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The origins of the enigmatic Falkland Islands wolf

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The origins of the extinct Falkland Islands wolf (FIW), Dusicyon australis, have remained a mystery since it was first recorded by Europeans in the seventeenth century. It is the only terrestrial mammal on the Falkland Islands (also known as the Malvinas Islands), which lie ~460 km from Argentina, leading to suggestions of either human-mediated transport or overwater dispersal. Previous studies used ancient DNA from museum specimens to suggest that the FIW diverged from its closest living relative, the South American maned wolf (Chrysocyon brachyurus) around 7 Ma, and colonized the islands ~330 ka by unknown means. Here we retrieve ancient DNA from subfossils of an extinct mainland relative, Dusicyon avus, and reveal the FIW lineage became isolated only 16 ka (8-31 ka), during the last glacial phase. Submarine terraces, formed on the Argentine coastal shelf by low sea-stands during this period, suggest that the FIW colonized via a narrow, shallow marine strait, potentially while it was frozen over.
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ARTICLE
Received 4 Oct 2012 |Accepted 1 Feb 2013 |Published 5 Mar 2013
The origins of the enigmatic Falkland Islands wolf
Jeremy J. Austin1,2,*, Julien Soubrier1,*, Francisco J. Prevosti3, Luciano Prates4, Valentina Trejo5,
Francisco Mena6& Alan Cooper1
The origins of the extinct Falkland Islands wolf (FIW), Dusicyon australis, have remained a
mystery since it was first recorded by Europeans in the seventeenth century. It is the only
terrestrial mammal on the Falkland Islands (also known as the Malvinas Islands), which lie
B460 km from Argentina, leading to suggestions of either human-mediated transport or
overwater dispersal. Previous studies used ancient DNA from museum specimens to suggest
that the FIW diverged from its closest living relative, the South American maned wolf
(Chrysocyon brachyurus) around 7 Ma, and colonized the islands B330 ka by unknown means.
Here we retrieve ancient DNA from subfossils of an extinct mainland relative, Dusicyon avus,
and reveal the FIW lineage became isolated only 16ka (8–31 ka), during the last glacial phase.
Submarine terraces, formed on the Argentine coastal shelf by low sea-stands during this
period, suggest that the FIW colonized via a narrow, shallow marine strait, potentially while it
was frozen over.
DOI: 10.1038/ncomms2570
1Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, The University of Adelaide, North Terrace, Adelaide, South Australia 5005.
Australia. 2Department of Sciences, Museum Victoria, Carlton Gardens, Melbourne, Victoria 3001, Australia. 3Divisio
´n of Mastozoologı
´a, Museo Argentino
de Ciencias Naturales ‘Bernardino Rivadavia’—CONICET, Avenida Angel Gallardo 470, Buenos Aires C1405DJR, Argentina. 4Divisio
´n of Arqueologı
´
a, Museo
de La Plata, La Plata 1900, Argentina. 5Martı
´
n Alonso Pinzo
´n 6511, Las Condes, Santiago, Chile. 6Centro de Investigacio
´n en Ecosistemas de la Patagonia
(R10C1003), I. Serrano 509, Coyhaique, Chile. * These authors contributed equally to this work. Correspondence and requests for materials should be
addressed to A.C. (email: alan.cooper@adelaide.edu.au).
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During the Pleistocene (ca. 2.6 Ma–12 ka) South America
supported a diverse canid fauna including large hyper-
carnivorous species (that is, Theriodictis spp., Protocyon
spp.,1–4), as well as smaller species such as the Falkland Islands
wolf (FIW) and a related fox found on the mainland, Dusicyon
avus (Fig. 1). Most of these species became extinct during the
Pleistocene, with D.avus extinct by the late Holocene, and the
FIW extinct in the nineteenth century following human
hunting4–7. The origin of the FIW has been a natural history
mystery for over 320 years8. Following his encounter with the
species in 1,834, Darwin commented ‘As far as I am aware, there
is no other instance in any part of the world, of so small a mass of
broken land, distant from a continent, possessing so large a
quadruped peculiar to itself’9. The mystery is deepened by the
absence of any other terrestrial mammals on the islands. While
the flora and fauna of the islands show overwhelming Patagonian
biogeographical affinities10, the physical isolation of the islands
(B460 km from the South American mainland) has resulted in a
number of theories to explain the origin of the FIW. These
include semi-domestication and transport of a continental
ancestor by humans sometime after their arrival in southern
South America B13–14.5 ka (refs 11,12) or natural dispersal via
rafting on ice or over a land bridge during Pleistocene glacial sea
level minima1,13–15. Recently, ancient DNA analysis revealed the
FIW to be a unique South American endemic only distantly
related to the living South American maned wolf (Chrysocyon
brachyurus)7. However, uncertainty about the canid phylogenetic
tree and ambiguous and/or imprecise fossil calibrations limited
the power of the analysis to an approximate divergence date for
the FIW and maned wolf of 6.7 Ma (4.2–8.9 Ma). The molecular
evolutionary rate estimated in this analysis was also used to
calculate the time to the most recent common ancestor (TMRCA)
of five museum specimens of the FIW as 330 ka (95% highest
posterior density (HPD) 70–640 ka), and this date was used as a
proxy for the colonization age of the Falkland Islands. However,
even if this date estimate was approximately accurate, the
colonization event could have been considerably older or
younger than this depending on the demographic history, any
subsequent lineage extinctions16, and the relationship between
the FIW and unsampled mainland relatives17. Furthermore, the
use of external fossil calibrations to date recent events is known to
be problematic due to the temporal dependency of molecular
rates18,19. Either way, while the estimated 330 ka date and wide
error margins (70–640 ka) clearly predates human arrival, it
provided little evidence as to how the FIW might have colonized
the islands.
