Ancient spider cousin had 5 sets of jaws for crushing its prey

Ancient spider cousin had 5 sets of jaws for crushing its prey

Back when the ancient seas teemed with trilobites, the little ones may have had nightmares about this fearsome predator.

It now appears that Habelia optata , which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

, which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

Those animals belong to a group of animals called chelicerates, and the new study on Habelia tells us a lot about what their ancestors may have been like.

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the study published this week in BMC Evolutionary Biology . “It’s like a centipede or perhaps an insect that would have not one pair of mandibles, but five.”

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the

Each of those powerful pairs of jaws was equipped with sharp teeth and designed to crush the protective shells of its prey, likely small trilobites. In fact, a similar fossil predator was recently found in Australia with chopped up trilobite remains in its gut, said Aria, who studied Habelia during his PhD at the University of Toronto and is now a postdoctoral researcher at the Nanjing Institute of Geology and Paleontology in China.

Fossil specimen of Habelia optata from the Royal Ontario Museum. This specimen shows some of the very large jaws under the head shield. Note also the long dorsal spines on the thorax. (Jean-Bernard Caron/Royal Ontario Museum)

But despite its fearsome features, Habelia was only two centimetres long and likely an appetizing snack for larger predators, as evidenced by its defensive features, said Jean-Bernard Caron, senior curator of invertebrate paleontology at the Royal Ontario Museum and Aria’s supervisor.

“It has a crazy number of spines along the body — pretty much from tail to head it has spines everywhere.”

The first specimens of Habelia were collected from the Burgess Shale of Yoho National Park in B.C. and described in 1912 by Charles Walcott. He was the paleontologist who first discovered the 508-million-year-old fossil beds that contain beautifully preserved remains of a huge variety of creatures that lived in a shallow sea during the middle Cambrian period.

New specimens of Habelia optata were collected during field expeditions to a fossil site called the Walcott Quarry in Yoho National Park in B.C. in the 1990s. (Desmond Collins/Royal Ontario Museum)

But for more than a century, paleontologists were puzzled as to what, exactly, Habelia was. Its mandibles made it look like it could be related to insects, which belong to a group of creatures called mandibulates. But many of its other features were unusual.

In the 1990s, ROM paleontologist Desmond Collins led several expeditions to the Walcott Quarry and collected dozens of other specimens.

Caron chose specimens in different orientations and, peering through a microscope, used microengraving tools to painstakingly chisel the rock away from its minuscule appendages to reveal hidden features.

The creatures had been squashed flat during the fossilization process, so figuring out what they looked like in 3D was a challenge.

Aria carefully measured the distances between different appendages and worked with artist Joanna Liang, a master’s student in the biomedical communications program at the University of Toronto, to reconstruct the animal in 3D. In some ways, he said, it’s similar to the way a dinosaur fossil might be put back together from its individual bones. In this case, he said, the art was a crucial step in the science.

A very small pair at the very front called “chelicerae” that are found in all chelicerates, including spiders. They were key to placing Habelia in what researchers believe is its rightful place in the tree of life.

Simplified phylogeny (tree of life) showing the relationship of Habelia with other groups of arthropods. The study shows that it is an early relative of chelicerates — a group including spiders, scorpions, horseshoe crabs and mites. (Cedric Aria)

While the head appendages on modern chelicerates are used for walking, Habelia appeared to use them for grasping and crushing prey.

It had another five pairs of legs for walking on its thorax, a part of the body where spiders and scorpions don’t have any appendages.

While animals that lived far in the past and are high up in their family trees are often described as primitive, Aria says that word is definitely not appropriate for Habelia: “It’s more complex than a lot of chelicerates that live today.”

The study was funded by the University of Toronto and the Natural Sciences and Engineering Research Council of Canada.

See Also : ancient spider cousin had 5 sets of jaws for crushing its prey 2.html

Back when the ancient seas teemed with trilobites, the little ones may have had nightmares about this fearsome predator.

It now appears that Habelia optata , which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

, which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

Those animals belong to a group of animals called chelicerates, and the new study on Habelia tells us a lot about what their ancestors may have been like.

