zoological journal
of the Linnean
Society (1994), 110: 207-217
Non-indigenous Acari of Antarctica and the
sub-Antarctic islands
P. J. A. PUGH, F.L.S.
British Antarctic Survey, JVatural Environment Research Council, High Cross,
Madingley Road, Cambridge C B 3 OET, U.X.
Received March 1993, revised and accepted M a y 1993
Approximately 70 species out of a total of more than 520 Acari recorded from Antarctica and the
sub-Antarctic islands may originate from other continents, especially Australasia, South America
and Europe. Although some species have probably been carried into the region on migrant birds,
most may have been introduced as a result of human activity, in particular by whalers and sealers.
The majority of species appear to originate from imported sheep, rabbits, rats and fowl, and a few
from vegetation, soil and ship’s stores.
ADDITIONAL’KEY WORDS:-Introduced
species - islands - mites - Southern Ocean.
CONTENTS
Introduction . . . .
Discussion
. . . .
The Antarctic biota .
Dispersal. . . .
Survival of alien species
Adaptations . . .
Control of alien species
Acknowledgements . .
References . . . .
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INTRODUCTION
Many animal and plant species occurring on mainland Antarctica and the
sub-Antarctic islands are not indigenous and have been introduced either
accidentally or deliberately as a result of human activity. The majority of studies
o n Antarctic and sub-Antarctic introduced or ‘alien’ organisms have
concentrated on flowering plants, birds and mammais. There is little information
on insects and virtually nothing on Acari, which is one of the largest and most
important groups within the region (Pugh, 1993).
T h e present study considers how alien Acari may have been introduced into
the Antarctic and sub-Antarctic regions by examining dispersal of other
Antarctic biota and the dispersal of Acari in other regions of the world.
0024-4CW4/94/030207
+ 1 1 808.00/0
207
@ 1994 The Lnnean Society of London
208
P. J. A. PUGH
DISCUSSION
The Antarctic biota
Compared with adjacent continents, the Antarctic biota is depauperate and
comprises an indigenous component and a more recent and numerically
dominant introduced one (Ellis-Evans & Walton, 1990).
The indigenous biota
This is composed of pre-glacial relict species and post-glacial immigrants.
Relict species. Geological evidence suggests that Antarctica was once joined to
South America and Australasia, and this hypothesis is supported by the
similarity in the present collembolan and oribatid mite faunas of these three
continents (Wallwork, 1973). As Antarctica moved south and became
increasingly colder, glaciated and isolated, the majority of species became
extinct, although a few adapted and survived around ice-free nunataks (EllisEvans & Walton, 1990), as relict representatives of the original Gondwanan
biota.
Immigrant species. The alternative hypothesis is that all the original ‘relict’
Antarctic biota were killed by low temperature and loss of habitat; subsequent
recolonization was by post-Pleistocene immigrants from adjacent temperate
continental South America and Australasia (Gressitt, 1961, 1967).
The introduced biota
This may form a significant proportion of the fauna and flora, especially on
sub-Antartic islands. Nine of the 26 insect species on Marion and Prince Edward
Islands are introduced (Crafford et al., 1986) and although the two islands have
slightly different faunas, there is, because of their proximity, the potential for
continual interchange of species between them (Crafford & Scholtz, 1987).
Further study of Southern Ocean island biota may reveal that, as is the case on
some non-polar oceanic islands, few species are truly indigenous and dispersal of
‘aliens’ is more widespread than currently realized (Schatz, 1991:I.
Introduced Acari
Between 50 and 80 species of terrestrial Acari recorded from the Southern
Ocean are probably introduced (Table 1). These species cover a broad
taxonomic range, though there is difficulty in determining the status of some
species, particularly those for which there is only one record. Despite the number
of recorded alien Antarctic and sub-Antarctic Acari there is little information
regarding their dispersal, survival and adaptations. Considerably more data exist
for non-indigenous plants, birds, mammals and insects, many of which may be
means of introduction of, or suitable models for comparison with, the introduced
Acari.