Critically, the genetic analysis of the FIW did not include the
putative mainland close relative D.avus, whose phylogenetic
relationship has not been tested. D. avus had a clear archae-
ological association and temporal overlap with modern humans
until extinction B3 ka (ref. 6), raising the possibility of human
transport to the islands. To investigate this issue and characterize
the genetic diversity and phylogenetic relationships of this extinct
canid, we extracted and sequenced 1,069 bp of ancient
mitochondrial DNA from six specimens of D. avus collected
across Argentina and Chile, ranging in age from 7,800–3,000 yr
BP. We also compared the genetic signals with morphological
data from nearly all the extinct and living species of South
American canids, a wide sample of other Caninae, and fossils of
the extinct subfamilies Hesperocyonidae and Borophaginae3.
Results
Sequences. A combined 1,069 bp of mtDNA COII and cyto-
chrome b sequence was obtained from six out of seven D. avus
specimens (Supplementary Table S1). No samples produced PCR
products for the four nuclear gene targets. All PCR and sequen-
cing results were replicated at least once and produced identical
results, suggesting a minimal contribution from damage-related
artefacts.
Topology. Both ML and Bayesian molecular analyses confirm
that D. avus is the closest relative of the FIW with strong sta-
tistical support (Fig. 2), and only six fixed transition differences
separate the FIW and D. avus (Supplementary Fig. S1, uncor-
rected sequence divergence ¼0.56%). Ten additional variable sites
defined three haplotypes within the FIW (from five individuals)
and four haplotypes in D. avus (from six individuals)
(Supplementary Fig. S1). Strong support is also recovered for the
monophyly of the Dusicyon clade, its sister taxon relationship to
the maned wolf, with the bush dog as the outgroup. While
D. avus was weakly recovered as paraphyletic in the topological
analysis, D. avus and the FIW were strongly supported as reci-
procally monophyletic in analyses using a molecular clock and
either external or internal calibrations (Fig. 3 and Supplementary
Fig. S2). The combined morphological and molecular analysis
recovered one most parsimonious tree of 3782.017 steps
(Supplementary Figs S3 and S4), similar to the one previously
reported by Prevosti3, but D.avus and FIW are a monophyletic
group sister of C. brachyurus (Supplementary Figs S2, S3 and S4).
Dating. Date randomization tests confirmed that sufficient signal
existed in the dated concatenated sequences from the six D. avus
individuals to provide appropriate internal temporal calibration
Falkland Islands wolf
(Dusicyon australis)
Dusicyon avus
Chile
Argentina
Falkland
Islands
12
3
4
5
Figure 1 | Specimen distribution. Distribution map of the specimens
analysed. The geographical locations of Dusicyon avus specimens known
from late Pleistocene to late Holocene sites are shown as orange dots, with
the localities of sampled specimens as orange stars. The latter are (1): La
Marcelina 1, Rio Negro, Argentina; (2): Loma de los Muertos, Rio Negro,
Argentina; (3): Ban
˜o Nuevo-1 Cave, Chile; (4): Perro 1 site, Tierra del Fuego,
Argentina. Five specimens of Falkland Islands Wolf (FIW) (red) from
museum collections were previously analysed, (5): Falkland or Malvinas
Islands.
ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2570
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information at the tip of the tree to facilitate molecular dating19
(Supplementary Fig. S5). Bayesian analyses estimate that the FIW
diverged from D. avus recently, around 16.3 ka (95% HPD 7.9–
31.1 ka). The TMRCA for the D. avus, and for the FIW,
specimens were estimated at 15.0 ka (95% HPD 8.5–25.2 ka) and
4.5 ka (95% HPD 0.6–10.6 ka), respectively (Fig. 3). In contrast,
analyses using several deep, and poorly constrained canid fossil
calibration points originally used in Slater et al.7produced much
older age estimates, close to those previously reported
(Supplementary Figs S2, Supplementary Table S2 and
Supplementary Methods).
Interestingly, while the low intra-specific mtDNA diversity
within the FIW (p¼0.0021) was consistent with that of other
insular species, the diversity within D. avus was similar
(p¼0.0026), despite the much larger geographic and ecological
range of the samples. Furthermore, no phylogeographic patterns
were evident within the D. avus sequences (Supplementary Fig.
S1) despite the wide sampling area.
Discussion
The estimated latest Pleistocene date of 16 ka (8–31 ka) for the
origin of the FIW is consistent with events associated with the
0.03
0.83/0.99
1/1
0.9/0.82
1/0.95
1/0.95
0.46/0.76
D. avus SCBN1-2A-3-O369
D. avus PR1-287
D. avus PR1-089
D. avus MAR1
D. avus LM1.E2.2
D. avus SCBN1-2A-1/2-6
D. australis D557
D. australis VT2369
D. australis NHM69.2.24.3
D. australis ANSP588
D. australis NHM37.3.15.4
7
Maned wolf
Crab-eating fox
Hoary fox
Small-eared dog
Culpeo fox
Pampas fox
Bush dog
Sechuran fox
Figure 2 | Phylogenetic position of D. avus and FIW. Phylogenetic tree showing the topological placement of D. avus, FIW and South American canids,
based on the analysis of 1,069bp of mitochondrial cyt b and COII genes. The same topology was recovered using Bayesian (MrBayes, black support values)
and maximum likelihood analyses (PhyML, grey support values). The FIW forms a monophyletic group separate from the mainland D. avus, which is weakly
supported as being paraphyletic, with the maned wolf as the closest living relative to the extinct Dusicyon clade.
