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Peckhamia 76.1 Salticidae of the Antarctic land bridge 1
PECKHAMIA 76.1, 7 October 2009, 1―14 ISSN 1944―8120
Salticidae of the Antarctic land bridge
David Edwin Hill 1
1213 Wild Horse Creek Drive, Simpsonville, South Carolina 29680
Introduction
At the breakup of Gondwanaland (~130―110 Ma), Africa, India, and Madagascar moved away from South
America and Antarctica, with Australia still firmly conjoined to the latter (Figure 1). This movement left
South America connected to Antarctica―Australia by a long isthmus (Ithmus of Scotia) of the southern
Andes, at least from the Late Cretaceous (Campanian) through the Eocene (Yanbin 1998, Lawver et al.
1999, Sanmartín 2002). This Antarctic land bridge remained in place from the late Cretaceous through
the Paleocene (65.5 ± 0.3 Ma to 55.8 ± 0.2 Ma) and Eocene (55.8 ± 0.2 to 33.9 ± 0.1 Ma) epochs. At times
it may have included short island arcs at either the Australian (Tasmanian) or South American ends. This
land bridge, associated with a tropical to temperate Antarctic climate (Francis et al. 2008), was thus
available to support the dispersal of plant and animal species for about 75 to 95 million years after the
separation of Africa, a very long time. One very interesting aspect about this interval is that it also
brackets the mass extinction event at the end of the Cretaceous. According to Penney et al. (2003),
however, that event did nothing to reduce the diversity of spiders as a group.
Figure 1. Cretaceous (top) and Eocene
(bottom) reconstructions of Earth
topography and bathymetry. Although
these reconstructions provide a good
view of the separation of Africa,
Madagascar, and India from the rest of
Gondwanaland, they do not depict the
land connections beween Antarctica
and either South America, or Australia,
respectively, that are thought to have
persisted well into the Eocene (Yanbin
1998, Lawver et al. 1999, Sanmartín
2002, Francis et al. 2008, and others).
© by Ron Blakely, NAU Geology.
Noncommercial use with attribution
permitted.
Peckhamia 76.1 Salticidae of the Antarctic land bridge 2
Near the end of the Eocene, at the Eocene―Oligocene boundary, the Australian plate, including New
Guinea, separated from Antarctica and began its long journey toward the north. This opened up the
Tasmanian Seaway, allowing the cold Antarctic Circumpolar Current to isolate Antarctica, leading to the
formation of a permanent ice sheet over that continent by ~33.5 Ma (Exon et al. 2000, 2004, Pollard and
DeConto 2005). Whether this opening, and the subsequent opening of the Drake Passage between South
America and Antarctica (Bohoyo et al. 2007, Maldonado et al. 2007, Miller 2007, Smalley et al. 2007,
Eagles et al. 2009), can fully account for the rapid cooling of Antarctica at the end of the Eocene is still an
open question, and the decline in atmospheric CO2 at that time may have been more important (DeConto
and Pollard 2003, Huber et al. 2004, Barker and Thomas 2004, Livermore et al. 2004, Barker et al. 2006).
Ocean floor presently separating South America and Tasmania, respectively, from Antarctica was
deposited after this time, beginning at the Eocene―Oligocene boundary (Torsvik et al., 2008). In any case,
rapid cooling did follow the end of the Antarctic land bridge between Australia and South America, and
contributed to the subsequent isolation of the two great continental faunas. The Eocene was followed by
more extensive cooling and the growth of ice sheets in the Oligocene (Miller et al. 2008). For reference,
Lawver et al. (1999) provide a useful animated reconstruction of the breakup of Gondwanaland, and both
Brown et al. (2006) and Torsvik et al. (2008) have published plate tectonic reconstructions (maps) of the
entire Cenozoic transition around Antarctica. The age of the oceanic lithosphere (Figure 2) provides a
concise graphic overview of the timing of separation of the continents since the break-up of Pangaea
(~175 Ma).
Figure 2. Age of the ocean lithosphere (Ma). Image created by Elliot Lim, Cooperative Institute for Research in Environmental
Sciences, NOAA National Geophysical Data Center (NGDC) Marine Geology and Geophysics Division. Data and images available
from http://www.ngdc.noaa.gov/mgg/. Data Source Müller et al. (2008).
A south-polar view of this chart (Figure 3) also depicts the relatively recent (since the Eocene ~33 Ma)
separation of Antarctica from Australia (upper right, Tasmania) and South America (lower left, Andes to
Transantarctic Range). Also note the extensive sea-floor spreading (green areas) to the left, between
Antarctica and Africa (upper left), and between South America and Africa, associated with the early
break-up of Gondwana (~130—110 Ma).
million years (Ma)
280
0 20 40 60 80 100 120 140 160 180 200 220 240 260
Peckhamia 76.1 Salticidae of the Antarctic land bridge 3
Although much of present-day Antarctica. particularly in the West (Western Hemisphere), is below sea-
level (Figure 4), models of the Eocene―Oligocene transition that have been corrected for thermal
contraction resulting from tectonic extension and for erosion and sedimentation since 34 Ma indicate that
most of Western Antarctica was actually above sea level at that time (Wilson and Luyendyk 2009). Even
after more extensive glaciation in the Miocene (Jamieson and Sugden 2008), a tundra habitat persisted in
Antarctica as recently as 14.1―13.8 Ma (Lewis et al. 2008).
280 20 40
0 60 80 100 120 140 160 180 200 220 240 260
million years (Ma)
Figure 3. Age of the ocean lithosphere, from a south
polar view with Antarctica at the center. Australia is to
the upper right, South America to the lower left, and
Africa to the upper left. Sea-floor spreading in green is
primarily associated with the break-up of Gondwana.
Note the presence of an earlier rift to the south of
Australia, before later sea-floor spreading separated
Tasmania from Antarctica. Sea-floor spreading between
the southern Andes of South America and the
Transantarctic Range also took place primarily in the
post-Eocene timeframe. Images by R. D. Müller and P. W.
Sloss, NOAA-NESDIS-NGDC. Data and images available
from http://www.ngdc.noaa.gov / mgg/ . Data Source
Müller et al. (2008).
Figure 4. Subglacial topography and bathymetry of
Antarctica. Although much of Western (Western
Hemphere, to the left) Antarctica is now below sea level,
corrected models now indicate that most of this area was
above sea level in the Eocene (Wilson and Luyendyk
2009). At upper left, the long Transantarctic Mountain
Range approaches the southern Andes of South America.
