irregularbooks.

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Acknowledgements
We are thankful to our families and friends, whose love and support helped us through the
making of this book. Several of our colleagues have provided feedback and discussion that
helped shape our ideas. In particular Mike Taylor and Ma Wedel were generous with their
encouragement and discussion.

We thank the late Dan Varner for discussion on the history of palaeoart; and Tim Isles, Luis
Rey, Mark Wi on and Steve White for thoughts on palaeobiology and palaeoart.

We thank John's wife, Jenny, for her support and proof-reading.

C. M. Kosemen would like to extend special thanks towards his parents, brother and sister
for their unconditional love, support and friendship.
About the Authors
John Conway
Artist & Author

John Conway is a palaeontological and fine artist, who's work has

been used for National Geographic, Discovery Channel and the
American Museum of Natural History, among others. His work has
most recently appeared in Dinosaur Art: the World's Greatest Paleoart.
John's interest in the methodology and culture of reconstructing of
palaeontological subjects is the genesis of this book.

Website: johnconway.co
Twi er: @nyctopterus
Facebook: facebook.com/nyctopterus

C.M. Kosemen
Artist & Author

C. M. Kosemen holds a Media and Communications Masters' degree

from Goldsmiths College, and has worked as an editor in Bene on
Company's Colors magazine. He has had several exhibitons of his
evolution-themed fine art at galleries and science festivals
internationally. Kosemen's areas of specialization are speculative &
real zoology, history and unusual things in general. His previous
work includes Snaiad, a self-initiated web project about life on an
alien planet.
Website: www.cmkosemen.com
Facebook: facebook.com/memo.kosemen

Darren Naish
Author

Darren Naish is a science writer, technical editor and palaeozoologist.

Darren works mostly on theropod and sauropods dinosaurs, but also
works on pterosaurs, marine reptiles, and other tetrapods. With
colleagues, he named the dinosaurs Eotyrannus, Mirischia and
Xenoposeidon. Darren has wri en several books, including Walking
With Dinosaurs: The Evidence (co-authored with David M. Martill),
Great Dinosaur Discoveries, and more recently Tetrapod Zoology Book
One. His blog, Tetrapod Zoology, is widely considered the world's
foremost zoology blog.

When not writing about tetrapods, Darren can be found pursuing his
interest in modern wildlife and conservation and its resulting
adventures in lizard-chasing, bird-watching and li er-collecting.

Website: blogs.scientificamerican.com/tetrapod-zoology/
Twi er: @TetZoo

Sco Hartman
Illustrator

Sco Hartman is a palaeontologist and illustrator who specialises in

creating skeletal diagrams of dinosaurs and other animals to the
highest degree of scientific accuracy. Sco 's skeletal drawings form
the basis of many other artist's work.
Website: skeletaldrawing.com
Twi er: @skeletaldrawing
Dedicated to Dan Varner
 All Yesterdays
Unique and Speculative Views of Dinosaurs and Other Prehistoric
Animals

By John Conway, C. M. Kosemen and Darren Naish with skeletal

reconstructions by Sco Hartman
Introduction, by Darren Naish
A desire to imagine the long-extinct organisms of the past as living,
moving animals has long inspired artists and scientists to clothe
bones and other fossil tissues in muscles, skin, fur and feathers. In
other words, to bring fossil animals 'back to life' in art. We need to be
clear from the start that-while there are many things that we're surely
ge ing right-there are many other things that have to be regarded as
'known unknowns', and even as 'unknown unknowns'. It is these
areas of doubt and speculation that form the focus of this book, the
first ever devoted entirely to the more speculative aspects of
palaeoart.1

It is well known that the process of reconstructing a fossil animal

involves a marriage of both ‘hard’ data as well as a degree of
informed speculation. That ‘hard’ data involves such things as the
lengths and widths of bones and other hard parts, and the positions
of specific muscle groups present in living animals. While the
creation of a bone-and-muscles-only reconstruction should be seen as
the first step in the depiction of a fossil animal (and even as a
presumably inescapable part of creating a reconstruction!), readers
may be surprised to learn that many people who have reconstructed
extinct animals have frequently done so without recourse to these
vital steps.

We in fact know that this was true of some of the greatest and most
influential palaeoartists of all time. The Czech master of ancient
animals and landscapes Zdenîk Burian (1905-1981), for example, best-
guessed the life appearance of dinosaurs and other vertebrates by
fleshing out museum-mounted skeleton on paper without the use of
measurements. Rudolph F. Zallinger’s (1919-1995) animals – most
famously depicted in the Zallinger mural at Yale’s Peabody Museum
of Natural History – were clearly done with only a superficial
reference to the skeletons of the species concerned. The pieces of art
generated by these individuals remain brilliant, beautiful and
wonderful, but the techniques they used were damaging to the
contention that the reconstruction of fossil animals involves science
as much as it does art. Indeed, this concept is reflected in the
paraphrased claim “There’s more than one way to reconstruct a
dinosaur”, and in the general idea that dinosaurs and other fossil
animals can only be reconstructed approximately, or with substantial
doubt about the most basic issues remaining.

So we must note to begin with that reconstructing a fossil animal is

not a speculative process that has many possible outcomes, but a
rigorous and evidence-led one where informed artists produce a
technically accurate musculoskeletal reconstruction for a given
animal. The problem comes with the integument – the covering that
involves the skin and all the things a ached to it (scales, feathers,
hairs and so on) – as we'll see.

Most people interested in palaeoart are aware of, and follow, the
high-fidelity musculoskeletal reconstructions produced by researcher
and artist Greg Paul. Many of Paul's hypotheses and arguments
about archosaur2 biology and evolution are the topic of argument
and disagreement, but his explanations and illustrations of archosaur
anatomy and the way he restores an animal's musculoskeletal system
remain important. Paul's 1987 article 'The science and art of restoring
the life appearance of dinosaurs and their relatives: a rigorous how-to
guide'3 remains a classic, and (while now very dated) it is probably
the best and most useful introduction to the sort of information an
artist would need (though read on).
Paul's initial forays into the accurate reconstruction of archosaur
musculature were the result of communication with Robert Bakker;
in turn, both Paul and Bakker were inspired by Charles R. Knight's
(1874-1953) discussions and depictions of animal anatomy. Knight is
most famous for his many paintings of fossil dinosaurs and
mammals, but he also illustrated living animals. His book—Animal
Drawing: Anatomy and Action for Artists—should be obtained by
anyone seriously interested in the subject.4 Therein, we see Knight's
excellent a ention to anatomical detail (especially in mammals), his
knowledge of musculature, and his pioneering use of silhoue ed
outlines to show the extent of soft tissues around the skeleton. Here is
the origin of the 'anatomically rigorous' movement in palaeoart.
However, Knight was a paradox. He understood well the link
between osteology and musculature, yet he gave dinosaurs small,
slender muscles that did not match their bones (dinosaurs actually
seem to have had enormous, more bird-like muscles), and frequently
drew dinosaurs freehand-style, again with what looks like poor
a ention to the proportions and nuances of the actual skeletons.

Allosaurus (left) and Diplodocus (right) by Charles Knight. Note the slender, lizard-like
thighs.
This paradox - this inherent contradiction - has remained throughout
the history of palaeoart. Yes, there are efforts to be as rigorous as
possible and to put in an enormous amount of unseen background
research on the detailed anatomy of those fossil animals being
reconstructed, but there are also quick and dirty ways of doing things
where research is minimal. A large number of popular books about
prehistoric animals use the work of individuals who do no research
whatsoever, creating their digitally reconstructed animals simply by
copying those depicted beforehand by other artists.

Paul's massive influence means that Mesozoic archosaurs-dinosaurs

and pterosaurs especially-are nowadays frequently depicted in high-
fidelity skeletal form before, or at the same time as, fleshed-out life
reconstructions appear. Outside of archosaurs, li le similar work is
so obvious, bar the several anatomist-artists (including Jay Ma ernes
and Adrie and Alfons Kennis) who have worked so hard to
reconstruct the faces and bodies of fossil hominids. Mauricio Antón is
now well known as an excellent and anatomically rigorous restorer of
fossil mammals and other vertebrates: his books and technical articles
are as much about the detailed science of anatomical reconstruction
as they are about evolution and palaeobiology.5,6,7,8

What needs to be made clear at this point, then, is that palaeoart of

the sort discussed and depicted in this book is firmly grounded in a
sceptical, rigorous, evidence-led effort to study and depict anatomy:
the approach promoted by Paul, Antón and the like. Several other
palaeoartists of the modern era-Jason Brougham, Mark Halle , Sco
Hartman, Bob Nicholls, Emily Willoughby and Mark Wi on come to
mind-are similarly part of the 'anatomically rigorous' movement.
Others are not, and it shows.