Dispersal
Invertebrates (and plants) are dispersed throughout the Southern Ocean by
mechanisms beyond the control of the individual organisms, and as a
consequence their arrival at any site, especially in the more southerly continental
regions, is a rare event (Ellis-Evans & Walton, 1990). Drifting wood and other
ANTARCTIC ACARI
209
plant material, wind and ocean currents, and transport by other animals,
including man, are the major routes by which plants and invertebrates have
colonized temperate and tropical oceanic islands ( Palmen, 1944; Gressitt, 1964;
Lindroth, 1970; Hammer, 1982; Schatz, 1991). Such mechanisms must be
considered with reference to the Antarctic and sub-Antarctic faunas.
Dispersal by water and wind
Drift wood is a very improbable means of transport within the Southern
Ocean (Gressitt et al., 1960; van Zinderen-Bakker, 1971), and although wind has
been shown to transport some plant propagules from South America to some
parts of the Antarctic Peninsula (Smith, 1991), and large numbers of air-borne
drifting insects have been collected far out at sea (Bowden &Johnston, 1976),
prevailing wind direction can be a major dispersal barrier for some islands
(Heywood, 1977).
‘Ballooning’ or ‘parachuting’ juvenile spiders are common features of the
‘aerial plankton’ and some species have been recorded at considerable distances
from land (Gressitt & Yoshimoto, 1963). Erigone autumnalis Emerton, a native of
North America, was found ballooning on the Ross Sea coast, continental
Antarctica, and may originally have been imported to the nearby McMurdo
base (Forster, 1971). Fragments of another spider, probably belonging to the
family Micryphantidae, were also found near the Ross Sea coast, and are
attributed to a cast exuvium that drifted into the region on air currents (Forster,
1970).
Evidence from temperate and tropical isolated island groups suggests that
Acari are probably not distributed over long distances by ocean currents (Jacot,
1934), and air currents appear to be equally unlikely. This leaves transport by
other animals as the most probable mechanism of dispersal for the majority of
non-indigenous or alien mite species.
Dispersal by other animals
Alien birds and mammals have been deliberately introduced to Crozet,
Heard, Kerguelen, Macquarie, Marion, South Georgia, the South Shetland and
other islands by sealers, whalers and research station personnel (Law & Burstall,
1956; Van Zinderen-Bakker, 1971; Headland, 1984; Leader-Williams, 1988).
Introduced birds and mammals have probably carried with them numerous
alien Acari (Tables 1 and 2); indeed their introduction is an unavoidable
consequence of human colonization (Schatz, 1991). The more northerly islands
of South Georgia, Marion, Campbell and Macquarie, which have a long history
of commercial exploitation and habitation by sealers and whalers (Headland,
1989), have the most numerous records of alien mite species (Table 1).
Europeans were largely responsible for the manning and maintenance of the
majority of the early sealing and whaling stations within the Weddell Sea,
Atlantic and Indian sectors of the Southern Ocean, and conducted much of the
research within these regions. Records of alien species in these areas are mainly
of European or cosmopolitan distribution (Table 1). Similarly, the alien Acari
(and other groups) on Macquarie Island probably originate from Australia,
under whose jurisdiction the island falls. Yet, with the possible exception of
South Georgia, none of the island groups shows any particular affinity with
South America, and there are no species within the Southern Ocean of definite
P. J. A. PUGH
210
TABLE
1 . Records of all known and suspected non-indigenous Acari from the Southern Ocean,
mentioned in the literature. Key to area records:
Mainland Antarctica
A. Maud Sector
B. Enderby Sector
C. Wilkes Sector
D. Scott Sector
E. Byrd Sector
F. Ronne Sector
G. Palmer Land
H. Graham Land
Family
Southern Ocean Island groups
-
1
3
3
-
I. South Shetland Is.
J. South Orkney Is.
K . South Sandwich Is.
L. South Georgia
M. Bouvetaya
N. MarioniPrince Edward Is.
0. Iles Crozet
1
5
3
9
-
1
P. Iles Kerguelen
Q. Heard Is.
R. Macquarie Is.
S. Campbell 1s.
T. Ballany/Scott Is.
U. Peter I 0 y
7
~~
45
12
~
-
2
2
Species
A. Mesostigmata ( = Gamasidaj
Ichthyostomatogasteridae Asfernolaelaps sp.
Parasitidae
Eugamasus sp. (5)
Pergamasus sp. (2)
Veigaiidae
Veigaia claggi Hunter
Veigaiidae (gen. et sp.)