D. avus SCBN1-2A-3-O369
D. australis D557
D. australis VT2369
D. australis NHM69.2.24.3
D. avus PR1-287
D. avus PR1-089
D. avus MAR1
D. australis ANSP588
D. australis NHM37.3.15.4
7
D. avus LM1.E2.2
D. avus SCBN1-2A-1/2-6
15.0
(8.5–25.2)
16.3
(7.9–31.1)
4.5
(0.6–10.6)
051015
1
0.41
202530
1
0.9
0.66
1
0.32
0.87
0.96
Ka
Figure 3 | Dated phylogeny of D. avus and FIW (D. australis). Unconstrained phylogeny of D. avus and FIW using a molecular clock, with the radiocarbon-
dated ancient samples as temporally proximate calibration points. The timescale is given in thousand years (ka). The two Dusicyon species are strongly
supported as reciprocally monophyletic clades in this analysis, and when external canid fossil calibration points are used (Supplementary Fig S2). The two
species are estimated to have diverged during the last glacial phase (7.9–31.1 ka), with the upper bound overlapping with the first human presence in
Patagonia11,12. The topology is a maximum clade credibility tree from BEAST with 95% HPD of the calculated node ages represented as grey bars. The use
of ancient samples to provide tip-date calibrations led to considerably younger TMRCA estimates for the Dusicyon clades than previous estimates
(B330 ka for FIW7), which relied on deep canid fossil calibrations of imprecise phylogenetic and temporal position.
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Last Glacial Maximum (LGM) ca. 26–19 ka (ref. 20), and is
considerably younger than the previously estimated date of 330 ka
(ref. 7). The accuracy of molecular divergence date estimates is
directly related to the quality of calibration data, and their
proximity to the date of interest. In this regard, the use of deep
fossil calibration points has been shown to be inappropriate for
intra-specific and recent inter-specific divergences18,19, such as
those within Dusicyon. The radiocarbon dates associated with the
ancient sequences of D. avus provide much closer calibration
points to the late Pleistocene events under consideration than the
early canid fossils used in Slater et al.7, and importantly do not
suffer from the same uncertainty over taxonomic identification
and phylogenetic position. Although additional internal
calibration points would no doubt help refine the inferred
dates, the randomization analyses confirm that the
heterochronous Holocene D. avus sequences contain sufficient
temporal information to calibrate the recent evolutionary history
of Dusicyon (Supplementary Fig. S5 and Supplementary
Methods).
The data do not support a recent origin for the FIW via human
transport from a source population of D. avus on the South
American mainland due to the estimated 16 ka divergence date
and the reciprocal monophyly of the two species in calibrated
analyses. Although the confidence interval (8–31 ka) for the
divergence event overlaps with the earliest human presence in
Patagonia (13–14.5 ka, (refs 11,12), and human agency cannot
therefore be ruled out, the genetic isolation of the FIW
(presumably via transfer to the Falkland Islands) would have
had to occur only once during the earliest phases of human
occupation in Patagonia (that is, 48 ka), which seems
improbable. Furthermore, the estimated 16 ka divergence date is
unlikely to be an overestimate due to the existence of undetected
and phylogenetically closer source populations of D. avus,
because the geographically widespread D. avus samples have
low nucleotide diversity and lack phylogeographic structure.
Indeed, the young TMRCA of B15 ka for D. avus and the lack of
geographic structure and low genetic diversity suggest that the
range of this species has recently expanded across the Patagonian
and Pampean regions, potentially from a LGM refugium.
A pre-human dispersal of the FIW requires either a land bridge
during low sea level stands or a marine crossing, but the absence
of other terrestrial mammals on the Falkland Islands strongly
I
II
III
IV
0 500 km
Figure 4 | Submarine terraces between the mainland and the Falkland Islands. The four submarine terraces (levels I–IV) along the continental shelf (light
blue) between the South American continent and the Falkland Islands: level I¼25/ 30 m isobath; level II ¼85/ 95 m; level III ¼110/ 120 m
and level IV ¼130/ 150 m (ref. 21). Terrace I has been dated ca. 11 ka, and Terrace II ca. 15 ka (ref. 21), with the latter being consistent with the
global Meltwater Pulse 1A, associated with the West Antarctic Ice Sheet20. Terraces III and IV are undated but are thought to reflect earlier phases
of the LGM, with the depth differential matching the rapid 10 m rise from the LGM lowstand caused by the global Meltwater Pulse at 19–20 ka (ref. 20). The
projected sea level position at 140 m is drawn in grey, representing the peak low sea-stand during the LGM (data from Ponce et al.21). At this point
the Falkland Islands would have been separated from the mainland by a shallow marine strait estimated to be as little as 20 km wide and only 10–30 m
deep. It is suggested that this is likely to periodically have been covered by ice, or facilitated ice rafting, during peak glacial conditions.
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argues against a continuous land bridge connection to the
mainland. The particularly shallow slope of the Argentine
continental shelf means that lowered sea levels dramatically
reduce the size of the marine strait separating the Falkland
Islands, which currently has a minimum depth of ca. 160 m
(ref. 21). During the height of the LGM (26–19 ka) global sea
levels were around 130 m lower, which exposed an enormous
coastal plain off the Argentine coast, while the Falkland islands
landmass was about four times larger than the present21–23. Four
pronounced submarine terraces detected on the coastal shelf
(Fig. 4) are thought to record low sea-stands at various points
during the LGM24. The two shallowest have been dated ca. 11 ka
(35/ 40 m isobath) and 15 ka ( 80/ 90 m) (ref. 21), with
the timing of the latter being consistent with Meltwater Pulse 1A,
associated with the onset of the retreat of the West Antarctic Ice
Shelf20. Two deeper terraces situated at the 110/ 120 m and
the 130/150 m isobaths remain undated, but are thought to
represent low sea-stands during earlier phases of the LGM. The
deepest terrace ( 130/150 m) is presumably the oldest and
corresponds to the lowest isostatic conditions, as it has not been
degraded through subsequent near-shore activity. The depth of
the terrace matches the estimated minimal sea level during the
LGM (ca. 130 m), but could also relate to earlier glacial
events24. However, the difference between the 110/120 m and
130/150 m terraces is consistent with a rapid 10 m rise from
LGM lowstand sea levels associated with the 19–20 ka Meltwater
Pulse caused by the widespread retreat of northern hemisphere
ice sheets20. Either way, the molecular date estimate for the
divergence of the FIW of 16 ka (range 8–31 ka) is consistent with
colonization during the LGM, when the submarine terraces
indicate the marine strait was both narrow and shallow.
