of the Antarctic
Ecosystem) research programme. KH and PC are supported by NERC core funding to the British Antarctic
Survey’s Polar Science for Planet Earth core programmes ‘Environment Office—Long Term Monitoring
and Survey’ (EO-LTMS) and ‘Ecosystems’.
Open Access This article is distributed under the terms of the Creative Commons Attribution License
which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the
source are credited.
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... The continent of Antarctica and its offshore islands are today the part of the planet with the lowest presence and impact of terrestrial biological invasions globally (Hughes et al. 2015b). The near-pristine status of the Antarctic region underpins calls for strengthening its conservation management regime , Wauchope et al. 2019. ...
... Nevertheless, transplant experiments continued through the 1960s and early 1970s (Edwards 1980). Although such experiments appear to have ceased after that as environmental protection concerns rose (but see Braun et al. 2012, Hughes et al. 2015b, it was not until the negotiation of the Protocol on Environmental Protection to the Antarctic Treaty in 1991 -which formally came into force in 1998 -that strict regulations controlling any such deliberate introduction of non-native species were adopted, requiring permitting and confirmed removal of such material at the end of any experiments. This legislation has effectively banned deliberate introductions, but it remains unclear how it applies to accidental introductions associated with human activity (Hughes & Convey 2014) or to microorganisms (Hughes et al. 2015a). ...
... We know little to nothing of the introduction histories of non-native Collembola. Nonetheless, the high number of non-native species now known to occur on Deception Island , Hughes et al. 2015b, which includes Whalers Bayone of the most visited sites in Antarctica -further reinforces the probable importance of the connectivity hypothesis of non-native invasion pathways to Antarctica. In contrast, we have few clues on the origins of the non-native Acari (Russell et al. 2014). ...
Article
Understanding the success factors underlying each step in the process of biological invasion provides a robust foundation upon which to develop appropriate biosecurity measures. Insights into the processes occurring can be gained through clarifying the circumstances applying to non-native species that have arrived, established and, in some cases, successfully spread in terrestrial Antarctica. To date, examples include a small number of vascular plants and a greater diversity of invertebrates (including Diptera, Collembola, Acari and Oligochaeta), which share features of pre-adaptation to the environmental stresses experienced in Antarctica. In this synthesis, we examine multiple classic invasion science hypotheses that are widely considered to have relevance in invasion ecology and assess their utility in understanding the different invasion histories so far documented in the continent. All of these existing hypotheses appear relevant to some degree in explaining invasion processes in Antarctica. They are also relevant in understanding failed invasions and identifying barriers to invasion. However, the limited number of cases currently available constrains the possibility of establishing patterns and processes. To conclude, we discuss several new and emerging confirmatory methods as relevant tools to test and compare these hypotheses given the availability of appropriate sample sizes in the future.
... Deception Island (South Shetland Islands) unsurprisingly, the Antarctic Peninsula and off-shore islands at its northern tip have more recordings of marine (Table 2.1) and terrestrial (Hughes et al., 2015;McGeoch et al., 2015) non-native species than anywhere else in continental Antarctica. While this may reflect a sampling bias, it is nonetheless an area worthy of particular focus for a monitoring Tourist vessels operate from October to April with on average 4062 ship days per year and repeatedly target the same coastal areas (Bender et al., 2016;Lynch et al., 2010). ...
... Terrestrial Antarctica is predicted to become increasingly ice-free throughout the century, particularly on the northern Antarctic Peninsula in Antarctic Conservation Biogeographic Regions (ACBRs) 1, 2 and 3 (Lee et al., 2017). These regions, due to warmer temperatures, increasing water availability, increasing ice-free habitat and increased human visitation, have the highest risk of colonisation by non-native vascular plants (Chown et al., 2012;Hughes et al., 2015). In the marine realm, since 1957, sea-ice extent and concentration in Antarctica increased in some areas while decreasing in others (Gutt et al., 2015;Steig et al., 2009), particularly the western Antarctic Peninsula (Steig et al., 2009). ...
... Here we outline the main regulatory frameworks relevant to NNMS in Antarctica (Figure 2.6). concern has grown over biosecurity measures in place for Antarctica, particularly for terrestrial environments (Chown et al., 2012;Hughes et al., 2011Hughes et al., , 2015Hughes & Convey, 2010Lewis et al., 2003;McGeoch et al., 2015) and a decisionmaking process for dealing with suspected introduced species has been formulated (Hughes & Convey, 2012). While response plans for eradication and management of nonnative terrestrial species are in development, the limited information about non-native marine species has created difficulties for the development of marine-focused policies (Hughes & Pertierra, 2016). ...