Rhodacaridae
Gamasellus sp.
Iphidozercon sp.
Ascidae
Lasioseius sp.
Phytoseiidae
Sejulus plumosus Oudemans
Eviphidiidae
Alliphis siculus (Oudemans)
Thinoseius hirschmanni
Hunter
Pachyseius adeliensis
Pachylaelapidae
Womersley
Ayersacarus plumapilis
Hunter
Laelapidae
Eulaelaps sfabularis (Koch)
Hypoaspis euansi (Hunter)
Haemogamasus pontiger
Berlese
Laelaps grahamensis
( Womersley)
Leplolaelaps macquariensis
(Worn.)
sp. # I
sp. # 2
Rhinonyssus rhinolethrum
Rhinonyssidae
(Trouess.)
Cercomegistidae
sp. # 1
sp. # 2 (Celaenogamasus sp. J
Polyaspididae
SP.
Piracarus crozefeasis Richters
Uropodidae
B. Prostigmata ( = Ac&edida)
Penthaleidae
Halotydeus sp.
Penthaleus major (Duges)
Tydeidae
Ereynetoides wafsoni Fain
LorTia lepfonycholes
Womersley
L . polaris Womersley
Tydeus sp.
Pvgmephoridae
Neopygmephorus arvorum
(Jacot)
JV. panonicus (Willmann)
JV. sellnicki (Krczalj
N. togatus (Willmannj
Recorded habitat
Nest of blue petrel
All from vegetation litter
From vegetation litter
White-chinned petrel nest
No data
No data
Penguin nests, vegetation
No data
Moss
Sheepdung nr. station
Under rocks on shore
Sites
Other records
N
Rl
R'
L
L
K
R'
J
0
RS"
L
Washed out to sea
C
Fowl debris, bird nests
RS"
Eur.
Fowl debris
S
-
R
O n rats. food and in nests
R
Under stones
H
Between tidemarks
R
Ectoparasite of rabbits
Ectoparasite of rabbits
O n ducks
R
R
L
Among vegetation
Among vegetation
Soil around roots of Poa
Washed out to sea
R
R
R'
C
O n sea-sprayed rocks
Among grass by a stream
In leaf litter
In gut of seal (dislodged?)
R'
Washed out to sea
Grassland
Litter and soil
D
R
N.Am. Eur
Vegetation and soil
Vegetation and soil
Vegetation and soil
R
Aust.
Eur. Asia
Eur.
S
R
D
cosmo.
cosmo.
cosmo.
cosmo.
R
R'
R
ANTARCTIC ACARI
21 1
Table 1-continued
Family
Cheyletidae
Cheyletiellidae
Raphignathidae
Stigmaeidae
Tetranychidae
Trombiculidae
Trombidiidae
Species
Recorded habitat
Acaropsis sp.
Cheyletus eruditus (Schrank)
c. sp.
Cheyleteiella parasitivorax
(Megnin)
Raphigna fhusjohnstoni
Womersley
Six different species
Bryobia praetiosa
(C.L. Koch)
Bryobia sp. (praefiosa?)
7etranychus sp.*
Eutrombicula hirsti
(Sambon)
Microtrombidium kaniensis
Womersley
-
c. Cryptostigmata ( = Oribatida) and Astigmata (
Andacarus watsoni Trave
Archaekoth;idae
Eobranchychthoniu oudemansi
Brachychthoniidae
v.d. Hammen
Camisiidae
Camisia segnis (Hermann)
Gressittoppia pe$fensis
Oppiidae
Hammer
Edwardzetes denfifcr
Ceratozetidae
Hammer
7ofobates elegans Hammer
Oribatulidae
Galumnidae
Galumna alata Hermann
Acaridae
Acarus sir0 Linnaeus
Aleurobius farinae DeCeer
Rhizoglyphus echinopus Fum.
& Robin
R . sp. (echinopus?)
lyiophagus casfelini Hirst
7.longior (Gervais)
Carpogly phidae
GIycyphagidae
Hyadesiidae
Histiostomatidae
Listrophoridae
-
Stored food and debris
-
Ectoparasites of rabbits
Sites
J
R'
R
C-D
On vegetation
On shore under stones
S'
LP
Bird nests and moss
cosmo.