A low sea-stand between the 130 and 150 m isobaths
would drastically reduce the size of the marine strait separating
the Falkland Islands, potentially to just 20–30 km (Fig. 4), with an
estimated minimum depth of 10–30 m. It is possible that the
ancestors of the FIW were transported across this strait by ice
rafting during this period, as has been suggested, but it seems
more likely that such a shallow marine strait would be
periodically frozen over by continuous sea ice and/or glacial
outflow to produce an ephemeral connection that could act as a
filter, rather than a corridor. Individuals or packs of the ancestor
of FIW pursuing marine food sources (for example, seals,
penguins, seabirds) on the margins of the ice would have an
increased likelihood of colonization compared with other South
American endemic mammals that would not cross large ice-fields
due to either habitat or behaviour. An interesting question is why
previous glacial maxima did not lead to other mammal
colonizations of the Falkland Islands, as the marine strait is
thought to have been shallower during earlier phases of the
Pleistocene before the erosion of soft sediments21. It is possible
that if there were such previous residents they may have gone
extinct without leaving a fossil record.
The greatly improved resolution provided by genetic data from
the closest mainland relative allows us to conclude that the
ancestor of the FIW likely colonized the Falkland Islands via
mobile or static ice, crossing a narrow strait during the Last
Glacial Maximum in Patagonia.
Methods
Samples. Seven D. avus teeth representing different individuals (Supplementary
Table S3) were obtained from four sites in Patagonia: La Marcelina, Rio Negro (one
tooth); Loma de los Muertos, Rio Negro (one tooth) and Perro 1 site; Tierra del
Fuego (two teeth) in Argentina; and Ban
˜o Nuevo-1 Cave (three teeth) in Chile. One
premolar from Ban
˜o Nuevo-1 cave has been dated to 7,860±78 cal. yr BP
(UCIAMS-19490, Supplementary Table S3), while specimens from Perro 1, Loma
de los Muertos and La Marcelina 1 were dated to 3,085±133, 3,072±151, and
3,814±117 cal. yr BP (AA75297, AA83516, AA90951; (refs 6,23) Supplemen tary
Table S3), respectively. A second specimen from Ban
˜o Nuevo-1 cave was undated
but assigned a prior mean age of 7,500 cal. yr BP (6,000–9,000 cal. yr BP 95% range,
Supplementary Table S3) based on an archaeological association with dated
remains from the same site. Teeth were obtained from recently described speci-
mens that have diagnostic morphological features of D.avus for example, large
lower carnassial, a lower fourth premolar with a second distal accessory cusp and a
narrow distal cingulum4,6. The FIW exhibits differences in dental morphology
from D. avus including a more reduced protocone in the P4, smaller metaconid in
the m1, and taller and more acute principal cusps of the premolars6. These
characters are generally associated with a more carnivorous diet25,26 and
potentially reflect the limited dietary breadth of the FIW27,28.
Molecular analyses. To avoid the potential for contamination of D. avus samples
with contemporary canid DNA or previously amplified FIW PCR products7,
all pre-PCR work was performed in a dedicated ancient DNA laboratory
geographically separated (by B1.5 km) from post-PCR and other molecular
biology laboratories at the Australian Centre for Ancient DNA, University of
Adelaide, South Australia. No contemporary canid DNA had ever been present in
the pre-PCR laboratory. The ancient DNA facility includes high-efficiency
particulate air-filtered positive air pressure with one-way air flow, overhead
ultraviolet (UV) lights, individual work-rooms, the use of dead-air glove boxes with
internal UV lights for DNA extractions and PCR set-up, regular decontamination
of all work areas and equipment with sodium hypochlorite, personal protective
equipment including full body suit, face mask, face shield, boots and triple-gloving
and strict one-way movement of personnel (shower4freshly laundered
clothes4ancient DNA laboratory4post-PCR laboratory).
A negative extraction control was included with every set of DNA extractions
and all extractions were carried out in small sets and generally included samples
from phylogenetically divergent non-canid species. DNA was extracted using a
modified silica-based method29 designed to maximize recovery of PCR-amplifiable
DNA from ancient bone and tooth specimens while minimizing coextraction of
PCR inhibitors.
Short fragments of mitochondrial (166–240 bp) and nuclear (108–173 bp) DNA
were targeted by PCR, to assemble a 652-bp fragment of the mtDNA COII gene, a
394-bp fragment of the mtDNA cytochrome b gene, and four nuclear loci (CH21,
VANGL, VTN(SNP), VTN(indel)7, (Supplementary Table S4). One microlitre of
extract, in parallel with extraction controls and negative PCR controls, was
amplified in a 25-ml PCR containing: 1 Platinum Taq High Fidelity Buffer
(Invitrogen), 2 mM MgSO
4
, 0.4 mM each primer, 0.25 mM each dNTP, 0.5 U
Platinum Taq DNA Polymerase High Fidelity, 1 mg ml 1RSA (Sigma-Aldrich)
and sterile H
2
0. PCRs were run on a Palmcycler (Corbett Research) under the
following condit ions: initial denaturation at 94 °C for 1 min; 50 cycles of
denaturation at 94 °C for 15 s; primer annealing at 55 °C for 15 s; elongation at
68 °C for 30 s; a final elongation step at 68 °C for 10 min. PCR products were
visualized under UV light on a 3.5% agarose gel stained with ethidium bromide.
Successful amplifications were purified using Ampure (Agencourt) according to the
manufacturer’s instructions and sequenced directly using Big Dye chemistry and an
ABI 3130XL Genetic Analyzer (Applied Biosystems). All positive PCR and
sequencing results were repeated to ensure reproducibility.