It should be noted that there are some disagreements at the level of

reconstructing skeletons and musculature, and that improvements
and tweaks are frequently being made. We mostly agree on the
positions of muscles, for example, but the sizes of some of the
muscles involved are variable in living animals and there is
sometimes no reliable way of determining their size in fossil animals.
This is well illustrated by Hutchinson et al.'s discussion of muscle
mass and body size in Tyrannosaurus rex9 where competing
possibilities made the same T. rex specimens look either svelte,
muscular or ridiculously muscular. The forelimb posture of bipedal
dinosaurs has been extensively revised in recent years as workers
have shown that palms faced inwards, not downwards,10,11,12 and
reassessment of dinosaurian tails have made it clear that Paul's
dinosaurs are typically not bulky enough in the tail region.13

Recent evidence suggests that dinosaurs were far more muscular in the tail region than
previously assumed. This reconstruction of Tyrannosaurus rex, produced by Sco
Hartman, has an appropriately chunky tail.
The problem with integument
The real complication in reconstructing fossil animals – the reason
we’re here, with a published book on the subject – is that there’s all
the soft stuff that goes on top of the musculoskeletal system.
Integument is the great unknown for many fossil animals; its
preservation is rare and infrequent, and even when it’s preserved, it
may be massively distorted or rearranged relative to its position in
life. This is important, since the shape and size of the integument can
radically change the appearance of the living animal relative to its
underlying musculature and skeleton. The skeletons of modern birds
– owls and parrots, for example – have long, slender neck skeletons,
but overlying skin and thick feather coverings obscure these entirely.
Extravagant head, wing and tail feathers present in some birds are
not reflected in the underlying osteology either, and the manes, ruffs,
thick furry coats and extensive amounts of skin linking the body with
the limbs in many modern mammals are, similarly, not suggested at
all by osteology.

If these observations extend across a wide diversity of living animals,

would they have done so for extinct ones as well? We have li le to go
on, but what we know suggests that, yes, integumentary coverings
may have effectively obscured much of the underlying anatomy that
we’ve worked so hard to reconstruct. Notably, dinosaurs found with
soft tissues (namely skin impressions and feathers) are flamboyant.
Feathered dinosaurs are not only covered in feathers (with feathering
extending from the middle of the snout all the way to the tip of the
tail and even down to the ankles or toes), they have especially long,
showy feathers growing off their arms and hands, the end parts of
their tails, and even (in cases) from their thighs, shins and feet. Fossil
mammals with body outlines and fur show a thick halo of tissues
surrounding the skeleton, meaning that the skeleton was deeply
submerged and effectively invisible in the live animal, as is typically
the case in modern species.

We are therefore presented with a huge diversity of ‘known

unknowns’ and ‘unknown unknowns’ – the gate is open for all
manner of bizarre possibilities as goes the life appearances of fossil
animals.

It is these speculative possibilities that John Conway and C. M.

Kosemen have explored in this book. Palaeontologists and
palaeoartists talk about these sorts of ideas all the time-about the
possibility that extinct animals were insanely flamboyant, that they
had super-sized genitalia, or that they were insulated from the cool or
even cold environments they sometimes inhabited by fat, thick skin,
or fuzzy coats-but this is the first time ideas of this sort have been
extensively discussed in print.

John Conway and C. M. Kosemen are two of the most exciting of

modern artists who are depicting extinct animals. Both combine
remarkable a ention to detail and technical accuracy with an
understanding of art and art history and a desire to move forwards-to
do something new, something innovative. Notably, both produce
work in the digital realm and the internet is the true home of their
many creations. A selection of John's outstanding works has recently
been showcased in the stunning Dinosaur Art volume14 and he is
rapidly becoming well known as the face of the future. His dinosaurs
and pterosaurs are not garish, flamboyantly pigmented, or shown in
a stereotypical landscape of palm trees and volcanoes, but
realistically muted and subtly hued, fi ing into backgrounds that
have been compared by some to impressionist paintings or Chinese
watercolours. C. M. Kosemen is best known for his remarkable
speculative and 'alternative' animals, some created as inhabitants of
speculative future evolutionary scenarios (see his All Tomorrows
project, available for free online, on human evolution) or of other
planets (see his Snaiad project). His speculative 'smart dinosaur'- later
labelled Avisapiens saurotheos and created as an antidote to the green,
scaly humanoids imagined by some palaeontologists-became an
internet sensation.

Above, Two aliens, a predatory Kahydron >and an armored, herbivorous Allotaur, from
C. M. Kosemen's Snaiad. Below, the intelligent Avisapiens, also by C. M. Kosemen.

To the future!
This book goes beyond some speculative possibilities about the life of
the past; it also indulges in a fun thought-experiment about a
hypothetical future. We humans have struggled to interpret the often
scrappy remains of fossil reptiles and other animals – it has taken us
decades, and a huge amount of luck and careful detective work, to
understand the true life appearance of small, predatory dinosaurs,
giant marine reptiles like plesiosaurs and mosasaurs, and other such
creatures. Think of the many mistakes we made along the way:
plesiosaurs with the insanely long neck misinterpreted as the tail,
Iguanodon imagined as a rhino-shaped, reptilian ‘pachyderm’, bird-
like predatory dinosaurs depicted more as big lizards or even as giant
turtles (this happened with Therizinosaurus), and so on.

Now consider what might happen were hypothetical non-human

scientists confronted with the remains of modern animals. If we
imagine that these curious and scientifically advanced creatures tried
to reconstruct modern birds, mammals and other animals, would
they end up with reconstructions that approach reality? It’s an
interesting thought experiment. Indeed, I tried it myself some years
ago, wondering just how modern cetaceans might be reconstructed if
hypothetical future scientists looked at them without knowing of
mammalian soft tissues. Many cetaceans have bizarre tall spines
growing upwards from their vertebrae; such distinctive structures as
the fa y melon on the top of the head and tail flukes are wholly
missing in fossils, and there’s no clear evidence for a thick covering of
blubber. Here, then, are my 1997 efforts to imagine a bo lenose
dolphin and Dall’s porpoise, pretending all the while that I’m a non-
human, non-mammalian palaeontologist from the future…
However, certain characteristics known as osteological correlates 15
can be used, with some reliability, to deduce soft tissue or integument
from skeletal details. Trunked mammals like elephants have unusual
bony recesses and other structures around their enlarged bony nasal
openings that help anchor the musculature associated with a
proboscis, and also possess a specially enlarged braincase opening for
one of the nerves that supplies the trunk. The melons and spermaceti
organs of toothed whales are housed in concave basins on the skull
roof, and the presence of horizontal tail flukes in cetaceans are hinted
at by the shapes of the vertebrae at the tail’s end. However, these are
things that we’ve discovered with the benefit of hindsight, or have
investigated specifically because we knew about the soft tissue
structures in the first place. If we truly imagine that modern animals
are represented in a future fossil record by imperfect remains, just
like ancient fossils in the real, modern-day world, things could
indeed be very different.

Science and speculation are happy bedfellows, so long as we remain

grounded in our speculations, and so long as we state the core
evidence we have in the first place. When it comes to the
reconstruction of fossil animals, there will always be a great many
aspects of anatomy, behaviour and lifestyle that will remain
unknown, and for which a modicum of reasonable speculation will
be allowed, and even necessary. Enjoy the ride.
All Yesterdays
What images come to mind when you hear the word "dinosaur"?
Perhaps you picture enormous lumbering animals, lurking in the
swamps of old; or maybe you imagine lithe predatory dinosaurs
sprinting on lean muscular legs, perfect killing machines with
mouths agape in perpetual screams. In recent years, these views have
become opposing stereotypes, pu ing prehistoric animals in
straightjackets of appearance and behaviour.

We wanted a way out – a way that would show dinosaurs (and the
other extinct animals of the Mesozoic Era) as plausible, real animals
that engage in such complicated behaviours such as play, courtship,
and the expression of curiosity.

While it is reasonable to think that the predominant purpose of

palaeontological art is to display the latest knowledge about extinct
animals, it can also serve as an arena for the proposal of new
hypotheses, rather than the repetitive drawing of proven theories. In
the history of palaeontology, out-of-the-norm images have been
crucial in popularizing new ideas about the appearances and
behaviours of extinct animals.