© by Paul V. Heinrich. Use subject to Creative Commons
Attribution 3.0 Unported License.
Peckhamia 76.1 Salticidae of the Antarctic land bridge 4
The Antarctic climate during this transition is of great interest. Studies of plant fossils indicate that a
tropical to subtropical climate dominated during the late Cretaceous (85 Ma), with a mean summer
temperature of about 20oC (Francis et al. 2008). After some cooling, a generally warm, ice-free period
continued through the Paleocene, with some warming in the early Eocene. By the late Eocene, the climate
was much cooler, and temperate forests were dominated by Southern Beech (Nothofagus, with living
species in Australia, Tasmania, New Guinea, New Caledonia, New Zealand, Chile and Argentina), and
monkey puzzle trees similar to living Araucaria araucama (Cantrill and Poole 2005, Poole and Cantrill
2006, Francis et al. 2008, Jamieson and Sugden 2008). Araucaria species are now found in New
Caledonia, Norfolk Island, Australia, New Guinea, Argentina, Chile, and southern Brazil. Fossil marsupials,
related to those presently found in South America, have recently been reported from Antarctic rocks of
Eocene age (Woodburne and Zinsmeister 1982, Goin et al. 1999, 2007, Beck et al. 2008). There is even
good reason to believe that at least four endemic species of Antarctic springtails (Collembola) represent a
continuous Gondwanan line of descent that diversified in Antarctica during the mid- to late Miocene,
21―11 Ma (Stevens et al. 2006). Based on the requirements of modern salticids, we can safely say that
the Antarctic land bridge could have easily supported a diverse array of salticids during its long existence,
particularly up to the late Eocene.
For ease of reference, I will refer to the joined continents of Australia, Antarctica, and South America
collectively as Australamerica (late Gondwana). To find Australamerican clades that used the Antarctic
land bridge, we can look for groups that meet the following conditions: 1, The clade had to originate in
Australamerica, during the long time period of its existence (~130―34 Ma). 2, Sister groups within the
clade can now be found in both Australia and South America (at least in fossil form). 3, The presence of
members of the clade in other areas, if applicable, can be explained by secondary migration from either
Australia or South America.
The relationship of the Australian to the South American fauna, particularly with respect to the
distribution of marsupial mammals, has been long recognized. Most early explanations for this
relationship, before the current acceptance of continental drift and plate tectonics, were awkward, and
have little or no support today. As early as 1924, however, Launcelot Harris presented a very bold and
determined argument in support of the migration of marsupials directly over an Antarctic land bridge.
Several other groups of animals that, based on criteria 1―3, above, may be characterized as native
Australamericans, are identified in Table 1.
Table 1. Some clades of apparent Australamerican origin that appear to have migrated across the Antarctic land bridge before
their more recent diversification within continental boundaries.
Clade Australian sister group South American sister
group
Gondwanan outgroup
(more ancient clade)
References
Mammalia:
Marsupalia
(part)
all Australian
marsupials
all South American
marsupials
Mammalia: Marsupalia Woodburne and Zinsmeister
1982, Goin et al. 1999, Luo et al.
2003, Nilsson et al. 2004, Goin et
al. 2007, Beck et al. 2008
Aves:
Struthioniformes
(part)
Emu (Dromaius),
Cassowary (Casuarius),
Kiwi (Apteryx)
Tinamiformes:
Crypturellus, Eudromia,
Nothoprocta, Tinamus
Struthioniformes,
including African
Ostrich (Struthio)
van Tuinen et al. 1998, Cooper et
al. 2001, Gibb et al. 2007, Hackett
et al. 2008, Harshman et al. 2008
Testudines:
Chelidae
all Australian chelids all South American
chelids
Pleurodira, including
Podocnemidae and
Pelomedusidae
Gaffney 1977, Fujita et al. 2004,
Krenz et al. 2005
Anura: Hylidae
(part)
all Pelodryadinae all Phyllomedusinae Hylidae, including
Hylinae (Hyla)
Faivovich et al. 2005, Frost et al.
2006, Zeisset and Beebee 2008
Of these groups, the timing of the diversification of the Marsupalia (late Cretaceous to early Cenozoic)
has received the most attention, and is fully in line with the hypothesis of Australamerican origin for the
Peckhamia 76.1 Salticidae of the Antarctic land bridge 5
living species (Nilsson et al. 2004). This does not require that the first marsupial was Australamerican,
however (Luo et al. 2003). Recent work (Hackett et al. 2008, Harshman et al. 2008) places the flying
neotropical Tinamous (Tinamiformes) as a sister group to living flightless birds of Australia, New Guinea,
and New Zealand. South American Rheas (Rhea and Pterocnemia) are more closely related to this group
than the Ostrich (Struthio), and at least one view (Harshman et al. 2008, Fig. 2) supports the possibility of
a closer relationship between the Rheas and the Tinamiformes. The Kiwi (Apteryx) appears to have
arrived in New Zealand later than the extinct Moas, and is not closely related (Cooper et al. 2001). Chelid
fossils have never been found outside of Australamerica.
I know of no salticid spiders, living or fossil, that have ever been found in Antarctica. Yet it is quite
possible that a diverse population of salticids, ancestral to at least some of those that we can find today in
Australia and the Americas, did live in Australamerica, and were of Australamerican origin. Assuming
that Australasian ancestors (or clades based on these ancestors and their descendents) did exist, our
challenge lies in finding the corresponding clades among the known Salticidae.
Prehistory of the Salticidae
As noted by Hill and Richman (2009), the fossil record for the Salticidae is indeed sparse, and is limited to
the Cenozoic. Some of the fossils that have been found are reviewed in Table 2.
Table 2. Some fossil salticid genera. Under each description, similar recent salticids are identified in some cases in
parentheses.