Sequence alignment.COII and cytochrome b sequences were generated from six
D. avus specimens and aligned with the available sequences from five FIW
specimens, and the eight South American canids used by Slater et al.7(maned wolf—
Chrysocyon brachyurus, bush dog—Speothos venaticus, crab-eating fox—Cerdocyon
thous,small-eareddogAtelocynus microtis, sechuran foxLycalopex sechurae,
culpeo fox—Lycalopex culpaeus,pampasfoxLycalopex gymnocercus,hoaryfox
Lycalopex vetulus). The South American canids have previously been shown to be
monophyletic7.
Phylogenetic analyses. To study the phylogenetic placement of D. avus,we
performed both Bayesian (MrBayes30) and maximum likelihood (PhyML31)
analyses on the entire data set (D. avus, FIW and eight South American canids, as
described above). The best substitution model was selected through comparison of
Bayesian information criteria scores using ModelGenerator v0.85 (ref. 32).
In addition, a haplotype network showing genealogical relationships between all
D. avus and FIW sequences was generated using statistical parsimony implemented
in TCS v1.17 (ref. 33). Nucleotide diversity within D. avus and FIW was calculated
using DnaSP v5.10.01 (ref. 34).
We also performed a total evidence analysis combining the new mitochondrial
sequences with previously reported morphological, behavioural and genetic data
for South American canids35. The individual mitochondrial sequences of D.avus
and FIW were each combined into a single consensus sequence using the
programme Bioedit 7.0.5.3 (ref. 36), and the variable sites were scored as
polymorphic using the IUPAC code. The mitochondrial sequences were added to a
previously published ‘total evidence’ matrix3containing dental and skeletal
characters, behavioural and life history traits37, and 22 nuclear and 3 mitochondrial
genes from a wide sampling of living canids and several fossil representatives (for
whom only dental and skeletal data was available). Sequence gaps were coded as a
fifth state. The combined matrix was analysed under Maximum parsimony with
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the software TNT version 1.1 (ref. 38) under equal weighting (SI). Trees were
obtained from heuristic searches with 1,000 random-addition sequence replicates
and tree bisection-reconnection (TBR) branch swapping, supplemented by a TBR
round on the resulting shortest-length trees. Additional searches were conducted
using the sectorial-searches, tree-drifting and tree-fusing algorithms39, but they
found the same trees. Branch support was quantified with symmetrical-resampling
jackknifing frequencies and frequency differences (5,000 resamples; ref. 40). The
trees were rooted with the basal canid Hesperocyon gregarius. Branches were
collapsed following the rule number 1, where any branch with at least one
reconstruction with 0 changes is collapsed (minimal branch length ¼0; see ref. 41).
Calibration using fossils versus internal radiocarbon dates. To examine the
problems caused by the temporal dependency of molecular dates, we performed
molecular dating analyses using either internal (radiocarbon), or external (fossil),
dates. The advantages of using internal calibrations to study recent evolutionary
events are well established18,19, and given a divergence event thought to be late
Pleistocene (natural dispersal) or Holocene (human dispersal), the internal
radiocarbon dates for D. avus appear far more appropriate than external fossil
dates of 4–32 Ma (with largely unknown error margins) and uncertain taxonomic/
phylogenetic position.
Radiocarbon ages were converted to calibrated ages using the CALIB 6.0.1
software available at http://intcal.qub.ac.uk/calib/42,43, using the Southern
Hemisphere SHCal04 curve44 and two sigma ranges. A Bayesian phylogenetic
analysis was performed using the FIW and D. avus sequences to estimate the
TMRCA of each species, and their immediate ancestor, using the D. avus
radiocarbon dates as the sole calibration points. The best substitution model was
selected through comparison of Bayesian information criteria scores using
ModelGenerator v0.85 (ref. 32). Phylogenetic analyses were performed with BEAST
1.6.2 (ref. 45) using a strict molecular clock (analyses using an uncorrelated
lognormal relaxed clock could not reject the strict clock assumption), and the
Bayesian skyride demographic model46 to account for demographic changes
through time. The results were processed with Tracer v1.5 (ref. 47) to check that
each sampled parameter had an effective sample size over 200.
To test whether the signal from the radiocarbon dates associated with the ancient
sequences is sufficient to calibrate the Dusicyon phylogeny, a ‘date randomization
test’48 was conducted. This test consists of randomizing all dates associated with
the sequences (including modern ones), and then replicating the phylogenetic
analysis. Ten replicates of the BEAST phylogeny described in the main text were
performed using different iterations of randomized dates.
To examine the impact of using external canid fossils as calibration points, we
repeated the initial molecular analysis of the FIW by Slater et al.7, but with the
addition of six new sequences from D. avus (Supplementary Fig. S2).
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ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2570
6NATURE COMMUNICATIONS | 4:1552 | DOI: 10.1038/ncomms2570 | www.nature.com/naturecommunications
&2013 Macmillan Publishers Limited. All rights reserved.
46. Minin, V. N., Bloomquist, E. W. & Suchard, M. A. Smooth skyride through a
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Acknowledgements
We thank J. Rabassa and J. F. Ponce for comments and access to high-resolution versions
of published figures, and F. Martin, M. Salemme, F. Santiago, O. Palacios and M. Silveira
for access to the D.avus samples, and the Otago Museum for a sample of their FIW
specimen. This work was funded by the Australian Research Council, PICT 2011-309
(ANPCyT), PIP 1054 and PIP 201101-00164 (CONICET).
Author contributions
A.C. and J.J.A. conceived the study; F.J.P., L.P., V.T. and F.M. provided samples; J.J.A.,
J.S. and F.J.P. performed the experiments; and A.C., J.S. and J.J.A. wrote the manuscript
with input from all authors.
Additional information
Supplementary Information accompanies this paper at http://www.nature.com/
naturecommunications
Competing financial interests: The authors declare no competing financial interests.
Reprints and permission information is available online at http://npg.nature.com/
reprintsandpermissions/
How to cite this article: Austin, J.J. et al. The origins of the enigmatic Falkland Islands
wolf. Nat. Commun. 4:1552 doi: 10.1038/ncomms2570 (2013).
NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2570 ARTICLE
NATURE COMMUNICATIONS | 4:1552 | DOI: 10.1038/ncomms2570 | www.nature.com/naturecommunications 7
&2013 Macmillan Publishers Limited. All rights reserved.
... Although it is unclear, some authors (Austin et al., 2013;Slater et al., 2009) consider this last lineage to be related to the extinct genus Dusicyon mentioned previously. ...
... The only one of the now-extinct South American Pleistocene canids known to have had a distribution in Chile was Dusicyon avus, found in Chilean Patagonia and Tierra del Fuego, on the southern tip of Chile (e.g., Méndez et al., 2014;Prevosti et al., 2009). The genus Dusicyon, to which D. avus and D. australis (also known as the Falklands wolf) belonged, went extinct fairly recently, although the extinction process may have been initiated by reductions in its distribution beginning in the Pleistocene (Austin et al., 2013;Prevosti et al., 2011;Prevosti et al., 2015). In spite of this, there have been reported sightings of D. avus by English explorers as late as 1870. ...
... lone-hunting cursorial strategy in areas with tree cover, at least for certain prey. The maned wolf, Chrysocyon brachyurus, is also large (up to 23 kg) (Sheldon, 1992) and described as fox-like (Dietz, 1985), and in some accounts a sister genus of Dusicyon spp (Austin et al., 2013). ...
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Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe), are reported to be very susceptible to predation by pack-hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies, and hunting behavior. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviors of guanacos: predator inspection of ambush predators, for example, Puma concolor, and rushing at and kicking smaller cursorial predators, for example, Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group-hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
... When the first Europeans arrived in 1690, they found no evidence of people nor have any archaeological sites been discovered that support a pre-European human presence in the islands to date. It has thus been hypothesized that D. australis evolved from populations of D. avus that crossed the South Atlantic during the Last Glacial Maximum, when sea levels were ~125 m lower and the exposed continental shelf reduced the distance of a transoceanic crossing (4). However, this hypothesis is undermined by a paucity of evidence for a sea-ice bridge, as well as the lack of any other native terrestrial mammals in the Falkland Islands (4). ...
... It has thus been hypothesized that D. australis evolved from populations of D. avus that crossed the South Atlantic during the Last Glacial Maximum, when sea levels were ~125 m lower and the exposed continental shelf reduced the distance of a transoceanic crossing (4). However, this hypothesis is undermined by a paucity of evidence for a sea-ice bridge, as well as the lack of any other native terrestrial mammals in the Falkland Islands (4). Alternately, if D. australis diverged from D. avus in mainland South America, perhaps via domestication, then it could have arrived in the Falkland Islands during the Holocene via transport by Yaghan (Yámana) or other Indigenous peoples before European exploration of the South Atlantic (1). ...
... Ethnographic accounts and recent genetic studies both suggest that L. culpaeus underwent self-domestication through the construction of a new ecological niche shared with humans (33,36). While D. australis was not closely related to L. culpaeus, its closest relative, D. avus, is the most commonly identified canid in Fuego-Patagonian archaeological sites until its extinction by ~400 BP (3,4,34). At the Loma de los Muertos mortuary site, a D. avus individual (2792 ± 50 14 C BP) was found to be the primary subject of an interment, which suggests an intimate and mutualistic interspecies relationship (35). ...
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When Darwin visited the Falkland Islands in 1833, he noted the puzzling occurrence of the islands’ sole terrestrial mammal, Dusicyon australis (or “warrah”). The warrah’s origins have been debated, and prehistoric human transport was previously rejected because of a lack of evidence of pre-European human activity in the Falkland Islands. We report several lines of evidence indicating that humans were present in the Falkland Islands centuries before Europeans, including (i) an abrupt increase in fire activity, (ii) deposits of mixed marine vertebrates that predate European exploration by centuries, and (iii) a surface-find projectile point made of local quartzite. Dietary evidence from D. australis remains further supports a potential mutualism with humans. The findings from our study are consistent with the culture of the Yaghan (Yámana) people from Tierra del Fuego. If people reached the Falkland Islands centuries before European colonization, this reopens the possibility of human introduction of the warrah.
... The 13 contemporary South American canids are divided into two lineages (Perini et al. 2010), one of species similar to Old World foxes (genera Lycalopex, Atelocynus, and Cerdocyon ), and the other of species similar to Old World wolves (Chrysocyon and Speothos ). Although it is unclear, some authors (see Austin et al. 2013;Slater et al. 2009) consider this last lineage to be related to the extinct genus Dusicyon mentioned previously. ...
... Méndez et al. 2014;Prevosti et al. 2009). The genus Dusicyon , to which D. avus and D. australis (also known as the Falklands wolf) belonged, went extinct fairly recently, although the extinction process may have been initiated by reductions in its distribution begining in the Pleistocene (Prevosti et al. 2011;Austin et al. 2013;Prevosti et al. 2015). In spite of this, there have been reported sightings of D. avus by English explorers as late as 1870. ...
... Yet, as for L. pictus (Fanshawe & Fitzgibbon 1993), they might be equally successful with a lone-hunting cursorial strategy in areas with tree cover, at least for certain prey. The maned wolf, Chrysocyon brachyurus is also large (up to 23 kg) (Sheldon 1992) and described as fox-like (Dietz 1985), and in some accounts a sister genus of Dusicyon spp (Austin, 2013). While it has never occurred in Chile to date (Torres et al. 2013), it would have coexisted with Lama spp., rheas (Rhea spp. ...