Thesis
Full-text available
Invasive non-native species are a major threat to global biodiversity. For at least 15 million years coastal Antarctica has been poorly connected to nearby temperate ecosystems due to physical and physiological barriers. Yet, Antarctica is experiencing significant environmental change and becoming increasingly exposed to ship-borne human activity that crosses the physical barriers. These factors may facilitate the establishment of non-native marine species. This doctoral research adds insight into the risk of non-native marine species being transported to Antarctica via ships’ hulls and internal seawater systems, with particular focus on pathways of introduction and species found within those pathways. To begin my research, I assessed the current knowledge of non-native marine species in the Antarctic region: the physical and physiological factors that resist establishment of non-native marine species; changes to resistance under climate change; the role of legislation in limiting marine introductions; and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non-native marine species was limited: up to 2019 just four marine non-native and one cryptogenic species that were likely introduced anthropogenically had been reported free-living in Antarctica or in the sub-Antarctica islands, but no established populations have been reported. An additional six species had been observed in pathways to Antarctica that are potentially at risk of becoming invasive. I estimated there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are not recorded comprehensively. In response to the scarcity of data on ship movements into the Southern Ocean, I obtained data on ship activity in the Southern Ocean from 2014-2018 inclusive and developed a ship traffic network for Antarctic-going vessels. I analysed the ship movements and conducted a spatially-explicit assessment of introduction risk for non-native marine species in all Antarctic waters. I found that vessels connect Antarctica via an extensive network of ship activity to all global regions, and especially South Atlantic and European ports. Ship visits were more than seven times higher to the Antarctic Peninsula and the South Shetland Islands than elsewhere around Antarctica. I found that, while the five recognised ‘Antarctic Gateway cities’ are important last ports of call, an additional 53 ports had vessels directly departing to Antarctica from 2014-2018. I identified ports outside Antarctica where biosecurity interventions could be most effective and the most vulnerable Antarctic locations where monitoring programmes for high-risk invaders should be established. Biofouling communities within the major pathway to Antarctica from Europe via the South Atlantic, identified in the network analysis, became my next focus. I obtained biofouling samples from the polar research vessel RRS James Clark Ross and found that niche (protected) areas of the hull represent significantly greater colonisation (species richness) and propagule pressure (individual abundance) than exposed areas of the hull. The composition of the biological communities did not differ among exposed and niche areas, but did change significantly among the three surveys conducted. Only six species were found on the ship’s hull in Antarctica, but they included a known invasive bryozoan, Tricellaria inopinata, and barnacles that have no counterparts in Antarctica. While the role of hull fouling is recognised as a globally important vector for introductions of non-native marine species, the role of a vessel’s internal pipework has been overlooked. I conducted the first comprehensive study of biofouling macrofauna living inside an Antarctic vessel’s internal seawater systems, finding breeding communities of Jassa marmorata (Amphipoda) and mytilid mussels throughout the internal pipework system. I found fouling communities that occluded ~9-17% of a pipe’s cross-sectional area, increasing running costs for ships. Since ships are constantly pumping their water through their pipework, they are likely to be releasing propagules at all stages of their voyages, including in polar regions. Before I started my research, Antarctic operators and policy-makers were unaware of the total number of vessels that visit Antarctica. Now, I have provided comprehensive insight into the most traversed routes to Antarctica and identified Antarctic locations that are the most likely recipient locations for non-native marine species. I found that non-native species from temperate regions can survive passages through polar areas and that sheltered sections of the hull and internal systems are especially important sites for both propagule and colonisation pressure. Together, these results demonstrate that Antarctica is well connected to worldwide marine ecosystems and that biofouling on ships poses an important and growing introduction risk to Antarctica.
... To mechanistically forecast Antarctic biodiversity will require a renewed effort to target basic natural history information and fill many of these data gaps. One particularly important threat to Antarctic biodiversity is the increasing rates of non-native species arrival and establishment [68]. Thus, incorporating novel biotic interactions ( Figure IA) into models will be increasingly important to predict the potential effects of invaders on resident biodiversity. ...
Article
Antarctic ecosystems are under increasing anthropogenic pressure, but efforts to predict the responses of Antarctic biodiversity to environmental change are hindered by considerable data challenges. Here, we illustrate how novel data capture technologies provide exciting opportunities to sample Antarctic biodiversity at wider spatiotemporal scales. Data integration frameworks, such as point process and hierarchical models, can mitigate weaknesses in individual data sets, improving confidence in their predictions. Increasing process knowledge in models is imperative to achieving improved forecasts of Antarctic biodiversity, which can be attained for data-limited species using hybrid modelling frameworks. Leveraging these state-of-the-art tools will help to overcome many of the data scarcity challenges presented by the remoteness of Antarctica, enabling more robust forecasts both near- and long-term.