Era/Epoch Locality and Source Description References
Eocene
~54―42 Ma
Baltic Sea: Baltic
Amber
Gorgopsina (~hisponine), Prolinus (~hisponine),
Eolinus (~Cyrba, Portia), Paralinus (~spartaeine?),
Almolinus, Cenattus, Distanilinus
Prószyński and Żabka 1980, Keiser
and Weitschat 2006, Maddison and
Zhang 2006, Dunlop et al. 2009,
Wolfe et al. 2009
Oligocene to
Miocene
~30―20 Ma
Chiapas, Mexico:
Chiapas Amber
Lyssomanes García-Villafuerte and Penney 2003
Miocene
~20―15 Ma
Dominican Republic:
Dominican Amber
Lyssomanes, Nebridia, Thiodina, Corythalia,
Descangeles, Descanso, Pensacolatus
Cutler 1984, Iturralde-Vinet and
MacPhee 1996, Dunlop et al. 2009
Given our necessary reliance on fossil amber, we need to recognize that the absence of a group from that
record does not establish the fact that this group did not exist. It may simply mean that members of this
group did not live on the trunks of trees that produced that amber, or that, for behavioral reasons, that
group was not likely to be captured in amber. For example, Penney (2007) has reported an unusual lack
of any salticid fossils from a deposit of lower Eocene amber in the Paris Basin (France). Although this
finding is consistent with the hypothesis that salticids did not occur in Europe until later in the Eocene, it
does little to establish that hypothesis as credible. In addition, we have not found any definitive
intermediate fossils, or proto-salticids, to clarify the evolution of the family.
Żabka (1995) referred to the major influence of continental isolation in the distribution of major salticid
groups that appeared to become highly diversified in the late Cretaceous to Eocene period. However, we
have no Cretaceous or Paleocene fossils from this group to support any hypotheses related to their
radiation. From the few available records (Table 2), we may assume that a diverse group that included
hisponines and possibly spartaeines (or their close relatives), not greatly different from existing species in
Africa or Asia, could be found in a much warmer northern Europe during the Eocene. For much of this
time the climate there was paratropical (Andreasson and Schmitz 2000, Harrington 2001, Kvaček 2002,
Huber and Caballero 2003). About 20 My later, by the early Miocene, we find an essentially modern fauna
on a Caribbean island. We have no transitional records to explain the emergence of diversity in either
Peckhamia 76.1 Salticidae of the Antarctic land bridge 6
area. With few fossil records, however, hypotheses relative to the Salticidae of Australamerica will have to
rely primarily on the current distribution of salticids, and their known (or supported) phylogeny. For
example, with the current center of hisponine diversity in Africa (Maddison and Needham 2006,
Maddison and Zhang 2006), the presence of Eocene hisponine fossils in Europe, and the lack of
hisponines in Australia and the Americas, it is possible that this group evolved in the Laurasia-African
supercontinent in a post-Gondwana time frame. Given the diversity of spartaeines in Southeast Asia
(Wijesinghe 1990), it is likewise tempting to think that this group also evolved in Laurasia-Africa.
However, as we will discuss below, there is also some evidence for a post-Australamerican origin for this
group, from Australasian ancestors. It is important to note that all living salticids, whether basal or
salticoid (Maddison and Hedin 2006), still represent modern groups. Within the Salticidae, although
some groups have been termed primitive, evolution proceeds in many directions, and there has been
more than one line of descent leading to either an increase in the acuity of the anterior medial eyes, or to
reduction of the posterior medial eyes (Blest 1983, Blest and Sigmund 1984, 1985, Blest et al. 1990, Hill
and Richman 2009).
It has been notoriously difficult to pinpoint the time of emergence of any major group from the fossil
record. Even well-known groups can turn out to be much more ancient than previously assumed (Table
3).
Table 3. Some new fossil discoveries that have pushed back the timeframe for emergence of respective animal groups.
Group Previous discoveries New discovery Reference
Age Formation Age Formation
Mammalia:
Metatheria or near-
marsupials
~75 Ma
(skeletal)
~125 Ma Lower Cretaceous Yixian
Formation, China.
Luo et al. 2003
Sauria: feathered
theropod
~150―145 Ma Jurassic Solnhofen
Limestone in Bavaria
(Archaeopteryx)
~160 Ma Earliest Late Jurassic
Tiaojishan Formation of
western Liaoning, China
Hu et al. 2009
Testudines: Chelidae ~23―5 Ma Miocene (?) and later
fossils from Australia
and South America
~105 Ma Lower Cretaceous (Lower
Albian), Patagonia
Lapparent de Broin
and de la Fuente
2001; see also de la
Fuente 2003
Squamata:
Gekkonidae
~54―42 Ma Eocene Baltic amber 110―97 Ma Lower Cretaceous (Albian)
amber from Myanmar
Bauer et al. 2005,
Arnold and Poinar
2008
Araneae: Araneidae ~45 Ma Middle Eocene oil
shales of the Messel
pit, Hesse, Germany
121―115 Ma Lower Cretaceous (Aptian)
amber from Alava, Spain
Penney 2003, Penney
and Ortuño 2006; see
also Peñalver et al.
2006, 2007
Araneae: Dipluridae ~54―42 Ma Eocene Baltic amber 125―112 Ma Lower Cretaceous (Aptian)
Crato Lagerstätte of Cearà
Province, north-east Brazil
Seldon et al. 2006
Araneae:
Linyphiidae
~54―42 Ma Eocene Baltic amber 135―125 Ma Lower Cretaceous (Upper
Neocomian–basal Lower
Aptian) amber, Kdeirji/
Hammana outcrop,
Lebanon
Penney and Selden
2002
Araneae:
Mecysmaucheniidae
Recent Living species found in
southern South
America and New
Zealand
~100 Ma Lower Cretaceous (Late
Albian) amber of Charente-
Maritime, France
Saupe and Seldon
2009
Araneae: Pisauridae ~54―42 Ma Eocene Baltic amber 107―100 Ma Lower Cretaceous (Albian)
amber from Myanmar
Penney 2004
Peckhamia 76.1 Salticidae of the Antarctic land bridge 7
As noted by Seldon et al. (2009), almost every new specimen of spider from the Palaeozoic and Mesozoic
eras . . . can drastically alter our perception of spider phylogeny. Proposed molecular clocks for
evolutionary sequences are often based on synchronization with the fossil record, and these have to be
reset when discoveries of earlier forms upset the underlying assumptions. The lack of fossil markers for
salticids, in particular, makes this task even more challenging. When we examine the fossil record for
emergence of a clade, we need to be cautious, given the fact that one or several uncommon, early
representatives of that clade may have been evolving in relative isolation, or in a different area for a long
time. Thus available fossils only set an upper bound for emergence, and with new fossil discoveries we
can only expect that this bound will move lower, to an earlier time.