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the study published this week in BMC Evolutionary Biology . “It’s like a centipede or perhaps an insect that would have not one pair of mandibles, but five.”

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the

Each of those powerful pairs of jaws was equipped with sharp teeth and designed to crush the protective shells of its prey, likely small trilobites. In fact, a similar fossil predator was recently found in Australia with chopped up trilobite remains in its gut, said Aria, who studied Habelia during his PhD at the University of Toronto and is now a postdoctoral researcher at the Nanjing Institute of Geology and Paleontology in China.

Fossil specimen of Habelia optata from the Royal Ontario Museum. This specimen shows some of the very large jaws under the head shield. Note also the long dorsal spines on the thorax. (Jean-Bernard Caron/Royal Ontario Museum)

But despite its fearsome features, Habelia was only two centimetres long and likely an appetizing snack for larger predators, as evidenced by its defensive features, said Jean-Bernard Caron, senior curator of invertebrate paleontology at the Royal Ontario Museum and Aria’s supervisor.

“It has a crazy number of spines along the body — pretty much from tail to head it has spines everywhere.”

The first specimens of Habelia were collected from the Burgess Shale of Yoho National Park in B.C. and described in 1912 by Charles Walcott. He was the paleontologist who first discovered the 508-million-year-old fossil beds that contain beautifully preserved remains of a huge variety of creatures that lived in a shallow sea during the middle Cambrian period.

New specimens of Habelia optata were collected during field expeditions to a fossil site called the Walcott Quarry in Yoho National Park in B.C. in the 1990s. (Desmond Collins/Royal Ontario Museum)

But for more than a century, paleontologists were puzzled as to what, exactly, Habelia was. Its mandibles made it look like it could be related to insects, which belong to a group of creatures called mandibulates. But many of its other features were unusual.

In the 1990s, ROM paleontologist Desmond Collins led several expeditions to the Walcott Quarry and collected dozens of other specimens.

Caron chose specimens in different orientations and, peering through a microscope, used microengraving tools to painstakingly chisel the rock away from its minuscule appendages to reveal hidden features.

The creatures had been squashed flat during the fossilization process, so figuring out what they looked like in 3D was a challenge.

Aria carefully measured the distances between different appendages and worked with artist Joanna Liang, a master’s student in the biomedical communications program at the University of Toronto, to reconstruct the animal in 3D. In some ways, he said, it’s similar to the way a dinosaur fossil might be put back together from its individual bones. In this case, he said, the art was a crucial step in the science.

A very small pair at the very front called “chelicerae” that are found in all chelicerates, including spiders. They were key to placing Habelia in what researchers believe is its rightful place in the tree of life.

Simplified phylogeny (tree of life) showing the relationship of Habelia with other groups of arthropods. The study shows that it is an early relative of chelicerates — a group including spiders, scorpions, horseshoe crabs and mites. (Cedric Aria)

While the head appendages on modern chelicerates are used for walking, Habelia appeared to use them for grasping and crushing prey.

It had another five pairs of legs for walking on its thorax, a part of the body where spiders and scorpions don’t have any appendages.

While animals that lived far in the past and are high up in their family trees are often described as primitive, Aria says that word is definitely not appropriate for Habelia: “It’s more complex than a lot of chelicerates that live today.”

The study was funded by the University of Toronto and the Natural Sciences and Engineering Research Council of Canada.

The rare animal Habelia optata , which had originally been described in 1912, had remained one of the most problematic fossils from the middle Cambrian Burgess Shale–the 508 million years old exceptional fossil deposit of British Columbia (see Habelia optata on the ROM Burgess Shale website for an overview). Our new study is part of a wider reappraisal of Burgess Shale arthropods–invertebrate animals with segmented and articulated bodies–, which I had worked on as part of my doctoral degree at the University of Toronto/Royal Ontario Museum (ROM). My former supervisor Jean-Bernard Caron , senior curator of invertebrate paleontology at the ROM and I, have now redescribed this species in detail mainly based on abundant new material housed at the ROM and also the original Walcott collection at the National Museum of Natural History in Washington, D.C.