... One of the most concerning threats to Antarctic ecosystems and biodiversity, as is the case globally (Pyšek et al. 2020), is the introduction through anthropogenic assistance of new species (potentially including some providing new ecological functions) that may be able to become established in the region, with unknown and unpredicted consequences for the native biota (Frenot et al. 2005, Amesbury et al. 2017, Robinson et al. 2018, Convey & Peck 2019. Many examples of this already exist in the sub-Antarctic, with increasing numbers of records becoming available from the Maritime Antarctic (Frenot et al. 2005, Hughes et al. 2015. Some of these were clearly able to become established in parts of Antarctica under pre-warming or current conditions, but the predicted continued warming is likely to act in synergy with increasing levels of human activity to increase the risks of both transfer events and successful establishment (Convey & Peck 2019). ...
Article
Full-text available
We report the first formal record of the Indian meal moth Plodia interpunctella from a location within the Antarctic Treaty area, with the capture of a live adult male within the Brazilian Comandante Ferraz research station on King George Island, South Shetland Islands. This species is a well-known pest of stored products and is widely recorded in synanthropic situations such as food stores globally. No other adults or immature stages have been observed on the station. While there is no suggestion that P. interpunctella could survive or establish in the natural environment beyond the station, this observation highlights the ever-present threat of unintended anthropogenically assisted transfer of non-Antarctic species into human facilities on the continent, with some such species proving extremely difficult to eradicate if they successfully establish within these facilities.
... However, the exact impacts of habitat expansion on Antarctic soil biology are largely unknown. This uncertainty is reflected in the current environmental policy situation, with no internationally agreed response strategy for non-native species colonisation (Hughes et al., 2015). Therefore, as human activity and movement around the continent and between Antarctic and sub-Antarctic biogeographic regions continues to increase, with that comes the risk of intra-and inter-regional transfer of native and non-native species. ...
Article
Ice-free regions consist of less than 0.5% of the total area of Antarctica, but the soils that have evolved on these areas are of very significant scientific value as they concentrate most of the terrestrial biodiversity on the continent. In this paper, we discuss four main research priorities for soil science in Antarctica. These include: Bockheim (2015) improving information on soil properties; Brooks et al. (2019) quantifying and monitoring the impacts of climate change; Convey et al. (2014) develop an assessment of soil quality; and (Cox and Smith, 2019) improve the way soil science feeds into environmental policy in Antarctica.
... As a consequence of increasing human pressure, the Antarctic region has progressively become less isolated and more affected by human footprint (Chu et al., 2019;Joblin, 2020). Among others, the rising maritime traffic has resulted in an increasing risk of introducing non-native species to the Southern Ocean (Bender et al., 2016;Hughes & Convey, 2010), as already reported for terrestrial (e.g. the bluegrass Poa annua, the brachypterous chironomid Eretmoptera murphyi or the enchytraeid worm Christensenidrilus blocki, Hughes & Convey, 2010, Chown et al., 2012, Chwedorzewska et al., 2015 and marine environments (e.g. the seaweed Ulva intestinalis, the crab (Hughes et al., 2015;Lee & Chown, 2009b;Volonterio et al., 2013). ...
Article
Full-text available
Aim The Western Antarctic Peninsula is challenged by climate change and increasing maritime traffic that together facilitate the introduction of marine non‐native species from warmer regions neighbouring the Southern Ocean. Ballast water exchange has been frequently reported as an introduction vector. This study uses a Lagrangian approach to model the passive drift of virtual propagules departing from Ballast water hypothetic exchange zones, at contrasting distances from the coasts. Location Western Antarctic Peninsula. Methods Virtual propagules were released over the 2008–2016 period and at three distances from the nearest coasts: 200 (convention for the management of Ballast Water, 2004), 50 or 11 nautical miles (NM). Results Results show that exchanging Ballast water at 200 NM considerably reduces the arrival of propagules in proposed marine protected areas of the western side of the Antarctic Peninsula. On the eastern side, propagules can reach north‐eastern marine protected areas within a few days due to strong currents for all tested scenarios. Seasonal and yearly variations indicate that exceptional climate events could influence the trajectory of particles in the region. Ballast water should be exchanged at least 200 NM offshore on the western side of the Antarctic Peninsula and avoided on the eastern side to limit particle arrival in proposed marine protected areas. Focusing on Deception Island, our results suggested that the Patagonian crab (Halicarcinus planatus) observed in 2010 could have been introduced in case of Ballast water exchange at 50 NM or less from the coast. Main conclusions This study highlights the importance of respecting Ballast water exchange convention to limit the risk of non‐native species introduction. Ballast water exchange should be operated at least at 200 NM from the coasts, which further limits particle arrival in shallow water areas. This is especially important in the context of a more visited and warmer Southern Ocean.