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Artiodactyl prey species of Chile, especially guanacos ( Lama guanicoe ) are reported to be very susceptible to predation by pack hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies and hunting behaviour. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviours of guanacos: predator inspection of ambush predators, e.g. Puma concolor , and rushing at and kicking smaller cursorial predators, e.g. Lycalopex culpaeus . We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
... Questions like how, when and from where different animal taxa arrived in an area have fascinated researchers in different fields for a long time. These questions especially attract attention in areas surrounded by a natural geographical border where there is no obvious answer to how the animals arrived, as is the case with the fauna on Madagascar (Vences 2004;Samonds et al. 2012) or the Falkland Islands (Austin et al. 2013). Apart from human-mediated translocation, the occurrence of animals on remote islands is commonly explained by natural dispersals through rafting on floating debris, swimming, flying or migration over a land bridge which was ultimately submerged (Samonds et al. 2012;Storey et al. 2013). ...
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This thesis aims to study the interactions of pre-agricultural societies in Scandinavia with wild mammals, for example in terms of hunting and translocation. More specifically, the aim is to investigate the possibility of identifying examples of overexploitation, targeted hunting or translocation of wild mammals in prehistoric Scandinavia, and to discuss the implications this could have had for both the wild animals and the humans. The thesis also studies translocation to evaluate the feasibility of using it as a proxy for prehistoric human mobility, and to understand the motivation for this action. Although the focus is on the animals in this thesis, the ultimate purpose is to study humans and their interactions with animals in prehistory. The thesis applies genetic analyses to zooarchaeological material of various mammalian species from different Scandinavian sites, in order to study whether the genetic structures have changed in these species over time, and to assess whether these changes were induced by different human actions. The species studied in this thesis were selected on the basis of the importance they are considered to have had for prehistoric people. The dissertation comprises five studies. The first study investigates the occurrence of mountain hares on the island of Gotland, and discusses how they got there and where they came from. The second study explores the temporal genetic structure of the grey seal in the Baltic Sea, and discusses whether humans and/or climate were the drivers for the sudden disappearance of grey seals from the island of Stora Karlsö. The third study concerns a shift where moose apparently became less important as prey in northern Sweden at the end of the Neolithic period, and discusses whether humans targeted female moose in hunting. The fourth study analyses and discusses the history of the harp seal in the Baltic Sea. The fifth study is a methodological paper which involves identifying seals according to sex, using the dog genome. The overall result of the different case studies shows that there were major population fluctuations over time in all the species studied, and that in some cases, humans are likely to have contributed to this, e.g. through overhunting and translocation. The study also shows that the population fluctuations often occurred in connection with certain climatic events, though it was not possible to separate climatic effects from human impact in terms of the cause.
... Other studies (see Thomas 1914, Cabrera 1958, in Wozencraft, 2005 classified the culpeo in the genus of Dusicyon (Smith 1839), although this genus is considered distantly related to Lycalopex by several authors (Bininda-Emonds et al., 1999;Slater et al., 2009;Austin et al., 2013). However, Perini et al. (2010) still suggest that L. culpaeus and the extinct guará (Dusicyon australis) may be congeneric species, in which case the generic name Dusicyon would be more appropriate for culpeo. ...
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... Datos paleobotánicos indican que en las Malvinas existió un bosque de Podocarpus en el pre-Neógeno (Birnie & Roberts, 1986). Por otro lado, evidencia geológica sugiere que durante las diversas glaciaciones del Neógeno y el Cuaternario, las Malvinas estuvieron conectadas al continente (Posadas & Morrone, 2004;Austin et al., 2013). Esta evidencia permitió postular eventos de vicarianza y geodispersión entre las islas y el continente (Morrone & Posadas, 2005), dado que estuvieron conectadas a la masa continental la menos durante cinco eventos, en donde el nivel del mar fue de 100 m por debajo del nivel actual, permitiendo la conexión biótica. ...
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Evolutionary biogeography identifies areas of endemism and establishes their relationships with other areas, providing the information required to develop a hierarchical system of natural regionalization. The closely related geological and biological evolution of the planet is manifested in the existence of endemic biotas, as result of geographical and ecological barriers. The current challenge of evolutionary biogeography is to document the existence of biotas representing the evolutionary structure of ecosystems and their functionality, that could contribute to the establishment of conservation priorities. In this contribution, the fundamental characteristics of the biogeographic units of Argentina are described, and the respective maps are provided, as a general reference system for ecological, evolutionary and biogeographic studies, and in education and decision-making regarding conservation and sustainable use. For each biogeographic unit, we present its valid name and synonyms, its geographic location, distinctive characteristics, dominant landscape and vegetation types and its endemic species. Also, the typical landscapes and some endemic or characteristic species of each biogeographical unit are illustrated. Their biotic relationships, geobiotic evolution and regionalization to the district category are also discussed. In the scheme presented herein 16 provinces are recognized in the country, which in turn are grouped in the Neotropical region (Yungas, Parana, Araucaria Forest, Esteros del Iberá, Chaco and Pampean provinces), South American Transition Zone (Puna, Cuyan High Andean, Monte and Comechingones provinces) and Andean region (Patagonian, Maule, Valdivian Forest, Magellanic Forest, Falkland (Malvinas) Islands and Magellanic Moorland provinces). The Yungas province has three districts: Transition Forests, Montane Jungles, and Montane Forests. The Parana province includes the Campos and Mixed Forests districts. The Esteros del Iberá province stat. nov. includes three districts: Delta of Paraná stat. rev., Uruguay River stat. nov. and Paraná Flooded Savannas stat. nov. The Chaco province includes the Eastern Chacoan, Montane Chacoan stat. rev. and Western Chacoan districts. In the Pampean province five districts are recognized: Austral Pampean, Eastern Pampean, Espinal, Western Pampean and Uruguayan. In the Argentinean part of the Puna province the Jujuyan district is recognized. In the Cuyan High Andean province three districts are identified: Diaguita nom. nov., Cuyan ubic. nov. and Huarpe nom. nov. The Monte province includes four districts: Prepuna, Northern, Eremean and Southern. In the Patagonian province we recognize five subprovinces: Central Patagonian (Chubut and Santa Cruz districts), Fuegian, Payunia (Northern and Southern Payunia districts), Subandean (Meridional Subandean Patagonia, Austral High Andean, and Septentrional Subandean Patagonia districts) and Western Patagonian. In the Argentinean sector of the Maule province is present the Pehuén district. The Valdivian Forest province is represented by the Valdivian district. The Falkland (Malvinas) Islands province has two districts: Falkland Islands and South Georgia Islands.