While we are well aware that some of our reconstructions will

probably be falsified, some may actually "hit the spot," or may even
look modest when compared to new fossil discoveries. Only time will
tell!
Carnotaurus and Other Arm-Waggling
Abelisaurs
Perhaps owing to its "bull-like" horns, Carnotaurus has become one of
the more popular predatory dinosaurs in public knowledge. A
member of an unique lineage of mostly Southern Hemisphere
predators known as abelisaurids, Carnotaurus had a short, deep skull,
heavy-set limb bones and very short arms. Although Carnotaurus is
known from reasonably complete remains, almost every feature of
this animal's body is surrounded by a cloud of theories and debates.
Visible yet frustratingly unknown, Carnotaurus is the perfect
palaeontological enigma.

This reconstruction was conceived as we tackled of the more

interesting anatomical details of Carnotaurus, namely, its comically
stunted hands. Although short dinosaur arms immediately remind
one of Tyrannosaurus, Carnotaurus and other abelisaurids
independently evolved a wholly different and even stranger set of
arms, which still bore a full set of four fingers. The upper arm bones
(or humeri) were long and straight while the lower arm bones were
bizarrely short. The head of the humerus was rounded and ball-like,
a feature indicating that substantial motion was possible at the
shoulder joint. It seems that Carnotaurus and kin could stick their
arms out sideways in a manner completely unlike that of other large
predatory dinosaurs.

What was the reason for this arrangement, and why was it
maintained across abelisaurids rather than fading away? Perhaps the
baby arms still had a function, albeit a social one. Palaeontologists
Phil Senter and J. M. Parrish suggested exactly this, proposing that
Carnotaurus and its relatives most likely waved their arms and hands
around when displaying to mates or rivals.17 We have to note that we
came up this idea independently of that study, and only learnt about
its existence after creating the artwork you see here. It's difficult to
think of a be er role for such weird structures.

Here, you can see a bull Carnotaurus and a related form,

Majungasaurus, in full display, flashing brightly-colored arms and
facial wa les to potential mates, or rivals. Seen head on, Carnotaurus
looks more like a science-fiction creature rather than a dinosaur, yet it
must be remembered that the sideways-facing depictions we are
accustomed to are artifacts of scientific illustration, laid out for
maximum visibility and clarity. Real-life animals would have been
more complex, more three-dimensional.
Elasmosaurus in a Neck-Swinging Contest
Even if, one day, we had access to perfectly preserved fossils, a vital
aspect of animal life would still elude our grasp. Behaviour is almost
entirely lost in the fossil record. Imagine the richness and strange
wonder of animal life today. The eerie, ululating songs of whales, the
elaborate middens of bowerbirds and the surreal spectacle of a
peacock’s display could never be deduced from inanimate remains.

Likewise, some of the most spectacular sights of the past will never
be seen, or even guessed. In this painting, we have imagined one such
piece of behaviour, in this case as applied to elasmosaurs, long
necked marine reptiles that lived during the Cretaceous period.
Although they feature in many books about dinosaurs, elasmosaurs
were not members of the dinosaur family. Instead, they belonged to a
distinct lineage of marine reptiles known as plesiosaurs.

This particular group of male elasmosaurs are out in the open sea,
trying to see who is the toughest by lunging up from the depths and
waving their necks. While old depictions mistakenly portrayed
elasmosaurs as holding their necks swan-like above the water surface,
these animals actually had very dense bones and heavy necks that
could probably not be lifted out of the water when the animal was in
its standard, horizontal pose.16 Accordingly, the stunt seen here
would have been very difficult and energy-demanding, and could not
be sustained for more than a few seconds. What be er way is there
for males to prove their strength than through rigorous, energy-
intensive bouts of a ridiculously pointless activity? No record of such
behaviour could possibly exist – it is wholly and unashamedly
speculative – and yet things equally spectacular must have happened
throughout the history of life.
A Giant Centipede Snatches an Anurognathid
Pterosaurs are everybody's favorite prehistoric flyers. Anurognathus
was a member of an extraordinary group of pterosaurs known as
anurognathids. Members of this group were characterized by
extremely small size, short, broad wings and wide, frog-like mouths.
They are further unusual in possessing short tails, an independently-
evolved feature elsewhere seen only in the larger, more "advanced"
pterosaurs called pterodactyloids. Judging from their wing and skull
shape, anurognathids are thought to have lived like today's insect-
eating bats.

Small-bodied animals like bats and anurognathids are rarely

preserved in the fossil record. Although only several anurognathid
specimens are known to science, there may have been hundreds, even
thousands, of different anurognathid species living in the lost forests,
caves and islands of the past. These animals must have lived in a
world of danger, where they were vulnerable to predation, not just
from dinosaurs, birds and other pterosaurs, but also from smaller
animals like mammals, insects, spiders and centipedes.

Our illustration depicts the death of an anurognathid at the

formidable, jaw-like maxillipeds of a large scolopendrid centipede.
Centipedes have a poor fossil record, but scolopendrids are known
from the Cretaceous: in fact, some Cretaceous centipedes are virtually
indistinguishable from modern ones. It is wholly plausible that large
scolopendrids were snatching small, flying animals during Mesozoic
times, just as they do today.18
Allosaurus fragilis and Camptosaurus dispar
Palaeontology aims to obtain a clear, natural view of the past, but
gratuitous acts of predation and vicious monsters are an undeniable
factor in a racting people to the study of dinosaurs. The tradition of
palaeoart is full of such epic ba les, almost as canonical as the clash
of heroes in classical mythology. No children's book is complete
without scenes of Tyrannosaurus a acking Triceratops, matched in
miniature by a Velociraptor locked in mortal combat with a
Protoceratops, and so on.

While predation is indeed a vital (and violent) fact of nature, not all
predator-prey encounters end in cinematic bloody struggles. More
often than not, hunters give up on chasing their quarry. Predators
regularly ignore animals that won't be worth the energy to pursue
them, and herbivores may cautiously approach meat-eaters while
seeking common resources such as water. Curiosity, fear,
intimidation and exhaustion make predator-prey relationships far
more complicated than we typically picture them to be.

In this scene, set in the Late Jurassic, a herbivorous Camptosaurus is

seen approaching a resting Allosaurus in what appears to be a curious
social gesture. While Allosaurus was certainly a regular predator of
Camptosaurus, this encounter seems to be a peaceful exception to the
norm. In today's ecosystems, predatory big cats and herbivores have
also been observed interacting in similarly non-violent ways.
Tenontosaurus tille i Takes a Stroll
Like mythical figures cast into tragic roles, some dinosaurs suffer
terrible fates in palaeoart, experiencing the same deaths over and
over again. Tenontosaurus - a horse-sized member of the same group
as Iguanodon that lived in North America during the Early
Cretaceous-was one such dinosaur. Because its remains were found
in association with the meat-eating Deinonychus, scientists have
argued that Tenontosaurus was a frequent prey of the vicious, bird-like
predators.19

While this assumption is possibly correct, it has become a cliché in

fact, it is almost impossible to find a reconstruction of Tenontosaurus
where it isn't being viciously torn apart by a pack of Deinonychus.
Relegated to the role of "stock fodder," many unique features of
Tenontosaurus, such as its unusually long tail and interesting position
within the iguanodontian family tree, have been mostly overlooked
by the general audience.

In real-world ecosystems, predators are far less common than their

quarry. It is thus absolutely likely that Tenontosaurus spent most of its
time feeding and resting, not fending off Deinonychus a acks. To
illustrate this point, we depict Tenontosaurus happily walking along,
without a single Deinonychus in sight. This surely happened on a
regular basis.
Hypsilophodon foxii Eating a Millipede
Much like wildlife art, prehistoric art has unconsciously cast roles for
animals and their behaviour. Predators are always on the hunt, giants
are always seen in majestic repose, while small herbivores, depicted
as meek and innocent, are either shown grazing or in hurried flight
from some terrible hunter.

With small, herbivorous dinosaurs, this type-casting has a risk of

falling short of reality. To begin with, herbivorous animals are not
"innocent" of occasional predatory acts. Modern herbivores eat, or
will at least try to eat, a wide variety of things. Deer and sheep have
been observed chewing the heads and legs off seabirds,20, squirrels
often eat bird hatchlings for an extra helping of protein, and it turns
out that even ca le will eat bird's eggs and nestlings if they find
them.21 Moreover, one can only draw general connections between
anatomy and dietary adaptations in fossil animals. There is no doubt
many of these are correct assumptions but, then again, we may be
overlooking some fascinating possibilities.