, which had originally been described in 1912, had remained one of the most problematic fossils from the middle Cambrian Burgess Shale–the 508 million years old exceptional fossil deposit of British Columbia (see

on the ROM Burgess Shale website for an overview). Our new study is part of a wider reappraisal of Burgess Shale arthropods–invertebrate animals with segmented and articulated bodies–, which I had worked on as part of my doctoral degree at the University of Toronto/Royal Ontario Museum (ROM). My former supervisor

, senior curator of invertebrate paleontology at the ROM and I, have now redescribed this species in detail mainly based on abundant new material housed at the ROM and also the original Walcott collection at the National Museum of Natural History in Washington, D.C.

Fossil specimen of Habelia optata from the Royal Ontario Museum. This specimen spectacularly shows some of the very large jaws (“gnathobases”) under the head shield. Note also the long dorsal spines on the thorax. ROMIP64359.

from the Royal Ontario Museum. This specimen spectacularly shows some of the very large jaws (“gnathobases”) under the head shield. Note also the long dorsal spines on the thorax. ROMIP64359.

In total, 27 specimens of Habelia were collected in the 1990s by the ROM under several Parks Canada Research and Collection permits to retired Senior Curator Desmond Collins. Specimens come from the famous Walcott Quarry, which is the main Burgess Shale site, located in Yoho National Park, British Columbia. Specimens were mechanically prepared at the ROM to reveal concealed elements still buried in the matrix, and then carefully observed under a microscope and photographed using different lighting techniques.

were collected in the 1990s by the ROM under several Parks Canada Research and Collection permits to retired Senior Curator Desmond Collins. Specimens come from the famous Walcott Quarry, which is the main Burgess Shale site, located in Yoho National Park, British Columbia. Specimens were mechanically prepared at the ROM to reveal concealed elements still buried in the matrix, and then carefully observed under a microscope and photographed using different lighting techniques.

This is a small specimen showing the long, bipartite tailpiece (on the left) and most of the appendages in great detail–including jaws with long teeth under the head shield. The ramified projections at the very front (right) are interpreted as the partially “detached” outer branches of the head appendages. ROMIP64357.

Joanna Liang was then tasked to work on detail technical drawings, including 3D animations, as part of her Master of Science in Biomedical Communications, a program offered through the Institute of Medical Science in the Faculty of Medicine at the University of Toronto. The process took close to 18 months of back and forth between us, the palaeontologists and the illustrator, to ultimately yield what could be seen as one of the most technically challenging 3D reconstruction of a Cambrian animal, among those described thus far.

Joanna Liang was then tasked to work on detail technical drawings, including 3D animations, as part of her Master of Science

in the Faculty of Medicine at the University of Toronto. The process took close to 18 months of back and forth between us, the palaeontologists and the illustrator, to ultimately yield what could be seen as one of the most technically challenging 3D reconstruction of a Cambrian animal, among those described thus far.

Detailed diagrams of Habelia optata (type A). A. Ventral view of the head. Right posterior appendage removed to show the morphology of the jaws (“gnathobases”). B. Lateral view. C. Dorsal view. D. Isolated limbs of the thorax in frontal, lateral and posterior views (left to right).

(type A). A. Ventral view of the head. Right posterior appendage removed to show the morphology of the jaws (“gnathobases”). B. Lateral view. C. Dorsal view. D. Isolated limbs of the thorax in frontal, lateral and posterior views (left to right).

Animation of Habelia optata – Head, jaws, and limbs. Demonstrating the complexity of its feeding mechanism Credit: Animated by Joanna Liang, (c) Royal Ontario Museum

– Head, jaws, and limbs. Demonstrating the complexity of its feeding mechanism Credit: Animated by Joanna Liang, (c) Royal Ontario Museum

Animation of Habelia optata – 360 view. Illustrating the spectacular architecture of its body. Credit: Animated by Joanna Liang, (c) Royal Ontario Museum

– 360 view. Illustrating the spectacular architecture of its body. Credit: Animated by Joanna Liang, (c) Royal Ontario Museum