... Endemic island populations, of both plants and animals, are generally highly adapted and therefore often lack the defence mechanisms to resist an invasive competitor (Convey 1996). The introduction of a non-native species, even unintentionally, has irreversible ecological impacts (Hughes et al. 2015). The introduction and rapid, pervasive colonisation of South Georgia by the invasive flora is one such example. ...
... Endemic island populations, of both plants and animals, are generally highly adapted and therefore often lack the defence mechanisms to resist an invasive competitor (Convey 1996). The introduction of a non-native species, even unintentionally, has irreversible ecological impacts (Hughes et al. 2015). The introduction and rapid, pervasive colonisation of South Georgia by the invasive flora is one such example. ...
Book
Antarcticness joins disciplines, communication approaches and ideas to explore meanings and depictions of Antarctica. Personal and professional words in poetry and prose, plus images, present and represent Antarctica, as presumed and as imagined, alongside what is experienced around the continent and by those watching from afar. These understandings explain how the Antarctic is viewed and managed while identifying aspects which should be more prominent in policy and practice. The authors and artists place Antarctica, and the perceptions and knowledge through Antarcticness, within inspirations and imaginations, without losing sight of the multiple interests pushing the continent’s governance as it goes through rapid political and environmental changes. Given the diversity and disparity of the influences and changes, the book’s contributions connect to provide a more coherent and encompassing perspective of how society views Antarctica, scientifically and artistically, and what the continent provides and could provide politically, culturally and environmentally. Offering original research, art and interpretations of different experiences and explorations of Antarctica, explanations meld with narratives while academic analyses overlap with first-hand experiences of what Antarctica does and does not – could and could not – bring to the world.
Technical Report
Full-text available
Antarctica and the surrounding Southern Ocean are under increasing pressure from cumulative impacts of climate change, pollution, fisheries, tourism and a variety of other human activities. These changes pose a high risk both to local polar ecosystems and to the regulation of the global climate, as well as through global sea-level rise. Thus, long-term monitoring programmes serve to assess the state of ecosystems as well as to make projections for future developments. The Fildes Region in the southwest King George Islands (South Shetland Islands, Maritime Antarctica), consisting of the Fildes Peninsula, Ardley Island and several offshore islands, is one of the largest ice-free areas in the Maritime Antarctic. As a continuation of a long-term monitoring programme started in the 1980s, local breeding bird and seal populations were recorded during the summer months (December, January, February) of the 2018/19 and 2019/20 seasons and supplemented by individual count data for the 2020/21 season. This study presents the results obtained, including the population development of the local breeding birds. Here, some species showed stable populations in a long-term comparison (brown skuas, southern polar skuas) or a significant increase (gentoo penguin, southern giant petrel). Other species, however, recorded significant declines in breeding pair numbers (Adélie penguin, chinstrap cenguin, Antarctic tern, kelp gull) up to an almost complete disappearance from the breeding area (cape petrel). In addition, the number of seals at their haul-out sites was recorded and the distribution of all seal reproduction sites in the Fildes Region was presented. Furthermore, data on the breeding bird population in selected areas of Maxwell Bay were added. Additionally, the rapid expansion of the Antarctic hairgrass was documented with the help of a completed repeat mapping. The documentation of glacier retreat areas of selected areas of Maxwell Bay was updated using satellite imagery and considered in relation to regional climatic development. Furthermore, the distribution and amount of marine debris washed up in the Fildes Region and the impact of anthropogenic material on seabirds will are addressed. In addition, the current knowledge of all introduced non-native species in the study area and the need for further research are presented.
Article
Full-text available
Poa annua L. is the only non-native vascular plant that was successfully established in the maritime Antarctic. This project aimed to determine the amount of genetic and epigenetic variation within and between two populations of P. annua, one from South Shetland Is. (Antarctic) and the other one from Central Europe. We applied two AFLP marker systems, using endonucleases that recognised the same restriction site but differed in their sensitivity towards methylation. The Antarctic population differed from the Polish one both at the genetic and epigenetic levels. Genetic variability in the Antarctic population was lower than in the Polish one. Some loci in the Antarctic population showed signs of selection. The difference between Polish and Antarctic populations might be due to a weak bottleneck effect followed by population expansion. Using only epigenetic markers, the Antarctic population exhibited increased variation level compared to the Polish one. These may have resulted from plastic responses to environmental factors and could be associated with survival in extreme conditions.