Some hypotheses related to the origin of the Salticidae, and the large salticoid clade, are outlined in Table 4.
Table 4. Some major, alternative hypotheses and associated predictions related to the origin of major salticid clades. All are
consistent with the fossil record. Hypotheses 15 relate to origin of the Salticidae, 610 to the origin of the Salticoida).
Hypothesis Predicted fossils Predicted faunal distribution
1
Salticidae originated before the break-up of
Pangaea into Gondwana and Laurasia ~150
175 Ma
Basal Pangaean
lineages in
Gondwanan and
Laurasian fossils.
Major salticid lineages, except for relict groups, divided
across Gondwana and Laurasia, and subsequently divided
between Africa and Australamerica. Later lineages divided
between Australia and South America.
2
Salticidae originated in Laurasia after the
break-up of Pangaea ~150175 Ma and
before the initial break-up of Gondwana
~130110 Ma
Laurasian fossils
predate Gondwanan
fossils.
Basal lineages found in many non-glaciated Laurasian areas
as relict groups.
3
Salticidae originated in Gondwana after the
break-up of Pangaea ~150175 Ma and
before the initial break-up of Gondwana
~130110 Ma
Gondwanan fossils
predate Laurasian
fossils.
Basal lineages found in many non-glaciated Gonwanan
areas as relict groups.
4
Salticidae originated in Laurasia after the
initial break-up of Gondwana ~130110
Ma, but before the break-up of Australasia
~35 Ma
Laurasian fossils
predate Gondwanan
fossils.
Basal lineages found in many non-glaciated Laurasian areas
as relict groups. Relict groups should also appear in Africa.
No endemic relict groups in Australamerica.
5
Salticidae originated in Australasia after the
initial break-up of Gondwana ~130110
Ma, but before the break-up of Australasia
~35 Ma
Australasian fossils
predate any other
fossils.
Basal, relict groups primarily found in Australasia, in both
South America and Greater Australia. Most lineages in
Laurasia appear in post-Australasian timeframe, after
Australia and New Guinea approach Southeast Asia, or
through Central America-Caribbean archipelago migration.
6
Salticoida originated before the break-up of
Pangaea ~150175 Ma
Diverse, Pangaean
salticoid lineages in
both Gondwanan and
Laurasian fossils.
Major salticoid lineages, except for relict groups, divided
across Gondwana and Laurasia, and subsequently divided
between Africa and Australamerica. Later lineages divided
between Australia and South America.
7
Salticoida originated in Laurasia after the
break-up of Pangaea ~150175 Ma and
before the initial break-up of Gondwana
~130110 Ma
Laurasian fossils
predate Gondwanan
fossils.
Each major Gondwanan salticoid lineage traced back to
more basal Laurasian groups.
8
Salticoida originated in Gondwana after the
break-up of Pangaea ~150175 Ma and
before the initial break-up of Gondwana
~130110 Ma
Gonwanan fossils
predate Laurasia
fossils.
Multiple Gondwanan lineages diverge in Africa and
Australamerica. these lineages diverge later between
Australia and South America.
9
Salticoida originated in Laurasia after the
initial break-up of Gondwana ~130110
Ma, but before the break-up of Australasia
~35 Ma
Laurasian fossils
predate Gondwanan
fossils.
Multiple salticoid lineages can be traced from Laurasian
groups to groups in either Africa or Australasia. More early
salticoid lineages, and earlier fossils, in Africa than in
Australamerica.
10
Salticoida originated in Australasia after the
initial break-up of Gondwana ~130110
Ma, but before the break-up of Australasia
~35 Ma
Australasian fossils
predate any other
fossils.
Multiple salticoid lineages can be traced from Australasian
origins, with primary branches split between South America
and Greater Australia.
Peckhamia 76.1 Salticidae of the Antarctic land bridge 8
Unless many more fossils are discovered, any testing of these hypotheses must be based on the biogeography
and phylogeny of recent species. The great diversity of lineages crossing the Wallace Line (Southeast Asia into
the East Indies to New Guinea) is particularly problematic for this purpose, as these may have originated from
either the north (Laurasia) or from the south (Australamerica). A number of groups that are basal to both the
spartaeines and the salticoids (Tomocyrba, Massagris, hisponines, and perhaps Goleba; Maddison and Needham
2006, Maddison et al. 2008) in the recent Madagascar to East Africa fauna may support either a Pangaean
(hypothesis 1, above) or a Gondwanan (3) origin for the Salticidae. With respect to the origin of the Salticoida,
hypothesis (10) gets some support from the fact that one of the two major branches of the Salticoida, the almost
exclusively neotropical Amycoida (Maddison and Hedin 2003, Maddison et al. 2008), almost certainly has a
South American origin. The apparent failure of Amycoida to cross to Australia may be related to the fact that
these are primarily tropical salticids. The other major branch divided many times, and includes several major
groups that may have crossed the Antarctic land bridge (see below). The failure to find many salticids in early
Eocene Europe (Penney 2007), and no salticoids in later Eocene Europe (Table 2) is consistent with this
hypothesis. The Salticoida may have been largely confined to Australamerica in the Eocene, but we have no
salticid fossils of any kind with which to establish their presence. Again, we need to be very cautious in our
interpretation of a very fragmented, incomplete fossil record. We also must remember that even as diverse a
group as the Salticoida at one time consisted of a single species, a species that may have been neither widely
distributed nor abundant. It is almost certain that any important ancestor species like this would be
missing altogether from the fossil record. Only at a much later date, after it had diversified into a number
of competitive species, would there be any probability of a fossil presence. The relatively short interval
(~13—18 My) between the end of the Eocene and the emergence of modern neotropical genera, as well
the enormous diversity found within the major clades of modern salticoids, suggest that that there were a
number of salticoid species alive during the Eocene. These would include the ancestral species for the
major salticoid clades that we see today.
Tentative identification of some trans-Antarctic salticid clades
Some local or relatively endemic salticid groups from Greater Australia (including New Guinea) have been
matched with possible South American sister groups in Table 5.
Table 5. Some endemic or near-endemic salticid genera from greater Australia (including New Guinea) matched with possible
South American sister groups.