J
In gut of seal (dislodged?)
From vegetation in pool
Ectoparasite of rabbit
Other records
St. Paul, Eur.
Arctic
0
L
PR
S
Aust.
= Acarida)
Moss
On vegetation
RS"
K
S.Am. Eur.
Moss
R
I
S.Am. Eur.
S.Am.
K
S.Am.
On terrestrial algae
-
Under stones in a pool
Stores, nests, litter
Ship's stores only
Ship's stores
Yellow lichens
Ectoparasites of rabbits
Fowl house and rat debris,
grain stores
7.putrescentiae (Schrank) Stores, inc. ships
No data
Carpoglyphus neglectus
Cambridge
c. sp.
Hay, dung, nest debris
Caloglyphus sp.
Fowl-house debris
No data
Ghyphagus destructor
(Schrank)
G. domesticus (de Geer) Fowl-house debris, bird
nests
Ship's stores
G . spinipes (Koch)
G . sp.
No data
Schwebia talpa Oudemans Bird nests
On coastal rocks
Hvadesia SD.
Bonomia or Spinanoefus sp.?* Vegetation litter
Wichmannia sp.?*
Bird nests
Listrophorus gibbus
On rabbits
(Pagenstec her)
RL'
LP
R
LR
Eur.
cosmo.
cosmo.
cosmo.
L
cosmo.
R'
B-CR'
cosmo.
JK
Eur.
P
R
R'
J
R
-
R
R
R'
R
R
R
Figures show number of known 'alien' species recorded for each area.
Key to other records--Aust.: Australasia, cosmo.: cosmopolitan, Eur.: Europe; N.Am.: North America;
%Am. South America.
Other notes--*: not given in Pugh (1993);c: contamination in transit; e: not well established; i: quoted as an
indigenous species, but based on only one record.
212
P. J. A. PUGH
African, Asian or North American origin, which reflects the comparatively
minor involvement of people of these continents in the exploitation of the subAntarctic islands.
The acarine groups with parasiticlphoretic larvae and/or nymphs, namely the
families Parasitidae, Uropodidae, Trombididae, Erythraeidae, and Acaridae,
were probably introduced attached to alien insect hosts. Some free-living alien
mites were probably imported on wood, agricultural products, packing material,
livestock fodder, food and other ship’s stores, but the majority of species are
associated with either (a) livestock, particularly sheep, rabbits and chickens, or
(b) pests, especially rats and rabbits (Tables 1 and 2).
The obligate parasites of birds and mammals, notably the ticks (Metastigmata
or Ixodida) and the feather mites (Sarcoptiform Astigmata), have been omitted
from the present study, as many are dispersed by native and/or migrant birds
and cannot be distinguished from the truly introduced species. Some other
parasitic arthropods were certainly deliberately introduced to the sub-Antarctic
islands, for example mammal fleas (Hexapoda: Siphonaptera) to Macquarie
Island (Skira et al., 1983). One species in particular, namely Spilopsyllus culiculi
(Dale) was introduced as a vector for myxamotosis, in an attempt to control the
introduced rabbit population (Sobey et al., 1973).
The oribatid mites Eobrachychthonius oudemansi van der Hammen, Camisia segnis
(Hermann), Gressittoppia pepitensis (Hammer) and Edwardzetes dentzfer Hammer
may have been accidentally introduced to the sub-antarctic islands from South
America (Wallwork, 1973). The most likely route for these species is on imported
agricultural products, because Humerobates rostrolamellatus (Grandjean) and
Galumna australis (Berlese) were introduced to Hawaii from Europe and South
America respectively in the same manner (Jacot, 1934). Oribatids are
phytophages, or more usually mycetophages, i.e. feed upon fungi that develop on
decomposing vegetation, and are thus unlikely to have been introduced with dry
seeds.
Although some alien plants may have been introduced to the sub-Antarctic as
seeds in livestock dung (Headland, 1984), and some Astigmata may survive
passage through the vertebrate gut (Chmielewski, 1970), it is unlikely that alien
oribatids have been imported in this way. Direct introduction with dry hay and
straw fodder is also improbable, and a more likely route is with live rooted
plants, soil or leaf litter, all of which have been imported to South Georgia for
use in greenhouses and cold-frames (R. I. L. Smith, personal communication).