Here, we see Hypsilophodon, generally known as the ultimate small

dinosaurian herbivore. This particular Hypsilophodon is unaware of
future speculations on its remains, and is happily supplementing its
plant diet with small animals. However, this snack will give it a nasty
surprise-millipedes often cover their bodies with foul-tasting
chemicals.
Citipati osmolskae and Stegosaurus stenops :
Giant Penises and Violent, Confused Sexuality
Reproduction is one of the most important, if not the most important
driving force in evolution. Sexual display and sexual a raction have
resulted in the peacock’s dazzling tail feathers, the musical songs of
birds, and the various horns, crests and frills that adorn animals of
every size and shape. Socio-sexual display behaviours may even have
contributed to the evolution of language and human intelligence.

With reproductive acts playing such a vital role in the evolution of

life, it is surprising that the issue has not received more a ention in
palaeoart. If we assume that the sex habits of Mesozoic dinosaurs
were similar to those of their modern descendants (the birds), then a
dazzling, even bizarre, range of possibilities reveal themselves.
Among modern birds, ducks lead notoriously rough sex lives,
involving aggressive mobbing and gang-rape. In some duck lineages,
female vaginal canals and male penises have entered a bizarre sexual
“arms race”,22 one of the results being that the males of some species
have penises that can sometimes be as long as the head, neck and
body combined.23,24 We have imagined a similar scenario in this
particular reconstruction of Citipati, an oviraptorosaurian
maniraptoran theropod from the Late Cretaceous. This Citipati has
expired after a particularly rough season of mating, but at least its
genes have been safely passed on to the next generation.

Another possibility we have considered is interspecies mating. When

sexually aroused, excited or unable to find available members of their
own kind, animals mate with members of other species with
surprising regularity. Incidents of this sort are probably more
common than generally realised, and there is evidence from the
modern world that they occur increasingly during times of
environmental stress or as populations become reduced or brought
together due to changing conditions. When the species concerned are
closely related, hybrid babies can be the result: numerous such cases
are known from the modern world. However, matings between
distantly-related species also occur in the wild. These seem to serve
no function other than to relieve the frustration or boredom of at least
one of the participants. As unse ling as they may seem, such acts
may even be considered to be part of the animal's play behavior. In
one especially celebrated recent case, an apparently frustrated
Antarctic fur seal (Arctocephalus gazella) copulated with a King
penguin (Aptenodytes patagonicus).25 It is well known that modern
elephants are prone to a sort of seasonal sexual madness when they
go through a phase of heightened sexual aggression termed musth.
While in musth, elephants have been observed trying to forcefully
mate with members of different species, such as rhinos.
We combined ideas about interspecies mating events with both the
possibility of oversized sexual organs and of a seasonal 'sexual
madness'. The result: a bull Stegosaurus trying to mount an innocent
Haplocanthosaurus. In order to mate with females bearing a phalanx of
dangerous spines and armored plates, we imagined male stegosaurs
to have developed some of the largest and most frighteningly
dextrous penises of the dinosaur world.

Stegosaurus trying to mount an innocent Haplocanthosaurus.

Stegosaurus stenops skeletal
Glorious Mud— Camarasaurus grandis
Of all the kinds of behaviour that animals can engage in, it's probably
play that been depicted the most rarely. Plenty of pieces of art show
dinosaurs running, jumping, a acking or defending themselves from
other dinosaurs, and some even show dinosaurs mating and
defecating, but almost none show them playing. Celebrated artist
Luis Rey might be unique, then, in having published a scene where
dromaeosaurids and troodontids are shown playfully sliding down a
snowy slope in a Late Cretaceous winter,26 a speculation inspired by
behaviour seen in living magpies and crows. Other than Rey’s work,
no artist seems to have thought of dinosaurs in play. This lack of
interest is no doubt maintained through the idea that only “smart”
animals such as birds and mammals engage in play behaviour.
Interestingly enough, it is now known that play behaviour is not
limited to birds and mammals. Monitor lizards, turtles, crocodiles
and even fish and cephalopods have been reported to engage in
behaviours that do not seem to serve any other purpose than simply
having fun.27,28 If all these animals could play, we are certain that
Mesozoic dinosaurs could, too. Here a sub-adult Camarasaurus is
shown enjoying the soothing feeling and antiparasitic properties of a
good roll in the mud.

Camarasaurus was a large sauropod dinosaur from the Late Jurassic of

North America. Well-known for its boxy skull and stout proportions,
it is considered by some experts to be among the ugliest of all
sauropods.
Sleepy Stan— Tyrannosaurus rex
If popular depictions are to be believed, Tyrannosaurus spent most of
its life charging at hapless victims while roaring at the top of its
lungs. This image, immortalized by movie and comic book
depictions, is false on both fronts. To begin with, predators almost
never roar or scream while a acking. Stealth is vital in nature. A
hunter like Tyrannosaurus must have waited silently, and kept as
quiet as possible as it hunted. Even the slightest noise could have
scared its prey away. The only things its victims would have heard
would be the crashing of the vegetation as the giant predator charged
towards them.

Secondly, as we saw before with Allosaurus, hunts take up only a

minor part of a predator's lifetime. Most hunting animals spend long
days resting, either in order to conserve energy, or while digesting
the food acquired from a fresh kill. Like most warm-blooded modern
predators, the fearsome T. rex may have spent most of its time asleep.

Here, we portrayed a large specimen of Tyrannosaurus with an

unusually large head, also known as "Stan." In this picture, Stan has
just finished a big meal and is sleeping soundly as he digests his fill.
A Tyrannosaurus like Stan was probably capable of eating tonnes of
flesh in a single hunt, and would have eaten relatively infrequently. It
could have taken Stan several days to sleep off the exhaustion and
torpor of a meal.

The sleeping poses of tyrannosaurs have also been an interesting area

of speculation. Resting poses have been depicted for tyrannosaurs
before: Lawrence Lambe imagined tyrannosaurs lying flat on their
bellies, partly due to their boot-shaped pubic bones.29 Combine this
with the idea that giant animals cannot lie on their sides due to their
excessive weight and an artistic meme was born whereby
tyrannosaurs were never shown sleeping on their sides. As
demonstrated by elephants, however, even tyrannosaur-sized
animals can and do recline on their sides and sleep deeply.30 How,
when and where other big dinosaurs slept remains a fascinating area
of speculation.
Mesozoic Goats— Protoceratops andrewsi Do
What They Aren't Best At
Reconstructing the behaviour of extinct animals is challenging, even
if we assume a good correlation between anatomy and behaviour.
Unfortunately, this correlation does not always exist in the real
world. Elephants are excellent at swimming, crocodiles and alligators
sometimes eat fruit and leaves, juvenile iguanas sometimes jump up
at the moon at night, and goats in some areas often climb trees in
order to browse. Animals do what they do, not necessarily because it
is what they are good at, or even because their anatomy is suited to it,
but simply because they can. As a result, unexpected behaviours are
commonplace.
Here, the famous ceratopsian Protoceratops is engaged in something it
has no obvious adaptations or reason to do: climbing trees.
Protoceratops was a boar-sized herbivore that lived during the Late
Cretaceous in Mongolia. It is related to the larger horned dinosaurs of
North America such as Triceratops, and is thought to have retained
the general appearance of their common ancestor. Hundreds of
Protoceratops skeletons are known, many preserved in spectacular and
lifelike poses. One specimen is preserved locked in combat with a
Velociraptor; others died in poses showing that they were struggling
to dig out of the sand that buried them.
Majungasaurus Makes Like a Log
Crypsis is a theme rarely explored in palaeontological art. Many
present-day animals develop complex coloration schemes to disguise
themselves from predators, prey, or both. Other animals match their
appearance and behavior to different, usually poisonous species and
mimic them for protection. While crypsis and mimicry are seen most
often in smaller creatures, large-bodied animals also exhibit strange
and confusing body pa erns for disguise. A tiger’s orange and black
stripes might look dazzling in a zoo but, in its natural environment,
they render it nearly invisible in the tall grass. Likewise, crocodiles
blend in among swamps and riverbanks with their outline-breaking
scutes and mo led coloration. At least some dinosaurs must have had
similar camouflage schemes.

Can you see the Majungasaurus in this picture? Majungasaurus was a

Cretaceous abelisaurid theropod from Madagascar, a strangely-
proportioned relative of the Carnotaurus we saw earlier. Unless there
is something very wrong with the fossils we have, Majungasaurus had
an extremely long body, combined with very short legs and relictual
arms, giving it the proportions of a bipedal, dinosaurian daschund.
As you might have guessed, it is not clear how this predatory
dinosaur moved about and hunted with such an atypical body plan.
We imagined that camouflage, made possible by matching its pebbly
scales to the texture of rocks or trees, made things easier for the
strange old Majungasaurus.
A Plesiosaur Makes like Some Coral
Camouflage is not limited to land animals. Rivers, lakes and seas also
contain many risks, and many opportunities for hiding. The oceans
especially teem with dazzling examples of crypsis; sea dragons that
mimic strands and stalks of kelp, wobbegong sharks that lie down
like mats of sand and seaweed, poisonous stonefish that are
dangerously invisible among stones, and so on.