Clade Australian sister group South American sister group References
Spartaeinae
+ lapsiines
Spartaeinae: Mintonia, Portia lapsiines: Gallianora, Lapsias, Thrandina Maddison and Needham 2006,
Richardson 2006, Żabka 1994
Astioida +
Marpissoida
Astioida: Adoxotoma, Arasia, Astia,
Damoetas, Helpis, Holoplatys,
Jacksonoides, Ligonipes, Megaloastia,
Mopsolodes, Mopsus, Myrmarachne,
Ocrisiona, Opisthoncus, Rhombonotus,
Sandalodes, Simaetha, Simaethula,
Sondra, Tara, Tauala, Zebraplatys
Marpissoida: Beata, Bellota, Eris,
Hentzia, Itata, Maevia, Metacyrba,
Peckhamia, Psecas, Rhetenor, Rudra,
Sassacus, Tutelina, Zygoballus
Wanless 1988, Hedin and
Maddison 2001, Maddison and
Hedin 2003, Richardson et al.
2006, Maddison et al. 2008
Euophryinae Euophryinae (part): Ascyltus,
Athamas, Bathippus, Canama, Cytaea,
Ergane, Euryattus, Hypoblemum,
Jotus, Lauharulla, Lycidas, Maratus,
Margaromma, Prostheclina, Servaea,
Spilargus, Udvardya, Zenodorus
Euophryinae (part): Amphidraus,
Anasaitis, Asaphobelis, Belliena, Chapoda,
Chloridusa, Cobanus, Commoris,
Coryphasia, Corythalia, Ilargus, Maeota,
Mopiopia, Neonella, Ocnotelus, Pensacola,
Semnolius, Sidusa, Siloca, Stoidis, Tariona,
Tylogonus
Maddison and Hedin 2003,
Richardson et al. 2006,
Maddison et al. 2008, Hill 2009
Grayenulla +
Hisukattus
Grayenulla Żabka 1992: seven
species from Australia
Hisukattus Galiano 1987: four species
from Argentina, Brazil, and Paraguay
Galiano 1987, Żabka 1992,
Żabka 2002, Żabka and Gray
2002, Richardson et al. 2006
Peckhamia 76.1 Salticidae of the Antarctic land bridge 9
To identify clades that may have crossed the Antarctic land bridge, I began with an examination of the
Australian genera that do not appear to have migrated to that continent at a later time from southeast
Asia. Many of these genera are also endemic to New Zealand and can be grouped into two larger clades,
the Astioida (Maddison et al. 2008) and the Euophryinae (Prószyński 1976). Note that the clades divided
into Australian and South American groups here range in nominal size from a small group of genera to
major divisions of the Salticidae as a whole. Given the long interval over which the Australian land bridge
was in place, and our present uncertainty with respect to a timeline for the evolution of salticid groups,
this can be expected. Although modern genera are given as examples of the respective clades, it can be
expected that unknown, now extinct members of these clades actually participated in any actual
migration across Antarctica.
The modern distribution of spartaeines, with only a few known species from Australia and a center of
diversity in the East Indies, provides little support for the division of spartaeines and lapsiines across
Australamerica. One pre-spartaeine species may have migrated out of Greater Australia, however, and
subsequently diversified in the tropical West Indies and Southeast Asia. This division is included for
consideration because of recent evidence from gene sequencing (molecular phylogeny) that spartaeines
and lapsiines are sister groups (Maddison and Needham 2006).
The second division, between the large groups Astioida (Maddison et al. 2008) and Marpissoida
(Maddison and Hedin 2003), reflects the relatively close relationship between these groups that has been
suggested through comparative gene sequencing (Maddison et al. 2008). Both groups are now greatly
diversified, the former in the greater Australian area (including New Zealand) and the latter in both North
and South America. Both groups include a variety of convergent forms that range from the largest of
salticids (e.g., Mopsus and Phidippus), to flattened, cryptic forms (e.g., Holoplatys and Platycryptus), to ant
mimics (e.g., Myrmarachne and Peckhamia). Given the size, diversity, and regional importance of these
groups, the hypothesis that they diversified to current forms after the closing of the Antarctic land bridge
(after the Eocene) appears to be most consistent with the fact that no astioids are found in South America,
and marpissoids (while having a subsequent, smaller dispersal to the Palaearctic) are essentially
American.
The third division (suggested by Hill 2009) of the Euophryinae is based on the fact that this group has two
current centers of diversity, one in the Americas, and one that appears to radiate out of Australia,
including many endemic species in that area. Comparative gene sequencing (Maddison and Hedin 2003,
Maddison et al. 2008) has indicated a close relationship between euophryines in both areas, but the
detailed phylogeny of existing genera will require more study to determine if a single, or if multiple
divisions of the Euophryinae, can be associated with the closing of the Antarctic land bridge. Timing of
diversification in this group is also of great interest. As noted above, Australamerica was around for a
long time.
In addition to species with an affinity to either Asia or Australia, some very unusual endemic salticids can
be found in the vicinity of New Guinea. These include the basal cocalodines (Maddison 2009), as well as
highly unusual forms like Coccorchestes, Diolenius, and Furculatus (Balogh 1981, Żabka, 1994, Szűts 2003,
Gardzińska and Żabka 2006). Many of the endemics, and almost all of the genera shared with Australia,
can be placed in either the Astioda (e.g., Opisthoncus and Sandalodes) or the Euophryinae (e.g., Bathippus
and Euryattus). More than 200 widely distributed species, most from the tropics of Africa or the East
Indies, have been placed in the antlike genus Myrmarachne. Recently (Maddison et al. 2008) included this
large genus with the related Ligonipes in the Astioida. Almost all of the other Astioda are Australasian in
distribution, and this exception by a widely distributed genus that has also made its way to many tropical
islands (including Madagascar) should not affect our hypothesis with respect to an older Australamerican
origin for the Astioida as a group.
Peckhamia 76.1 Salticidae of the Antarctic land bridge 10
Finally, based on the suggestion (Żabka and Gray 2002) that distinctive Australian endemics of the genus
Grayenulla Żabka 1992 resembled the South American Hisukattus Galiano 1987, I have added this division
to Table 5 for consideration. In both genera the bulb of the male pedipalp is distinctively angulate or
bears unusual protuberances, and and a heavy, curved ebolus emerges laterally (Galiano 1987, Żabka
1992).
Between the astioids, the endemic euophryines, and Grayenulla, this brief review has thus treated most of
the endemic salticids in Australia, and supports the view (Richardson et al. 2006) that a significant
number of endemic species not closely related to the Asian fauna remain to be discovered with further
exploration of the Salticidae of that continent.