Bark-clad timber was also shipped to the whaling stations on South Georgia
from the U.K. and Norway (R. I. L. Smith, personal communication), and as
wood bark is a suitable habitat for several oribatid mites, including C. segnis
(S. Seniczak, personal communication), this and other oribatids may have been
introduced on such timber. The few records of potential alien oribatids indicates
that these mites are not easily distributed by man.
Suruiual of alien species
T o survive, aliens must not only arrive by suitable transport, but also find an
acceptable new microhabitat in which they can grow and reproduce successfully.
Alien plants and animals have become established on sub-Antarctic islands
(Holdgate, 1964; Edwards & Greene, 1973; Jenkin et al., 1982; Masse, 1982;
ANTARCTIC ACARI
213
TABLE
2. Habitat records for possible non-indigenous Acari in the Antarctic and sub-Antarctic
Category
Details of records
Plants
On live vegetation
Litter, plant debris and soil
Birds and mammals
Fowls and in fowl-house litter
Other birds (excluding Sarcoptiform Astigmata)
Wild bird nests
Rats
Rabbits
Sheep dung
Number of
species
17
14
5
4
10
1
5
1
Stores
On imported stores and debris
In ship’s stores
6
4
No precise data
Under stones
On shore
3
4
Others
‘Washed out to sea’
In gut of seal
No habitat data
3
2
12
Headland, 1984; Smith, 1984), especially Marion, Campbell and Maquarie,
which have comparatively mild climates and a large number of potential niches.
By contrast, extensive glaciation on South Georgia has resulted in ice-free areas
(and possible microhabitats) being widely separated, forming a ‘biological
archipelago’ (Leader-Williams, 1988), deterring the establishment of many
species. Few alien species have become established in the Antarctic because of
conditions imposed by the severe climate and consequent lack of niches
(Edwards, 1974; Block et al., 1984; Smith, 1984).
The status of alien animal species may be considered at two levels, namely
(1) ‘transient aliens’, which either fail to breed or cannot survive beyond the
immediate confines of human habitation or although able to endure average
winters are killed during an exceptional one, and (2) established or ‘naturalized
aliens’, which find a niche in the natural environment and compete successfully
with the endemic biota and breed in the natural environment.
Transient aliens
The majority of transient aliens survive in heated buildings, living on stored
food, etc., and are encouraged by accumulations of detritus, spillage and
spoilage of food material (Rounsevell, 1978). Transient aliens include spiders
(Araneae), weevils and ladybirds (Coleoptera), flies and mosquitoes (Diptera), a
lacewing (Neuroptera), cockroaches (Dictyoptera) and mites (Acarina) (Hagen,
1876; Cameron, 1972; Rounsevell, 1978; Headland, 1984; R. I. L. Smith,
personal communication).
The german cockroach Blatella germanica L., introduced to Grytviken whaling
station and probably other stations on South Georgia on foodstuffs imported
from South America, was a continual problem during the whaling era, but died
out when the station was abandoned (Headland, 1984). The facultative acarine
parasite Haemogamasus pontiger Berlese (Mesostigmata) is often associated with
human activity and has been reported in buildings at Davis Station, Enderby
214
P. J. A. PUGH
Sector on continental Antarctica, where it was introduced along with a beetle
(Rounsevell, 1978). This mite is naturalized on Macquarie Island, where it was
probably introduced by the early nineteenth century ‘oil gangs’ (elephant
sealers) (Rounsevell, 1978). An additional 30 or so poorly established alien
species have been found close to the settlement/research station on Macquarie
Island, associated with non-indigenous birds, sheep dung, stored food and grain
(Watson, 1967).