We imagined what sort of animals might have camouflaged

themselves in prehistoric seas, and turned to long-necked marine
reptiles known as plesiosaurs , as a possible example. Plesiosaurs
were not dinosaurs, and instead belonged to a major group of marine
reptiles called sauropterygians. The most successful, most diverse
and most long-lived sauropterygians, plesiosaurs included the
previously-mentioned elasmosaurs, the big-headed, macropredatory
pliosaurs as well as a number of groups that are intermediate in
shape and proportions.

We pictured this particular plesiosaur as a sit-and-wait hunter, lying

in shallow coral reefs and waiting for suitably sized prey to swim by.
A near total absence of information on plesiosaur skin morphology
means that we have no firm ideas about skin texture in these animals.
It is assumed that most had a smooth skin for hydrodynamic
efficiency, but we imagined this particular plesiosaur as a well-
camouflaged hunter with an exceptional sit-and-wait strategy.

As an air-breather, the dive time of our camouflaged lurker would be

limited, but could theoretically still be long enough to allow fruitful
hunting in areas of rich prey density. The dorsoventrally compressed
body shape present in at least some plesiosaurs31 renders it possible
that they could lie on the seafloor; intriguingly, the shallow-bodied
Jurassic form Tatenectes also has particularly dense, heavy bone
located on its underside and near its midline, suggested that it was
using bone as a form of ballast to stay near (or on?) the seafloor. We
speculate that some bo om-hugging plesiosaurs may have used their
long necks for suction feeding: they would open their mouths and
lunge up when a small fish or marine reptile wandered by. The rush
of water filling the animal's long throat cavity would create a brief
vacuum effect, helping it capture its victim.
Ouranosaurus : I'm Not Fat, it's the Spines!
Nothing quite says "prehistoric" like an animal with a skin sail
running down its back. This is possibly due to the "reptilian" nature
of such baroque accessories: the only animals with sails on their
backs today are cold-blooded, "exotic" lizards such as basilisks.
Whatever its cause, this trend is so entrenched in popular culture that
Dimetrodon, a sail-backed animal that lived millions of years before
dinosaurs and was more closely related to mammals, is frequently
lumped in with the ruling reptiles in children's books and toy sets.
Even within dinosaurs, the popular desire to have ridge-backed
monsters manifests itself with depictions of exaggerated skin sails in
species with extended vertebral spines, such as the herbivorous
ornithopod Ouranosaurus.
Tall vertebral spines on dinosaurs are usually interpreted as having
supported "sails", with li le flesh covering the bones. However, it is
possible that these spines supported other tissues, such as fat
deposits or humps. This is exactly the case in large herbivores today
such as bison, camels and rhinoceroses. Indeed, palaeontologist Jack
Bailey published some reconstructions in 1997 in which he made
Ouranosaurus and Spinosaurus look more 'hump-backed' than 'sail-
backed'.32 Perhaps, however, this approach is wrong and tall-spined
reptiles like chameleons, with narrow ridges that are neither sails nor
a humps, provide be er models for dinosaurs.33 Nevertheless, this
area remains understudied and we decided to reconstruct one of the
more popular "sail-backed" dinosaurs with an armored hump on its
back. Behold Ouranosaurus in its hump-backed glory.

Ouranosaurus was a large iguanodontian ornithopod that inhabited

northern Africa during Early Cretaceous times. Several other
distantly related dinosaurs from the same location also have
elongated vertebral spines, including the enormous theropod
Spinosaurus and the sauropod Rebbachisaurus. So perhaps, instead of
just Ouranosaurus, the region was inhabited by an entire cast of
unusual, hump-backed dinosaurs in real life. The reason for such
unusually synchronized convergent evolution, and indeed the very
nature of these spines in life, remains unknown. Perhaps climactic
factors or sexual selection could have played a role. Display is
certainly a likely purpose-and certainly fits with the general idea that
dinosaurs were flamboyant, social animals.
Parasaurolophus walkeri the Porker and
Lambeosaurus magnicristatus the Flapper
Since the old view of dinosaurs as lumbering and shapeless piles of
stocky lard was discarded, palaeontological artists have been keen to
portray most dinosaurs as slim, sleek animals where every contour of
the limbs, every muscle and even every bone can be clearly
discerned. No living mammal, reptile or bird has such a “visible”
anatomy. Living animals are replete with skin flaps, fat and saggy
bits that obscure the exact lines of their bones and musculature.
 Parasaurolophus skeletal

The famous “duck-billed dinosaurs” known as hadrosaurs have

provided us with some of the best demonstrations of the fact that
dinosaurs were not walking anatomical diagrams. Remarkable
mummified hadrosaur specimens provide us with an unprecedented
amount of information on hadrosaur soft tissue anatomy, and one of
the most interesting features of these fossils is a series of vertical
“shoulder folds” that cover the upper arm and shoulder region. Greg
Paul has argued that these were evidently present in life, and must
always be depicted in life reconstructions of these animals.34 Here,
however, we explore the possibility that these are actually artifacts of
desiccation, and that they supported a heavy padding of fat and
muscles in real life. Behold our fat Parasaurolophus and the solidly-
built Lambeosaurus with a flap-like throat pouch.

Parasaurolophus and Lambeosaurus are familiar duckbilled dinosaurs,

recognisable by their spectacular head crests. Duckbilled dinosaurs,
or hadrosaurs, were the dominant plant-eaters of the Late Cretaceous,
and had evolved one of the most advanced chewing mechanisms
seen in vertebrates. Their crests were hollow and almost certainly
would have been used to create loud honking sounds for
communication.

Lambeosaurus magnicristatus
Mountain of Feathers— Therizinosaurus
Some dinosaurs suffer from looking too exciting de-fleshed, leading
artists to depict them in exciting and garish ways. If one group of
dinosaurs could be singled out for their strangeness, the mystifying,
long-clawed therizinosaurs would win hands (or claws) down.

Therizinosaurs are a such a bizarre group of dinosaurs that, for a long

time, it was not possible to discern what they were, let alone what
they looked like. Originally they were known only from a set of giant,
mystifying claws, the longest of which were more than 70 centimeters
long. These claws prompted a whole range of theories and
hypothetical identities. Early ideas were that therizinosaurs might be
gigantic, turtle-like beasts or vast, vicious predators which slashed
open the bellies of sauropods and other large dinosaurian prey. Be er
remains led to suggestions that therizinosaurs were plant eaters:
perhaps late-surviving prosauropods or relatives of early
ornithischians, or even a completely unique lineage of dinosaurs
distinct from any other group. With newer discoveries and the rise of
cladistic methodology, it became clear that they were, in fact,
aberrant herbivorous theropods, reasonably closely related to birds.

The skeleton of Nothronychus mckinleyi shows the strange anatomy of therizinosaurs

Even after scientists established what they were, depictions of

therizinosaurs maintained a strong claw-based fetish. Therizinosaurus
– shown here in our life reconstruction – was one of the later, more
advanced forms. It was a large, elephant-sized animal with
tremendously wide hips and (presumably) enormous guts. To be fair,
it is difficult to ignore the sheer weirdness of these majestic animals;
in real-life, however, exciting skeletons, claws and bellies are all
covered up in a great big mounds of feathers, fur, or fat. Silhoue es
are the most distinctly recognizable visual a ributes of large animals
today. As a result, our Therizinosaurus are not brandishing their
meter-long claws in the viewer’s face. Our opinion on palaeoart is
that subtle references and hints of anatomical features are more
realistic, more in line with what we see in living animals. Portrayed
with style and in an appropriate se ing, they can leave a more
distinct effect on the viewer.
Heterodontosaurus tucki the Goes the Whole
(Hedge)Hog
Small ornithischians, the quintessential "harmless herbivores" of the
prehistoric world, have had one of the greatest image changes of the
dinosaur renaissance in the recent years. Traditionally, these animals
were usually portrayed as smaller versions of their larger relatives.
Unlike the meat-eating theropods, whose feathery integument is
becoming more-or-less common knowledge thanks to dozens of new
well-preserved fossil discoveries, artists and scientists were reluctant
to dress small ornithischians in feathers or "dinofuzz."