It is important to note that, although continental boundaries often appear to determine the distribution of
major groups of salticids (Żabka 1995, Maddison et al. 2008), these spiders are also capable of dispersal
over the ocean (e.g., Żabka and Nentwig 2002, Arnedo and Gillespie 2006). Successful transport of a
single female spider, or a small number of spiders, might result in their colonization and diversification in
a new area, particularly if they were not faced with serious competition. Statistically, the sheer number of
dispersal opportunities across a direct physical connection would appear to drive the larger picture, but
not the complete picture. The introduction of even one species with novel or innovative features, however
improbable, could lead to the radiation of many descendent species over time.
Acknowledgments
I would like to thank Drs. G. B. Edwards, Wayne P. Maddison, Jerzy Prószyński, and David B. Richman for their respective
reviews and meaningful feedback.
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... These circumstances have given rise to intricate scenarios that could explain the hyper diversity of the Neotropical region [71]. Additionally, this diversity could have been boosted by geological events through time, including: (i) continental drift [72,73]; (ii) the intermittent connection through bridges, such as the Antarctic bridge that connected the southern cone of South America with the Australian Region [74,75], and the Berigian bridge with a northern connection to the Palearctic Region [76]; and (iii) the complex geological dynamics in the northern Neotropical such as, for example, formation of the Isthmus of Tehuantepec and Panama [77][78][79]. ...
... Mya (95% HPD: 43-31.6 Mya), an age that matches with the end of the Antarctic bridge connection of the southern hemisphere, in the Late Eocene, at about 35 Mya [74,75]. The origin age of the diversification of the genus has been estimated at 74-60 Mya by Seabolt [23], which is close to the start of continental connections of the southern hemisphere at 65 Mya [74,75]. ...
... Mya), an age that matches with the end of the Antarctic bridge connection of the southern hemisphere, in the Late Eocene, at about 35 Mya [74,75]. The origin age of the diversification of the genus has been estimated at 74-60 Mya by Seabolt [23], which is close to the start of continental connections of the southern hemisphere at 65 Mya [74,75]. Both scenarios do not rule out a Southern Hemisphere fauna connection. ...
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Background: Amblyomma is the third most diversified genus of Ixodidae that is distributed across the Indomalayan, Afrotropical, Australasian (IAA), Nearctic, and Neotropical biogeographic ecoregions, reaching the Neotropic its highest diversity. There have been hints in previously published phylogenetic trees from mitochondrial genome, nuclear rRNA, from combinations of both and morphology that the Australasian Amblyomma or the Australasian Amblyomma plus the Amblyomma species from the southern cone of South America, might be sister group to the Amblyomma of the rest of the world. However, a stable phylogenetic framework of Amblyomma for a better understanding of the biogeographic patterns underpinning its diversification is lacking. Methods: We used genomic techniques to sequence complete and nearly complete mitochondrial genomes –ca. 15 kbp– as well as the nuclear ribosomal cluster –ca. 8 kbp– for 17 Amblyomma ticks in order to study the phylogeny and biogeographic pattern of the genus Amblyomma, with particular emphasis on the Neotropical region. The new genomic information generated here together with genomic information available on 43 ticks (22 other Amblyomma species and 21 other hard ticks–as outgroup–) were used to perform probabilistic methods of phylogenetic and biogeographic inferences and time‐tree estimation using biogeographic dates. Results: In the present paper, we present the strongest evidence yet that Australasian Amblyomma may indeed be the sister group to the Amblyomma of the rest of the world (species that occur mainly in the Neotropical and Afrotropical zoogeographic regions). Our results showed that all Amblyomma subgenera (Cernyomma, Anastosiella, Xiphiastor, Adenopleura, Aponomma, and Dermiomma) are not monophyletic, except for Walkeriana and Ambly- omma. Likewise, our best biogeographic scenario supports the origin of Amblyomma and its posterior diversification in the southern hemisphere at 47.8 and 36.8 Mya, respectively. This diversification could be associated with the end of the connection of Australasia and Neotropical ecoregions by the Antarctic land bridge. Also, the biogeographic analyses let us see the colonization patterns of some neotropical Amblyomma species to the Nearctic. Conclusions: We found strong evidence that the main theater of diversification of Amblyomma was the southern hemisphere, potentially driven by the Antarctic Bridge’s intermittent connection in the late Eocene. In addition, the subgeneric classification of Amblyomma lacks evolutionary support. Future studies using denser taxonomic sampling may lead to new findings on the phylogenetic relationships and biogeographic history of Amblyomma genus.
... America with the Australian Region [75][76] and Berigian bridge with a northern connection to with Palearctic Region [77], iii. and the complex geological dynamics in the northern Neotropical, e.g., formation of the Isthmus of Tehuantepec and Panama [78-80]. ...
... According LnL and AICc values, the DEC + J model was the best selected model (Table S5) and therefore, we focus on reporting the results under this model. Our best biogeographic scenario supports the origin of the diversi cation of Amblyomma in the southern hemisphere at the end of the Eocene (Fig. 3), potentially associated with the faunistic ow in the nal Antarctic Bridge connection [75][76]. This agrees with the South America and Australia lineages (here well represented) as the early divergent event in Amblyomma (Fig. 1) reached with high statistical support. ...
... Further work is needed to evaluate the time divergences of Amblyomma using solid topologies, perhaps including nuclear information and calibrations of endemic species of Islands (e.g., Galapagos, Hispaniola).The origin of diversi cation of Amblyomma was estimated to have occurred 36.8 Mya (95% HPD, 43-31,6), an age that match with the end of the Antarctic bridge connection of the southern hemisphere, in the Late Eocene, at about 35 Mya[75][76]. The origin age of the diversi cation of the genus has been estimated at 74-60 Mya by Seabolt,[23], close ages to the starting connection of the southern hemisphere at 65 Mya[75][76]. ...