... To exemplify this approach, we consider whether nonnative South American grey foxes (Lycalopex griseus, hereafter SAGF) act as de facto ecological replacements for extinct Falkland Island wolves (Dusicyon australis, hereafter FIW; Figure S1, Table S3). FIWs likely reached the Falklands from mainland South America naturally at least 16,000 years ago (Austin et al., 2013). Prior to their extirpation by man around 1876, they were the archipelago's only endemic land mammal and large terrestrial predator. ...
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Ecological replacement as a conservation tool presupposes that nonnative taxa can restore degraded ecosystems by performing the ecological functions of extinct taxa. This assumption is rarely tested however, largely because it is difficult to quantify the functions of species extirpated long ago. Here, we test whether feral South American grey foxes (SAGF), introduced to the Falkland Islands ∼90 years ago, act as unintended ecological replacements for endemic Falkland Islands wolves (FIW), extirpated during the 19th century. Using hair stable isotope ratios as proxies for diet, we show that the isotopic niche space of modern SAGFs almost completely encompasses that of archaic FIWs. However, the former's niche is larger so while SAGFs may play similar ecological roles to FIWs, they probably perform additional functions, which may or may not be desirable. In so doing, we illustrate a generalized framework for using comparative isotopic niche analysis to test for ecological replacement objectively.
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Over the past decade research into early domestication has been transformed by the genomics revolution and increased archaeological investigation. Despite clarification of the timing, locations and genetic processes, most scholars still envision evolutionary responses to human innovations, such as sickle harvesting, tilling, selection for docility or directed breeding. Stepping away from anthropocentric models, evolutionary parallels in the wild can provide case studies for understanding what ecological pressures drove the evolution of the first domestication traits. I contrast evolutionary trends seen among plants and animals confined on oceanic islands with the changes seen in the first cultivated crops and animals. I argue that the earliest villages functioned as habitat islands, applying parallel selective pressures as those on oceanic islands. In this view, the collective assemblage of parallel evolving traits that some scholars refer to as either an island syndrome or domestication syndrome results from similar ecological pressures of insularity, notably ecological release. © 2022 The Author. Oikos published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos.
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Stable isotopes from archaic Falkland Islands wolves ( Dusicyon australis ) indicate a high trophic, marine diet. Hamley et al. argue that this is consistent with mutualism with Yaghan people. However, most D. australis had similar isotopic signatures in the European era, despite human persecution. These data therefore neither support nor refute human-mediated introduction of D. australis to the Falklands.
Thesis
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Public awareness of nature and environmental issues has grown in the last decades and zoos have successfully followed suit by re-branding themselves as key representatives for conservation. However, considering the fast rate of environmental degradation, in the near future, zoos may become the only place left for wildlife. Some scholars argue that we have entered a new epoch titled the “Anthropocene” that postulates the idea that untouched pristine nature is almost nowhere to be found.1 Many scientists and scholars argue that it is time that we embraced this environmental situation and anticipated the change. 2 Clearly, the impact of urbanization is reaching into the wild, so how can we design for animals in our artificializing world? Using the Manoa School method that argues that every future includes these four, generic, alternatives: growth, discipline, collapse, and transformation3, this dissertation explores possible future animal archetypes by considering multiple possibilities of post zoo design.
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Events during the Late Oligocene of North America and the Pleistocene of South America, provide an opportunity for exploring possible causes of the evolution of hypercarnivory in canids. Plots of generic diversity gainst time for North American predators reveal a roughly inverse relationship between the number of hypercarnivorous canid taxa and the numbers of other hypercarnivores, such as creodonts, nimravids, mustelids, and amphicyonids. Similarly, the radiation of hypercarnivorous canids in South America occurred at a time of relatively low diversity of other hypercarnivores. Analysis of trophic diversity within the North American carnivore paleoguild before, during, and after the Late Oligocene reveals considerable taxonomic turnover among carnivores because of immigration and speciation. Late Oligocene hypercarnivorous canids appear to have been replaced first by amphicyonids and large mustelids, and then by felids. -from Author
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Estimating genealogical relationships among genes at the population level presents a number of difficulties to traditional methods of phylogeny reconstruction. These traditional methods such as parsimony, neighbour-joining, and maximum-likelihood make assumptions that are invalid at the population level. In this note, we announce the availability of a new software package, TCS, to estimate genealogical relationships among sequences using the method of Templeton et al. (1992) .
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Members of the Canidae are known from the late Pliocene and early Pleistocene (Uquian) through the Recent in South America. Ten genera and 28 species of wolves and foxes are represented. Cladistic analysis supports recognition of four monophyletic groups: 1) Urocyon; 2) Dusicyon; 3) Cerdocyon and 4) Chrysocyon including Canis. Zoogeographic implications of the cladistic hypotheses presented here are supported by the fossil record, suggesting the origin of canids in North America and their subsequent dispersal and extensive radiation in South America. The extinction of large canids in South America at the end of the Pleistocene is a consequence of extinction of their specialized large herbivorous prey. The current high diversity of South American foxes is, at least in part, the result of an opportunistic feeding strategy that utilizes small prey as well as fruits and grains. -from Author
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The main features of the phylogeny program TNT are discussed. Windows versions have a menu interface, while Macintosh and Linux versions are command-driven. The program can analyze data sets with discrete (additive, non-additive, step-matrix) as well as continuous ...
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The Age Calibration Program, CALIB, published in 1986 and amended in 1987 is here amended anew. The program is available on a floppy disk in this publication. The new calibration data set covers nearly 22 000 Cal yr (approx 18 400 14C yr) and represents a 6 yr timescale calibration effort by several laboratories. The data are described and the program outlined. -K.Clayton