When illustrated, these generic, naked herbivores were mostly shown

running, either away from a more interesting predator, or simply
scampering across the landscape with no purpose or aim. However,
recent discoveries have shown that there is far more to small
ornithischians than previously assumed. To begin with, they are no
longer naked: dermal quills are now thought to be present in almost
all small ornithischians after the discovery of the Chinese
heterodontosaurid Tianyulong (preserved with a covering of spike-
like body hairs35) and of the early ceratopsian Psi acosaurus (one
specimen is preserved with bristle-like structures on its tail36).
Another exciting discovery, this time of an ornithopod called
Oryctodromeus, has revealed that at least some small ornithischians
had burrowing habits, spending at least part of their time sheltering
together in dens.37 It seems that these dens were inhabited by family
units. Taken altogether, these discoveries paint a very different
picture of small ornithischians. No longer are they defenseless
bipedal ‘lizards’, but an unique group of animals with their own
extraordinary adaptations and social behavior.

This picture shows a family group of Heterodontosaurus near the entry

of their collective burrow. Taking the latest discoveries into account,
we reconstructed them with a full set of porcupine-like quills for
defence. Heterodontosaurus was a small, basal ornithischian from the
Early Jurassic of South Africa. Its name ("differing tooth lizard")
refers to its unusual dentition. Heterodontosaurus had the beak and
chewing teeth common to most ornithischian dinosaurs, but it also
had large "canines" and two other types of teeth that helped it chew
its food efficiently. Scientists still aren't sure if Heterodontosaurus was
purely herbivorous, or if it supplanted its diet with small animals as
well.
Let's get fluffy: Leaellynasaura amicagraphica
As seen before with Hypsilophodon, many small plant-eating
dinosaurs will have their popular images greatly revised thanks to
new discoveries of social behaviour and integument. Instead of
looking like two-legged iguanas, these animals will have to be re-
imagined with the extra possibilities offered by furry bodies and
communal living habits.

Here is one more small ornithischian re-interpretation, this time

featuring Leaellynasaura. This dinosaur was discovered in Australia,
where it lived during Early Cretaceous times, about one hundred and
ten million years ago. At that time, Australia was located close to the
Earth's geographic south pole. Although its climate was possibly not
as cold as today's Antarctic, the axial tilt of our planet meant that
Cretaceous Australia did not receive direct sunlight for long periods
of time, and almost certainly experienced sub-zero temperatures.

Aside from being a polar dinosaur, Leaellynasaura was also

extraordinary for its immensely long tail, which was almost three
times as long as its body. Nobody knows why Leaellynasaura had
such a long tail, or what it used it for. Theories range from the tail
being used as a climbing aid, a sexual and social display feature, or a
long, shaggy “scarf” that the dinosaur wrapped around itself as
protection from the cold. It has even been suggested that the tail
aided the animal in swimming!

Drawing on these facts, we reconstructed Leaellynasaura as a rotund

furball that scurried peacefully about in the polar forests of
Australia's past. We imagined its long tail as a thin, signalling
"flagpole" that helped it identify and keep close to members of its
herd. No doubt some people will find this reconstruction
preposterous, and perhaps they will be right. However, we felt that
not enough dinosaurs were reconstructed as "cute" beasts, whereas in
nature, polar animals can look quite pulchritudinous under layers of
fat, muscle, fur and other insulation.
Microraptor gui
Almost twenty years of fossil discoveries have firmly established that
certain small, meat-eating dinosaurs were covered in feathers, to such
a degree that even popular science books routinely depict these
animals with some sort of integument. Yet many palaeoartists, it
seems, still have difficulty picturing such small, feathered theropods
as bird-like creatures. Instead of picturing the coherent, well-rounded
forms that are found in nature, most artists over-focus on certain
features or details of preservation found in a single fossil, and repeat
and exaggerate them to picture "li le monsters" that look more like
film creatures than real animals. When examined in detail, this
typecasting can tell us a lot about trends in palaeoart.

The small theropod Microraptor is a case to the point. This small,

possibly flying relative of Velociraptor and Archaeopteryx was
described in 2003 as a "four-winged dinosaur"38 amid much media
hype. Since then, almost every reconstruction of this animal has
depicted it as a strange, dragon-like feathered glider with a reptilian
face. In almost every picture, Microraptor is virtually always shown
spreading its arms and legs, all lined with feathers that stick out as
visibly as possible, as if to prove to the viewer that it really had “four
wings.” Such illustrations might be educational, but they also help
popularize an image of the animal that was not real. It’s as if artists
are unable to conceive of dinosaurs that don’t look abnormal and
“alien” in some way.

When illustrating Microraptor, we wanted to “filter out” all popular

repetitions and approach it from scratch. The animal was indeed
remarkable for having wing-feather-like structures on its rear legs,
but we know from observations of living animals that flying species
rarely have their wings fully visible when resting. Our result looks far
more naturalistic, and far less like the spread-eagled fantasy creature
that popular audiences have come to associate with Microraptor.

On a side note, we gave our Microraptor s a nest in order to add depth

to their story. The nesting behaviour of dinosaurs is a complex issue
that deserves further discussion and speculation: it seems likely that
this varied immensely, just as it does in modern birds.
Triceratops horridus
Even the most familiar of dinosaurs may hold great surprises in their
life appearance. It seems that every time the soft tissue of a dinosaur
is discovered, our views of that animal, and usually all of its relatives
as well, are changed drastically. Such revelations show how artificial
our images of even the most well-known dinosaurs can be. What we
are drawing all the time may not be the "real" animals themselves,
but artifacts of an artistic tradition.
An example is Triceratops, arguably one of the most recognizable of
all dinosaurs. This animal has always been seen as a sort of
dinosaurian rhinoceros, with its iconic head frill, three horns and the
giant, plant-cropping beak. In recent years, however, scientists have
made several discoveries that may drastically revise our image of this
animal. Fossil skin impressions (mentioned online in 2010, but
unpublished at the time of writing) apparently reveal that Triceratops
had nipple-like protuberances located within the centres of the scales
that cover its back and tail: it is inferred that these structures
anchored quill-like structures of some sort. Moreover, other fossils
show that an earlier member of the horned dinosaur family,
Psi acosaurus, had a fringe of long, spine-like filaments lining the
upper surface of its tail.39 Since Psi acosaurus was close to the
ancestry of all large-bodied, horned dinosaurs, it would not be
unreasonable to suggest that all ceratopsians possessed some sort of
spiny integument.

Following this line of deduction, we produced this unfamiliar

depiction of the very familiar Triceratops. What if the unusual dermal
nipples of Triceratops were the bases of giant protective spines? While
we are not suggesting that this was the definitive look of this animal,
it wouldn't surprise us if Triceratops, or other, even less-expected
dinosaurs bore a covering of spiky hairs.
Everything Old is New Again:
Opisthocoelicaudia
The oldest dinosaur stereotype is that of the massively dumb, long-
necked sauropod, dragging itself slowly out of a fetid swamp. Once
pictured as too big to support themselves on land, sauropod
dinosaurs have long since been lifted out of the marshlands. Thanks
to new fossil discoveries and unbiased considerations of the
anatomical and palaeoenvironmental evidence,40 we now know that
most sauropods lived on land, moved about on erect legs and did not
drag their tails on the ground.41

Numerous recent discoveries have also revealed that sauropods were

far more diverse and complicated than originally assumed. There
were tall, long-limbed forms such as Brachiosaurus and Giraffatitan,
long and lean, whiplash-tailed form such as Diplodocus, and strangely
proportioned forms such as the short-necked Brachytrachelopan42 and
the weird Isisaurus43 with its wide, deep neck and unusually
proportioned limbs. Isolated island habitats led to the evolution of
dwarf, cow-sized sauropods such as Europasaurus44.

Within this diversity, how sure are we that sauropods were all
landlubbers? As we can see from hippopotamuses and certain Asian
rhinoceros species today, large herbivores don't need to be cold-
blooded dimwits in order to pursue partially aquatic lifestyles.
Perhaps the same was also true for members of some sauropod
lineages.