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Background Amblyomma is the second most diversified genus of Ixodidae that is distributed across the Indomalayan, Afrotropical, Australasian (IAA), Nearctic, and Neotropical biogeographic ecoregions, reaching in the Neotropic its higher diversity. There have been hints in previously published phylogenetic trees from mitochondrial (mt) genome, nuclear rRNA, from combinations of both and morphology that the Australasian Amblyomma or the Australasian Amblyomma plus the Amblyomma species from the southern cone of South America, might be the sister-group to the Amblyomma of the rest of the world. However, a stable phylogenetic framework of Amblyommafor a better understanding of the biogeographic patterns underpinning its diversification is lacking. Methods We used genomic techniques to sequence complete and nearly complete mt genomes –ca. 15 kbp– as well as the ribosomal operons –ca. 8 kbp– for 17 Amblyomma ticks in order to study the phylogeny and biogeographic pattern of the genus Amblyomma, with particular emphasis on the Neotropical region. The new genomic information generated here together with genomic information available of 43 ticks (22 other Amblyommaspecies and 21 other hard ticks –as outgroup–) were used to perform probabilistic methods of phylogenetic and biogeographic inferences and time-tree estimation using biogeographic dates. Results In the present paper, we present the strongest evidence yet that Australasian Amblyomma may indeed be the sister group to the Amblyomma of the rest of the world (species that occur mainly in the Neotropical and Afrotropical zoogeographic regions). Our results showed that all Amblyomma subgenera included, but Walkeriana and Amblyomma, are not monophyletic, as in the cases of Cernyomma, Anastosiella, Xiphiastor, Adenopleura, Aponomma, and Dermiomma. Likewise, our best biogeographic scenario supports the origin of Amblyomma and its posterior diversification in the southern hemisphere at 47.8 and 36.8 Mya, respectively. This diversification could be associated with the end of the connection of Australasia and Neotropical ecoregions by the Antarctic land bridge. Also, the biogeographic analyses let us see the colonization patterns of some neotropical Amblyomma species to the Nearctic. Conclusions We found strong evidence that the main theatre of diversification of Amblyomma was the southern hemisphere, potentially driven by the Antarctic Bridge's intermittent connection in the late Eocene. In addition, the subgeneric classification of Amblyomma lacks evolutionary support. Future studies using denser taxonomic sampling may take us to new findings on the phylogenetic relationships and biogeographic history of Amblyommagenus.
... The subfamily Asterophryinae is the most speciose group within Microhylidae, currently consisting of 327 species inhabiting the tropical forests of northern Australia, New Guinea, and adjacent Australasian islands westwards to Sulawesi, southern Philippines, and crossing the Wallace line in Bali (Frost, 2018). The original biogeographic hypothesis for this subfamily suggested that the common ancestor of Asterophryinae dispersed to Australia via an Antarctic land bridge (Hill, 2009;Savage, 1973), where it diversified and subsequently dispersed to New Guinea and adjacent Australasian islands. However, based on multilocus phylogenetic analyses, Kurabayashi et al. (2011) demonstrated that the enigmatic genus Gastrophrynoides from Sundaland (Borneo and Malay Peninsula) belongs to the subfamily Asterophryinae as a sister-lineage with respect to all Australasian taxa, suggesting that the basal split of the subfamily may not have occurred in Gondwana, but instead on the Eurasian mainland. ...
Article
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We report on the discovery of a new genus of microhylid subfamily Asterophryinae from northern and eastern Indochina, containing three new species. Vietnamophryne Gen. nov. are secretive miniaturized frogs (SVL<21 mm) with a mostly semi-fossorial lifestyle. To assess phylogenetic relationships, we studied 12S rRNA-16S rRNA mtDNA fragments with a final alignment of 2?591 bp for 53 microhylid species. External morphology characters and osteological characteristics analyzed using micro-CT scanning were used for describing the new genus. Results of phylogenetic analyses assigned the new genus into the mainly Australasian subfamily Asterophryinae as a sister taxon to the genus Siamophryne from southern Indochina. The three specimens collected from Gia Lai Province in central Vietnam, Cao Bang Province in northern Vietnam, and Chiang Rai Province in northern Thailand proved to be separate species, different both in morphology and genetics (genetic divergence 3.1%≤P≤5.1%). Our work provides further evidence for the "out of Indo-Eurasia" scenario for Asterophryinae, indicating that the initial cladogenesis and differentiation of this group of frogs occurred in the Indochina Peninsula. To date, each of the three new species of Vietnamophryne Gen. nov. is known only from a single specimen; thus, their distribution, life history, and conservation status require further study.
... Most previous works, though varying on taxon sampling and molecular data, suggested that Microhylidae are of Gondwanan origin and gave evidence supporting the "Antarctic route scenario" for the Australasian subfamily Asterophryinae, as suggested for several other vertebrate taxa that are distributed in Australia (Van Bocxlaer et al., 2006;Van der Meijden et al., 2007). According to this scenario, the basal split of Microhylidae took place in Gondwana and the ancestor of Asterophryinae dispersed to Australia via Antarctic land bridge (Hill, 2009), where the subfamily diversified (it comprises 323 recognized species to date, Frost, 2017) and subsequently dispersed to New Guinea and adjacent Australasian islands, but was unable to cross the Wallace line with exception of the genus Oreophryne Boettger, which is also known from the island of Bali (west from the Wallace line, see Fig. 1). ...
Article
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We report on a discovery of Siamophryne troglodytesGen. et sp. nov., a new troglophilous genus and species of microhylid frog from a limestone cave in the tropical forests of western Thailand. To assess its phylogenetic relationships we studied the 12S rRNA–16S rRNA mtDNA fragment with final alignment comprising up to 2,591 bp for 56 microhylid species. Morphological characterization of the new genus is based on examination of external morphology and analysis of osteological characteristics using microCT-scanning. Phylogenetic analyses place the new genus into the mainly Australasian subfamily Asterophryinae as a sister taxon to the genus Gastrophrynoides , the only member of the subfamily known from Sundaland. The new genus markedly differs from all other Asterophryinae members by a number of diagnostic morphological characters and demonstrates significant mtDNA sequence divergence. We provide a preliminary description of a tadpole of the new genus. Thus, it represents the only asterophryine taxon with documented free-living larval stage and troglophilous life style. Our work demonstrates that S. troglodytesGen. et sp. nov. represents an old lineage of the initial radiation of Asterophryinae which took place in the mainland Southeast Asia. Our results strongly support the “out of Indo-Eurasia” biogeographic scenario for this group of frogs. To date, the new frog is only known from a single limestone cave system in Sai Yok District of Kanchanaburi Province of Thailand; its habitat is affected by illegal bat guano mining and other human activities. As such, S. troglodytesGen. et sp. nov. is likely to be at high risk of habitat loss. Considering high ecological specialization and a small known range of the new taxon, we propose a IUCN Red List status of endangered for it.