Short film showing the process of reconstructing Habelia optata–from fossil observations to the 3D model. Credit: Animated by Joanna Liang, (c) Royal Ontario Museum

In brief, the extremely complex head of Habelia formed a surprisingly efficient sensory, raptorial and masticatory device suited to a durophagous diet (diet based on hard carapaces and shells). We interpreted the species as an active predator that lived at the bottom of the Cambrian seas and hunted small preys, possibly including small trilobites. Our study shows that Habelia optata belongs to chelicerates, one of the two main groups of modern arthropods (including spiders, scorpions, horseshoe crabs, mites) and illustrates a spectacular and unique morphological convergence with species of the other sub-group of modern arthropods: the mandibulates (including shrimps, crabs, centipedes, insects). Morphological convergence is a phenomenon that involves the evolution of similarities between features, despite different origins, usually because they fulfill comparable functions. Wings in pterosaurs, bats and birds, for instance, are convergent since they have evolved independently in each of these groups.

formed a surprisingly efficient sensory, raptorial and masticatory device suited to a durophagous diet (diet based on hard carapaces and shells). We interpreted the species as an active predator that lived at the bottom of the Cambrian seas and hunted small preys, possibly including small trilobites. Our study shows that

belongs to chelicerates, one of the two main groups of modern arthropods (including spiders, scorpions, horseshoe crabs, mites) and illustrates a spectacular and unique morphological convergence with species of the other sub-group of modern arthropods: the mandibulates (including shrimps, crabs, centipedes, insects). Morphological convergence is a phenomenon that involves the evolution of similarities between features, despite different origins, usually because they fulfill comparable functions. Wings in pterosaurs, bats and birds, for instance, are convergent since they have evolved independently in each of these groups.

Convergence goes even further in Habelia , because functional similarities with mandibulates mostly evolved from different limbs.

Figure showing convergent evolution of head features between Habelia (A) and predatory mandibulates (B: isopod, C: centipede). Although these traits in Habelia had a different evolutionary origin, they fulfilled similar functions. Green: sensory/tactile; Orange: masticatory; Blue: holding/manipulating.

had a different evolutionary origin, they fulfilled similar functions. Green: sensory/tactile; Orange: masticatory; Blue: holding/manipulating.

Habelia optata is a well-armoured arthropod of about 2 cm in length with a tail as long as the rest of the body, and made of two articulated pieces. Its exoskeleton is covered in numerous blunt spines/tubercles and long spines also adorn the back behind the head.

is a well-armoured arthropod of about 2 cm in length with a tail as long as the rest of the body, and made of two articulated pieces. Its exoskeleton is covered in numerous blunt spines/tubercles and long spines also adorn the back behind the head.

The body is divided into head, thorax and post-thorax (that is, not an abdomen in the technical sense). The head is particularly complex and contains three different types of appendages: a reduced anteriormost pair thought to be the precursor of chelicerae; a series of five pairs of highly modified appendages fulfilling various functions, and a seventh, posteriormost, larger pair. The appendages in the series of five are made of: a large plate with teeth (gnathobase) for mastication; a leg-like branch with setae and spines for grasping; and an elongate, slender branch modified as a sensory/tactile appendage. This sensory branch is not fully “attached” to the rest of the appendage as it should be (how it is attached to the head is not known), a very peculiar condition in arthropods.

The thorax bears five pairs of biramous walking legs. The post-thorax bears rounded appendages likely used in respiration.

Among chelicerates are very familiar animals: spiders, scorpions, harvestmen, mites and horseshoe crabs. They are united by a head bearing five pairs of legs and an anteriormost pair of chelate appendages used to cut their food: the chelicerae. These traits distinguish them from the other large group of modern arthropods, the mandibulates–such as shrimps, crabs, centipedes or insects–which by contrast have sensorial antennae and modified head limbs assisting in food mastication. Mandibulates that feed on other arthropods or shelly animals in particular (i.e. that are durophagous), such as lobsters, crabs or centipedes, usually have these head appendages forming series of plates (and especially the mandibles), providing them with strong crushing abilities.