If we could bet on the possibility of a certain sauropod being at least

partially aquatic, our money would be on Opisthocoelicaudia, a
titanosaur from the Late Cretaceous of Mongolia. Known from a
famous fossil that preserved a more-or-less complete body and limbs
but, frustratingly, has no head or neck preserved, it was surprisingly
rotund animal, with short legs and a massive, round body. Its tail
vertebra fit together via flexible ball-and-socket joints, and its tail
preserves evidence of massive ligamentous and muscular
a achments. It is not difficult to imagine these adaptations as
modifications for a mostly water-based mode of existence. While
today's Mongolia doesn't sound like the best place to find water-
dwelling dinosaurs, the area was a lush, fertile zone crossed by many
rivers during the time Opisthocoelicaudia lived. At the time of writing,
no technical study has properly addressed the possibility of aquatic
life for Opisthocoelicaudia, but its proportions certainly recall those of
fossil mammals suggested to be amphibious.45 Those who know their
palaeoart will of course recognise our illustration as a homage to part
of the Zallinger mural.
All Todays
Although the accumulation of discoveries and fossil insights have
given us a more-or-less clear view of some extinct animals, many are
still reconstructed with a rule-of-the thumb methodology. As we saw
in the previous section, this modus operandi does not take soft tissues,
integument, colour and behavioural elements into account. More
recently, palaeontological artists have began to notice these
shortcomings and have begun to include more lifelike (if slightly
speculative) elements in their reconstructive work.

In this section, "All Todays", we took the opposite approach and tried
to highlight what some reconstructions might be missing out by
"reconstructing" modern animals from their skeletal diagrams. We
took the position of speculative observers with some knowledge of
how vertebrate bones and muscles work, but with no information on
the lifestyles or the soft tissue integument of their subjects. We
applied this approach to a number of modern-day animals such as
birds, ungulates, carnivores, whales and reptiles, with extraordinary
and sometimes hilarious results.
Shrink-Wrapping: How Future Artists Will
Show that They Know the Anatomy
As we have already discussed, the modern, sleek versions of
dinosaurs have become a cliché. Dinosaurs have been stripped of
extraneous soft tissue, with every muscle and bone ridge carefully
portrayed as visible in life. It has become fashionable to reconstruct
theropods with every ridge and fossa on their skulls echoed by lines
of scales, with the joints between skull bones given maximum
visibility.

Perhaps palaeontological artists of the future will give modern

animals the same treatment. They will want to flaunt their knowledge
of mammalian anatomy by faithfully portraying every limb bone,
every finger, every indentation in the skulls. Some will be bolder than
the rest, and will decorate their creations with a light sprinkling of
hairs, or apply garish color schemes. To be honest, it won’t be easy to
blame them, since there is only the most tenuous of correlations
between bone textures and actual soft tissue in living animals.

Can you guess what the animals in the following pages are? The
answers, as explained by the unknown scientists that will discover
and depict them in the future, are visible next to each image.
"The killer stare of the Cat , a vicious, pack-dwelling hunter. This
dangerous predator brandished a set of not one, but FIVE
switchblade claws on each forefoot. Its fossils are usually found in the
concrete nests of long-limbed, bipedal Humans , suggesting that the
killer Cats wandered into their lairs before slaughtering their hapless
victims."
"Only the skull of the Hippopotamus is known, yet even that is
enough to tell that this apex predator was the most dangerous hunter
of its time. Its long teeth and heavy jaws were strong enough to chew
right through the metallic armor of Cars ."
"The Cow was a lithe, graceful herbivore that despite its size, could
easily outrun pursuing hunters."
"With its extraordinary skin sail, the Rhinoceros was among the
strangest of the Holocene herbivores. Researchers suggest that this
organ might have helped it shed excess heat."
"Many theories have been suggested to explain the unusual, single-
digit feet of equid perissodactyls. It was likely that these animals
bore muscular pads on their hands and feet to support their stance."
"The Spider Monkey was a small, possibly arboreal species of
Human with extraordinarily long fingers and big eyes. Unlike its
cursorial relatives, the Spider Monkeys were efficient, stealthy
predators."
"A Toad ambles merrily along the Holocene forest floor. This
reconstruction is now considered to be erroneous since their longer
hindlimbs indicate that Toads were possibly bipedal."
Covering Up is not so Easy: the Integument
Problem
If no feathers are preserved in the fossil record, will future
palaeontologists guess at their presence? It seems unlikely. They will
possibly recognise that birds had wings, but fill them in the simplest
way: a membrane of skin. What about the other way around? Will
errors of preservation, or of phylogenetic bracketing, lead to mistaken
assumptions of furry body covering? Only time will tell.

"This reconstruction of the Vulture reflects our latest theories about

the wing membrane extensions in Flying Birds ,those mysterious
beings which appeared suddenly on the skies of our planet a
hundred million years ago."
"Two Casque-Headed Hornbills duel for dominance. These strange
beaked animals grew elaborate head horns for competition among
during the mating season."
"Two Swans are seen with their long, scythe-like forelimbs, which
they must have used to spear small prey items. One of them has just
caught a Tadpole , one of the mysterious fish of the past."
"Unlike many others, this reconstruction of the Iguana is fully up-to-
date. Impressions of a furry integument have been discovered around
the skeletons of Rats a few years ago. It is likely that all small
vertebrates had such body coverings to protect against the cool
Holocene climate."
The Biomechanics of Rabbits
Palaeontology is an imperfect and difficult science. Many
methodologies are brought to bear, and many contingencies must be
considered. Since 1999, Kent Stevens and colleagues have argued that
the vertebral joints of sauropod dinosaur necks can be effectively
modelled in digital space using a specially designed piece of software
called Dinomorph. The overriding conclusions from this work are
that neck flexibility in sauropods was rather restricted, and that
sauropod habitually held their necks in a more or less horizontal
pose.46 However, other researchers have argued that this work fails to
take proper account of the soft tissue anatomy present in animals. It
ignores, for example, the substantial contribution that cartilage makes
to neck flexibility, nor is it informed by the fact that virtually all
living terrestrial amphibians, reptiles, birds and mammals habitually
hold their necks in an elevated pose relative to the long axis of the
dorsal vertebrae.47

Perhaps, not realizing that neck posture in living animals is not

properly reflected in the way neck bones fit together, future
palaeontologists will model such living animals as rabbits in the same
way, concluding that they, too, held their necks horizontally. This
illustration thus serves as a warning against overly mechanistic
approaches to anatomy.
The Unknown Unknowns: Too Conservative,
Too Cautious?
Fossils are rarely complete, and naturally enough, we tend to fill in
the missing bits with conservative parts from the closest relatives
whenever we can. The problem is, the defining anatomical features of
many living animals are entirely composed of soft tissue, which is
hard to predict, even with educated guesswork. How daring and
how correct will the palaeontologists of tomorrow be?

"A Python rears up in a colorful, defensive body display. We know

this strange animal from fragmentary remains only, but it is likely
that it supported its long body with stubby legs, much like those of
closely related Lizards ."
"A solitary Manatee is shown grazing in its mountain home. We only
know the skull of this enigmatic herbivore."
"A bull Elephant , one of the gigantic mammals of ancient Africa, is
shown here inflating its balloon-like facial sac. Some fanciful
reconstructions depict Elephants with even longer muscular
appendages projecting from their faces, but this is unlikely since no
modern mammals have such organs."
"The enigmatic Whales were some of the largest animals to swim in
the ancient oceans. Here, we can see a couple of predatory Sperm
Whales looking for prey.
"Another dangerous predator, the elongated, serpentine Bowhead
Whale, is visible swimming through a tangled underwater forest.
This extremely derived predator fed on animals as big as itself, which
it trapped with its gigantic, extensible jaws."
Strange Hypotheses
Modern palaeontology has its fair share of strange and outlandish
hypothesis floating around, and these frequently find their way into
artworks and illustrations. What kinds of strange and uninformed
hypotheses might be formulated about present day animals?

Venomous Baboons

Lacking any and all knowledge of the thick woolly coats and brightly
coloured, naked skin patches and large guts of living baboons, it
seems plausible that these lightly built, long-muzzled, fanged
primates would be reconstructed as gracile terrestrial predators.
Furthermore, those giant fangs have grooves running down their
sides - a feature normally regarded as as key signature of the ability
to produce venom, and inject it into the tissues of other animals.
Perhaps the complicated nasal sinuses of baboons might be
interpreted as spaces for venom glands.

The skull of Sinornithosaurus.

Baboons, of course, are not venomous, and quite why they have fang
grooves isn’t entirely clear. Knowing this, suggestions of venomosity
sound more than a li le preposterous. However, it will be surprising
to know that very similar theories have been proposed for quite a few
fossil animals: the most notorious recent example concerning the
bird-like theropod dinosaur Sinornithosaurus.48 This hypothesis was
later rebuked by a number of experts, but not before it had been
reported by a number of media sources as a legitimate discovery.
Thus, this case also illustrates the importance of proper science
reporting in facilitating or limiting the spread of untrue or unproven
ideas.
Hummingbird Parasites

The future will no doubt have its fair share of bizarre and poorly
supported hypotheses, and not all unusual ideas are equal. Would it
be obvious that the slender, tubular (and sometimes serrated) bills of
hummingbirds were used to probe deep into flowers and feed on
nectar, or would alternative hypotheses be entertained? There is
much reason to think so. A hummingbird's skeleton is among the
strangest in the animal kingdom. Their wings are tiny, in fact shorter
than their legs. The large, strongly curved foot claws hint at a lifestyle
involving clinging and climbing, as do small curved claws on the
distinctive elongate hands.