... (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.) than those with Africa (Hill, 2009), a greater similarity exists between the floras of the aforementioned continents (Axelrod and Raven, 1978). Numerous components of the Gondwanan flora have vanished from Africa, but still survive in the modern floras of Australia, New Zealand and Patagonia (Kooyman et al., 2014;Macphail, 1999;Macphail et al., 1993Macphail et al., , 1995. ...
Article
A multi-proxy study of an offshore core in Saldanha Bay (South Africa) provides new insights into fluvial deposition, ecosystems, phytogeography and sea-level history during the late Paleogene-early Neogene. Offshore seismic data reveal bedrock topography, and provide evidence of relative sea levels as low as -100 m during the Oligocene. 3D landscape reconstruction reveals hills, plains and an anastomosing river system. A Chattian or early Miocene age for the sediments is inferred from dinoflagellate taxa Distatodinium craterum, Chiropteridium lobospinosum, Homotryblium plectilum and Impagidinium paradoxum. The subtropical forest revealed by palynology includes lianas and vines, evergreen trees, palms and ferns, implying higher water availability than today, probably reduced seasonal drought and stronger summer rainfall. From topography, sedimentology and palynology we reconstruct Podocarpaceaedominated forests, Proto-Fynbos, and swamp/riparian forests with palms and other angiosperms. Rhizophoraceae present the first South African evidence of Palaeogene/Neogene mangroves. Subtropical woodland-thicket with Combretaceae and Brachystegia (Peregrinipollis nigericus) probably developed on coastal plains. Some of the last remaining Gondwana elements on the sub-continent, e.g., Araucariaceae, are recorded. Charred particles signal fires prior to the onset of summer dry climate at the Cape. Marine and terrestrial palynomorphs, together with organic and inorganic geochemical proxy data, suggest a gradual glacio-eustatic transgression. The data shed light on Southern Hemisphere biogeography and regional climatic conditions at the Palaeogene-Neogene transition. The proliferation of the vegetation is partly ascribed to changes in South Atlantic oceanographic circulation, linked to the closure of the Central American Seaway and the onset of the Benguela Current ~14 Ma.
... The initial Drake Passage would have been an extremely narrow, deep seaway at c. 30 Ma (Eagles and Jokat, 2014). It was also likely that a chain of islands persisted between South America and Antarctica during the opening of the Drake Passage and this may have facilitated biotic dispersal from Antarctica well into the Oligocene (Hill, 2009;Eagles and Jokat, 2014) (Fig. 2). So, the Antarctic land bridge may have at first existed as a terrestrial connection during the Jurassic and Cretaceous, but evolved into islands separated by narrow, shallow water bodies (first lakes, later seas) through the Eocene and perhaps until the Oligocene. ...
Article
Amphi-Pacific disjunct distributions between South America and Australasia are correlated with the breakup and changing palaeo-climate of Gondwana. For a long period, with a temperate climate, Antarctica formed a land bridge between Australia and South America, allowing species to disperse/vicariate between both continents. Dated phylogenies in the literature, showing sister-clades with a distribution disjunction between South America and Australia, were used for the correlation. The initiation of the Antarctic Circumpolar Current, and a change to a colder Antarctic climate is associated with the opening of the Drake Passage between South America and Antarctica at c. 30 Ma, and the final separation of Australia and Antarctica along the South Tasman Rise at c. 45 Ma. The distribution data highlighted the existence of a “southern disjunct distribution” pattern, which may be the result of continental vicariance/dispersal. This is strongly indicative of a connection between Antarctica, South America and Australia; which later provided a dispersal pathway and facilitated vicariance after break up. The taxa that likely dispersed/vicariated via Antarctica included all species with a more (sub)tropical climate preference. Twelve distributions, younger than 30 Ma, are interpreted as the result of long distance dispersal between South America and Australia; these taxa are suited to a temperate climate. The climatic signal shown by all taxa is possibly a consequence of the Australian plate's asynchronous rifting over tens of millions of years in combination with climate changes. These events may have provided opportunities for tropical and sub-tropical species to disperse and speciate earlier than what we observe for the more temperate taxa.
Article
Improved understanding of tick phylogeny has allowed testing of some biogeographical patterns. On the basis of both literature data and a meta-analysis of available sequence data, there is strong support for a Gondwanan origin of Ixodidae, and probably Ixodida. A particularly strong pattern is observed for the genus Amblyomma, which appears to have originated in Antarctica/southern South America, with subsequent dispersal to Australia. The endemic Australian lineages of Ixodidae (no other continent has such a pattern) appear to result from separate dispersal events, probably from Antarctica. Minimum ages for a number of divergences are determined as part of an updated temporal framework for tick evolution. Alternative hypotheses for tick evolution, such as a very old Pangean group, a Northern hemisphere origin, or an Australian origin, fit less well with observed phylogeographic patterns.
Chapter
The arachnofauna of various parts of the Earth is analyzed and the particularities, endemics, relicts, and the presumed ways of formation of the fauna are outlined. Also the northern limits of the groups in the Holarctic are indicated, and the connections in the geological time are analyzed.
Chapter
As factors of distribution of Arachnida are outlined paleogeography and paleodistribution, age of groups, barriers, bridges, ability to overcome them, phoresy, dispersal, climate, orography and many other fundamental concepts.
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The result of a three-year devoted work based on a couple decades of Antarctic research, this 368-page edition is an encyclopedic narrative of the principal topics related to Antarctica – nature, history, sovereignty and politics, Antarctic science, resources, fisheries, tourism and Antarctic names, naturally not forgetting Bulgarian participation. The book includes an extensive bibliography (with most of the items available online), and is amply illustrated with over one hundred photographs, old and new maps and paintings, some of them unique. Lyubomir Ivanov is a polar explorer, founding chair of the Bulgarian Antarctic Place-names Commission, and national representative of Bulgaria to the international Standing Committee on Antarctic Geographic Information (SCAGI). Nusha Ivanova has participated in four Antarctic expeditions, and was the first Bulgarian school student to visit Antarctica. A second, revised and expanded (electronic) edition of the book was published on 26 September 2014, ISBN 978-619-90008-2-3
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