In revisiting Habelia optata , notably with the help of unpublished fossil material from the Royal Ontario Museum, we found that this long problematic arthropod from the Burgess Shale was in fact a close relative of the ancestor of all chelicerates. Although they do not seem to form pincers, the anteriormost limbs of Habelia are small and distinct from a series of five other pairs of head appendages, and hence likely ancestral forms of the chelicerae. A seventh pair of appendages at the back of the head also helps explain the origin of corresponding modified structures in chelicerates, such as the small appendages called “chilaria” in horseshoe crabs.

, notably with the help of unpublished fossil material from the Royal Ontario Museum, we found that this long problematic arthropod from the Burgess Shale was in fact a close relative of the ancestor of all chelicerates. Although they do not seem to form pincers, the anteriormost limbs of

are small and distinct from a series of five other pairs of head appendages, and hence likely ancestral forms of the chelicerae. A seventh pair of appendages at the back of the head also helps explain the origin of corresponding modified structures in chelicerates, such as the small appendages called “chilaria” in horseshoe crabs.

Simplified phylogeny (‘tree of life’) showing the relationship of Habelia with other groups of arthropods. The study shows that it is an early relative of chelicerates — a group including spiders, scorpions, horseshoe crabs and mites.

with other groups of arthropods. The study shows that it is an early relative of chelicerates — a group including spiders, scorpions, horseshoe crabs and mites.

However, we also found that Habelia converged remarkably with durophagous mandibulates: the head appendages behind the “proto-chelicerae” are smaller and not walking legs, but spinose and raptorial, and are attached to exceptionally large masticating plates called gnathobases. Unlike horseshoe crabs for instance, these plates overlap and close parallel to the ventral surface of the head–much as in mandibulates. The outer branches of these appendages (exopods), absent in modern chelicerates, are here present and modified into slender sensory apparatuses, recalling the antennae of mandibulates. Furthermore, the seventh pair of appendages at the back of the head seems to have fulfilled a function analog to that of the maxillipeds–additional appendages in mandibulates assisting the other head limbs in the processing of food.

converged remarkably with durophagous mandibulates: the head appendages behind the “proto-chelicerae” are smaller and not walking legs, but spinose and raptorial, and are attached to exceptionally large masticating plates called gnathobases. Unlike horseshoe crabs for instance, these plates overlap and close parallel to the ventral surface of the head–much as in mandibulates. The outer branches of these appendages (exopods), absent in modern chelicerates, are here present and modified into slender sensory apparatuses, recalling the antennae of mandibulates. Furthermore, the seventh pair of appendages at the back of the head seems to have fulfilled a function analog to that of the maxillipeds–additional appendages in mandibulates assisting the other head limbs in the processing of food.

It is important to note that these resemblances are not entirely due to the proximity of Habelia to the point of divergence between chelicerates and mandibulates: they are in part secondary modifications of features that were already characteristically chelicerate in nature. Combined with other pieces of evidence, such as the high number of head appendages, the modifications of limb branches into pseudo-antennae, and the shape of trunk segments, this suggests that the origin of chelicerates was marked by a remarkable morphological plasticity.

to the point of divergence between chelicerates and mandibulates: they are in part secondary modifications of features that were already characteristically chelicerate in nature. Combined with other pieces of evidence, such as the high number of head appendages, the modifications of limb branches into pseudo-antennae, and the shape of trunk segments, this suggests that the origin of chelicerates was marked by a remarkable morphological plasticity.

From this outstanding head structure, as well as from well-developed walking legs in its trunk, we can picture Habelia and its relatives (known from less complete remains in the USA and Australia) as small active predators of the Cambrian sea floors, hunting for small or immature trilobites and other shelly animals. This builds onto the importance of carapaces and shells for evolutionary change during the Cambrian Explosion, and expands our understanding of ecosystems at this time, showing another level of predator-prey relationship and its determining impact on the rise of modern arthropods.

From this outstanding head structure, as well as from well-developed walking legs in its trunk, we can picture

and its relatives (known from less complete remains in the USA and Australia) as small active predators of the Cambrian sea floors, hunting for small or immature trilobites and other shelly animals. This builds onto the importance of carapaces and shells for evolutionary change during the Cambrian Explosion, and expands our understanding of ecosystems at this time, showing another level of predator-prey relationship and its determining impact on the rise of modern arthropods.