Combine all of these features with a lack of evidence for their true
lifestyle, and we imagine that future palaeontologists could
misinterpret hummingbirds entirely. Vampiric hummingbirds sound
unlikely, but similar theories have been proposed about vampiric
pterosaurs. In 2003, researcher David Peters claimed that he had
discovered “ra lesnake fangs” and other indications of a blood- and
egg-sucking lifestyle in a small anurognathid pterosaur known as
Jeholopterus. However, Peters’ controversial research methods
involved relying on manipulated images on a computer to discern
features that no other observer can see. Both this methodology and
Peters’ hypothesis of vampiric pterosaurs were consequently
disproved by other scientists.
An Ending and Self-reflection: Homo diluvii
Having shown you a fair share of speculations, oddities and unusual
theories, we conclude our tour with perhaps the oldest "wrong
reconstruction" of them all. Behold Homo diluvii: the Man of the
Deluge. Described in 1726 by the Swiss Scholar Johann Jakob
Scheuchzer from giant salamander remains, the creature was thought
to be an actual human being, fossilized after drowning in the
mythical deluge of the Bible.
This mistake seems so obvious now that even children could realize
that the skeleton was not that of a human being. But Homo diluvii
shows us how we take scientific reasoning and our repository of
evidence and theories for granted. Science was still at its infancy at
the time this fossil was discovered. People didn't understand what
fossils were and tried to explain them with concepts they had at
hand, which invariably came from religious myths at the time.
Scholars were incapable of even conceiving that animals could leave
their remains as fossils in rock, or that different animals and plants
lived at different times, or that organisms could go extinct. Thus, a
salamander ended up a man in their worldview.

Over the centuries, we have come a long way from Homo diluvii.We
now know what fossil animals are, and can reliably formulate their
relationships to living beings today. Almost with each passing day,
new discoveries are providing us with insights into their lifestyles,
forms, colour, diversity and evolutionary relationships. Indeed, our
repository is growing faster than we can assimilate and get used to its
implications. It is possible, though unlikely, that paradigm shifts may
one day make our current view of extinct animals as odd and dated
as Homo diluvii itself.

This is why we prepared this collection: to show you some of our

mistakes, and hint at what we might be missing on the way. We don't
pretend to have any conclusive answers-the only solid statement we
can offer in this work is that no one should "rest easy" with the facts
on hand. Our picture of the past is constantly evolving, and it will
continue to evolve as new discoveries change or sweep old certainties
away. We hope you enjoyed your ride with us through All Yesterdays
and All Todays.
Notes and References
1 A minor note on terminology. Some people argue that the term
palaeoart (or 'paleoart' if you're American) should best be applied to
art produced during prehistoric times, and that artwork depicting
prehistoric times and prehistoric organisms therefore needs a new
name. However, the only word yet on the cards-palaeontography-has
yet to catch on.

2 Archosaurs are the 'ruling reptiles': dinosaurs, crocodiles and all of

their relatives.

3 Paul, G. S. 1987. The science and art of restoring the life appearance
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S. J. & Olson, E. C. (eds) Dinosaurs Past and Present Vol. II. Natural
History Museum of Los Angeles County/University of Washington
Press (Sea le and London), pp. 4-49.

4 Knight, C. R. 1959. Animal Drawing: Anatomy and Action for Artists.

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Image Credits

"Venomous Baboon" - © all rights reserved, by C.M. Kosemen

"Elephant" - © all rights reserved, by C.M. Kosemen
"Hippopotamus" - © all rights reserved, by C.M. Kosemen
"Homo diluvii" - © all rights reserved, by C.M. Kosemen
"Hornbill" - © all rights reserved, by C.M. Kosemen
"Lambeosaurus magnicristatus" - © all rights reserved, by C.M. Kosemen
"Rhinoceros" - © all rights reserved, by C.M. Kosemen
"Zebra" - © all rights reserved, by C.M. Kosemen
"Dinosauroid" - © all rights reserved, by C.M. Kosemen
"Majungasaurus" - © all rights reserved, by C.M. Kosemen
"Kahydron" and "Allotaur" - © all rights reserved, by C.M. Kosemen
"Spidermonkey" - © all rights reserved, by C.M. Kosemen
"Stegosaurus mounts a Haplocanthosaurus" - © all rights reserved, by C.M. Kosemen
"Torosaurus" - © all rights reserved, by C.M. Kosemen
"Swans" - © all rights reserved, by C.M. Kosemen
"Allosaurus and Camptosaurus" - © all rights reserved, by John Conway
"Camarasaurus" - © all rights reserved, by John Conway
"Carnotaurus" - © all rights reserved, by John Conway
"Cat" - © all rights reserved, by John Conway
"Centipede and a Anurognathid" - © all rights reserved, by John Conway
"Citipati" - © all rights reserved, by John Conway
"Cow" - © all rights reserved, by John Conway
"Elasmosaurus" - © all rights reserved, by John Conway
"Griffon Vulture" - © all rights reserved, by John Conway
"Hesperornis" - © all rights reserved, by John Conway
"Heterodontosaurus tucki" - © all rights reserved, by John Conway
"Heterodontosaurus tucki Skeletal" - © all rights reserved, by John Conway
"Hummingbirds" - © all rights reserved, by John Conway
"Hysilophodon eats a Millipede" - © all rights reserved, by John Conway
"Iguana" - © all rights reserved, by John Conway
"Leaellynasaura" - © all rights reserved, by John Conway
"Majungasaurus Makes Like a Log" - © all rights reserved, by John Conway
"Microraptor gui" - © all rights reserved, by John Conway
"Opisthocoelicaudia" - © all rights reserved, by John Conway
"Ouranosaurus" - © all rights reserved, by John Conway
"Parasaurolophus" - © all rights reserved, by John Conway
"Plesiosaur" - © all rights reserved, by John Conway
"Protoceratops andrewsi" - © all rights reserved, by John Conway
"Python" - © all rights reserved, by John Conway
"Rabbits" - © all rights reserved, by John Conway
"Seacow" - © all rights reserved, by John Conway
"Sleepy Stan" - © all rights reserved, by John Conway
"Sperm Whale" - © all rights reserved, by John Conway
"Tenontosaurus" - © all rights reserved, by John Conway
"Therizinosaurus" - © all rights reserved, by John Conway
"Toad" - © all rights reserved, by John Conway
"Triceratops horridus" - © all rights reserved, by John Conway
"Whale" - © all rights reserved, by John Conway
"Allosaurus Skeletal" - © all rights reserved, by Sco Hartman
"Camarasaurus Skeletal" - © all rights reserved, by Sco Hartman
"Carnotaurus Skeletal" - © all rights reserved, by Sco Hartman
"Majungasaurus Skeletal" - © all rights reserved, by Sco Hartman
"Microraptor gui Skeletal" - © all rights reserved, by Sco Hartman
"Nothronychus mckinleyi Skeletal" - © all rights reserved, by Sco Hartman
"Ouranosaurus Skeletal" - © all rights reserved, by Sco Hartman
"Parasaurolophus Skeletal" - © all rights reserved, by Sco Hartman
"Tyrannosaurus rex (Stan) Skeletal" - © all rights reserved, by Sco Hartman
"Stegosaurus Skeletal" - © all rights reserved, by Sco Hartman
"Tyrannosaurus rex in a Hurry" - © all rights reserved, by Sco Hartman
"Triceratops horridus Skeletal" - © all rights reserved, by Sco Hartman
"Bo lenose Dolphin and Dall's Porpoise" - © all rights reserved, by Darren Naish
Rabbit skull and neck photographs - © all rights reserved, by Darren Naish
"Goats in a Tree" CC BY 2.0 by Marco Arcangeli, from Wikipedia
Hippopotamus photograph - CC BY-SA 3.0 from Wikipedia
Andrias scheuchzeri skeleton photograph - CC BY-SA 3.0 by "Haplochromis" , from Wikipedia
"Sinornithosaurus skull" - CC BY-SA 3.0 from Wikipedia
"Allosaurus" and "Diplodocus" - Public Domain, by Charles Knight