Naturalized aliens
Temperate free-living alien insects, including a lepidopteran, have become
established on Kerguelen and Marion Islands (Kidder, 1876; Crafford &
Chown, 1987) and in soil around Grytviken whaling station on South Georgia
(Vogel & Nicolai, 1983). However, few alien species have become adapted to the
Antarctic environment (Balfour-Browne & Tilbrook, 1966; Rounsevell, 1978;
Goddard, 1979). Notable examples include the midge Eretmoptera murphyi
Schaeffer (Diptera), an alien but unknown enchytraeid, introduced to Signy
Island in the South Orkney Islands, in soil attached to plants from South
Georgia or the Falkland Islands in 1967 (Burn, 1982; Block et al., 1984; Block &
Christensen, 1985). There are no naturalized alien Acari recorded in Antarctica,
though there are numerous known from several sub-Antarctic islands (Table 1 ) .
Adaptations
Evidence from the Kerguelen Islands suggests that native fleas have adapted
to introduced hosts, and that alien fleas from introduced mammals now occur on
other introduced and native host species (Smit, 1957). This is a comparatively
minor adaptation and a number of ‘naturalized aliens’, including Acari on subAntarctic islands, have undergone changes in behaviour or feeding strategies to
adapt to new habitats (Rounsevell, 1978; Block et al., 1984). The ‘selected’
habitats are, however, similar in nature to those on which they were probably
imported. For example, species introduced on sheep dung, hay and other fodder
have inhabited bird-nesting material, vegetable matter and litter, whilst some
facultative fowl parasites have adapted to a variety of mammal and bird nests as
well as debris in human habitation.
Some alien species are physiologically pre-adapted to the sub-Antarctic; for
example, the lepidopteran Agrotis ipsilon (Hufnagel),a transient alien recorded at
Bird Island, South Georgia, is considered to be cold-tolerant (Bonner & Honey,
1987). However, the most important pre-adaptations exhibited by naturalized
alien insects are those concerned with the life-cycle, for example parthenogenesis
and/or protracted larval development (Block et al., 1984; Crafford, 1986; Schatz,
1991; Norton & Palmer, 1991).
Control of alien species
The majority of introduced bird and mammal species have died out or have
been deliberately exterminated on most sub-Antarctic islands, though rats and,
to a lesser extent, mice, cats and rabbits remain a problem on some, and there is
a thriving population of reindeer on South Georgia (Headland, 1984;
Leader-Williams, 1988). Although a number of large domestic mammals have
ANTARCTIC ACARI
215
been introduced to the Kerguelen Islands, including reindeer, sheep, mouflon
and rabbits, only the sheep are subject to any form of management (Decante,
1987). The eradication or at least the control of surviving alien mammal
populations is being considered for a number of islands, and such measures have
been incorporated into current long-term management strategies (Bloomer &
Bester, 1982; Clark & Dingwall, 1985; Decante, 1987; Hunter, 1990). However,
sheep will be maintained on Campbell Island (Anon., 1983), where the recent
arrival of new alien bird species will be monitored before future control measures
are decided (Anon., 1991).
Mites have never been deliberately introduced to the Antarctic or subAntarctic and all alien species have been accidentally imported on other
material. The strict control of alien birds and mammals on sub-Antarctic islands
is the most obvious means of preventing the introduction of such species.
Although alien mites have been recorded and continue to be collected from ships
and bases on continental Antarctica, they have never survived beyond the
confines of human habitation and present neither threat nor problem. The most
likely source of potential alien mites is from imported soil, for example that on
the wheels and tracks of vehicles shipped to research stations (R. I . L. Smith,
personal communication). Such soil should be removed a t source. I n addition, at
Christmas 1991, the troops at the military garrison on South Georgia were
‘issued’ with young ‘Christmas trees’ in pots, a number of which were
subsequently planted around the garrison (H. A. MacAlister, personal
communication). The trees and their organisms in their potting material are
certainly alien to the sub-Antarctic. Similar introductions to the region must be
discouraged, indeed there is legislation against it in the Antarctic Treaty
(Annexe 2, Article A, Appendix B and C).
ACKNOWLEDGEMENTS
I would like to thank the British Antarctic Survey for providing support. I
am indebted to Dr W. Block for his active encouragement and Drs R. I. L.
Smith, J. Shears and M r H. MacAlister, of B.A.S.; M r R. Headland of the Scott
Polar Research Institute, Cambridge; Dr H. Schatz, Institut fur Zoologie,
Innsbruck, Austria, and D r S. Seniczak of the Dept. of Animal Ecology,
Academy of Technology and Agriculture, Mazowiecka, Poland, for their advice.
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