Back when the ancient seas teemed with trilobites, the little ones may have had nightmares about this fearsome predator.

It now appears that Habelia optata , which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

, which lived about 508 million years ago in what is now eastern B.C., was a relative of spiders and scorpions, researchers at the Royal Ontario Museum and the University of Toronto have discovered.

Those animals belong to a group of animals called chelicerates, and the new study on Habelia tells us a lot about what their ancestors may have been like.

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the study published this week in BMC Evolutionary Biology . “It’s like a centipede or perhaps an insect that would have not one pair of mandibles, but five.”

“If you’re looking for a scary Hollywood creature, it probably would be the perfect one,” said Cedric Aria, lead author of the

Each of those powerful pairs of jaws was equipped with sharp teeth and designed to crush the protective shells of its prey, likely small trilobites. In fact, a similar fossil predator was recently found in Australia with chopped up trilobite remains in its gut, said Aria, who studied Habelia during his PhD at the University of Toronto and is now a postdoctoral researcher at the Nanjing Institute of Geology and Paleontology in China.

Fossil specimen of Habelia optata from the Royal Ontario Museum. This specimen shows some of the very large jaws under the head shield. Note also the long dorsal spines on the thorax. (Jean-Bernard Caron/Royal Ontario Museum)

But despite its fearsome features, Habelia was only two centimetres long and likely an appetizing snack for larger predators, as evidenced by its defensive features, said Jean-Bernard Caron, senior curator of invertebrate paleontology at the Royal Ontario Museum and Aria’s supervisor.

“It has a crazy number of spines along the body — pretty much from tail to head it has spines everywhere.”

The first specimens of Habelia were collected from the Burgess Shale of Yoho National Park in B.C. and described in 1912 by Charles Walcott. He was the paleontologist who first discovered the 508-million-year-old fossil beds that contain beautifully preserved remains of a huge variety of creatures that lived in a shallow sea during the middle Cambrian period.

New specimens of Habelia optata were collected during field expeditions to a fossil site called the Walcott Quarry in Yoho National Park in B.C. in the 1990s. (Desmond Collins/Royal Ontario Museum)

But for more than a century, paleontologists were puzzled as to what, exactly, Habelia was. Its mandibles made it look like it could be related to insects, which belong to a group of creatures called mandibulates. But many of its other features were unusual.

In the 1990s, ROM paleontologist Desmond Collins led several expeditions to the Walcott Quarry and collected dozens of other specimens.

Caron chose specimens in different orientations and, peering through a microscope, used microengraving tools to painstakingly chisel the rock away from its minuscule appendages to reveal hidden features.

The creatures had been squashed flat during the fossilization process, so figuring out what they looked like in 3D was a challenge.

Aria carefully measured the distances between different appendages and worked with artist Joanna Liang, a master’s student in the biomedical communications program at the University of Toronto, to reconstruct the animal in 3D. In some ways, he said, it’s similar to the way a dinosaur fossil might be put back together from its individual bones. In this case, he said, the art was a crucial step in the science.

A very small pair at the very front called “chelicerae” that are found in all chelicerates, including spiders. They were key to placing Habelia in what researchers believe is its rightful place in the tree of life.

Simplified phylogeny (tree of life) showing the relationship of Habelia with other groups of arthropods. The study shows that it is an early relative of chelicerates — a group including spiders, scorpions, horseshoe crabs and mites. (Cedric Aria)

While the head appendages on modern chelicerates are used for walking, Habelia appeared to use them for grasping and crushing prey.

It had another five pairs of legs for walking on its thorax, a part of the body where spiders and scorpions don’t have any appendages.

While animals that lived far in the past and are high up in their family trees are often described as primitive, Aria says that word is definitely not appropriate for Habelia: “It’s more complex than a lot of chelicerates that live today.”

The study was funded by the University of Toronto and the Natural Sciences and Engineering Research Council of Canada.

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