UFOs: Demonic Activity & Elaborate Hoaxes

Meant to Deceive Mankind

Bro. Michael Dimond
A UFO or unidentified flying object is any flying object or phenomenon
that cannot be identified by the observer. 1 Over the last twenty-five
years, millions of people have claimed to see UFOs over every major
city in the world. Many polls (Roper, Gallup, PBS/Nova) claim that
almost 20 million Americans have claimed to see a UFO, 2 and over 4
million have claimed to have been abducted by aliens. 3 Some insurance
companies are even selling Alien Abduction Insurance. 4 People like
Jimmy Carter, 5 Barry Goldwater, and Prince Charles of England have
seen UFOs. 6 Reports of UFO sightings have been statistically proven to
be becoming more frequent with the passing of each decade. There are
now many universities that offer specialized studies on the subject of
UFOs. 7
The fascination with UFOs is a world-wide phenomenon. This
fascination is shown by the large numbers of people who went to see the
UFO film, Independence Day, which grossed more in its first week than
all other previous movies. 8 Over 90 percent of UFOs turn out to be
IFOs, that is, identified flying objects. 9 Naturally occurring phenomena
like balloons, searchlights, advanced aircraft, comets, bright meteors,
ball lighting, twinkling stars, 10 etc., are examples of things claimed to be
UFOs. But many other people have definitely seen a phenomenon that
cannot be explained naturally. The first part of this book will focus on
the UFOs that are truly unidentifiable. It will show that they are not
from the material world and are a spiritual phenomenon.
The Absolutely Impossible Flying Maneuvers of UFOs
UFOs have been tracked by scientists traveling at over 25,000 miles per
hour. 11 UFOs have also made right angle turns at an estimated 15,000
miles per hour. 12 They have made 90 degree turns in mid air 13 and they
have instantly reversed direction while moving at these high speeds.
Planes made by humans make sonic booms when they travel at such
speeds, but UFOs make no sonic booms. In fact, in most sightings of
UFOs, no noise is heard at all. 14 UFOs have sometimes been picked up
UFOs 1

on radar, and other times they cannot be picked up on radar. 15 UFOs

sometimes show up on photographs, but other times they do not. 16
While in flight UFOs frequently change size or shape in front of
people, 17 and they appear as strange lights. UFOs also change colors, 18
many times only when accelerating, with virtually every color of the
spectrum reported. 19 Sometimes UFOs instantly appear and disappear in
front of people and they will pass right through physical objects. People
have also seen multiple UFOs merge into one UFO, and one UFO turn
into many UFOs. There is no one that can explain, from a scientific
standpoint, how it is possible for any material craft to do these things.
Many have objected to the idea that UFOs are alien aircrafts precisely
because UFOs have performed flying maneuvers that are impossible for
any materially constructed craft. They violate the known laws of
science.
Physicist James McCampbell, speaking to an academic conference on
UFOs in 1975, stated: in flight, their startling departures, sudden
stops, and right angle turns at high speed require them to be virtually
massless. 20
Established in 1955, the British journal, Flying Saucer Review, is widely
recognized as the leading UFO publication in the world. Its authored by
over fifty experts from around the world, who conduct major
investigations of UFOs. 21 Gordon Creighton was the editor of the
journal until he died in 2003. 22 In 1996, Creighton made the following
official position statement on UFOs: There seems to be no evidence
yet that any of these craft or beings originate from outer space. The
whole phenomenon involves a mass of features that conflict with modern
science, and many researchers now believe that more than one type of
being may be involved, some of them originating from outer space and
some of them of an interdimensional nature, and consequently
possibly from some unknown aspect of our own world... 23 Many
UFO researchers have now abandoned the ETH (extraterrestrial
hypothesis) for the IDH (interdimensional hypothesis). 24 Creighton also
said about UFOs: I do believe that the great bulk of these
phenomena are what is called satanic. 25
Charles Bowen is the current editor of the Flying Saucer Review.
Speaking about UFOs and their occupants, he said: I have pondered
over the idea that the frightening, spooky creatures described by some
witnesses could be some sort of psychic projection. There are noticeable
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dreamlike qualities about the incidents described in these cases. Is it

possible that something from somewhere is coming here and by means
incomprehensible to us? 26
To find out more about what UFOs really are, we need to listen to what
the worlds top experts on UFOs have concluded about the UFO
phenomenon.
The Top Experts in the World on the UFO Phenomenon
Dr. Jacques Vallee is a computer scientist and astrophysicist. Vallee has
addressed the UN on the subject of UFOs and is widely recognized as the
premier expert on the subject. He has done computer analysis on tens of
thousands of UFO reports. He has looked for patterns that could be clues
to an explanation of UFOs. Vallee has also authored eight books on
UFOs and was the real life basis for the character of the French scientist
Claude LaCombe in the UFO movie Close Encounters of the Third
Kind. 27
In May 1955, Vallee saw a UFO flying over his home in France. 28 In the
mid-1960s, Vallee initially attempted to validate (like many other UFO
researchers) the Extraterrestrial Hypothesis or ETH. 29 But by 1969,
Vallees conclusions had changed. He publicly stated that the ET
Hypothesis ignored too much data. 30
Vallee: we are dealing with a yet unrecognized level of
consciousness, independent of man but closely linked to the earth. For
reasons that will become clear, I do not believe any more that UFOs are
simply the spacecraft of some race of extraterrestrial visitors. 31
Vallee: Consider what these [UFO] sightings have in common. In each
case the so-called spacecraft did not disappear by moving away, even at
high speed. It simply vanished on the spot, or it slowly faded away...
In other cases, UFOs have been reported to enter the ground. I hardly
need to point out that this behavior is contrary to what physical objects
do, and quite impossible to duplicate with our current spacecraft
technology. It is the behavior of an image, or a holographic projection.
Yet at other times the objects have left material traces. 32

UFOs 3

Vallee: Most witnesses are not primarily reporting an object or a craft;

they are reporting a light, a massive, multicolored, intense, pulsating
light, a playful, fascinating, impressive, hypnotic light 33
Vallee: I propose that the UFO we see is, among other things, a
device which creates a distortion of the witnesss reality; that it does
so for a purpose, which is to project images or fabricated scenes
designed to change our belief systems; and that the technology we
observe is only the incidental support for a worldwide enterprise of
subliminal seduction They are helping to create a new form of belief:
an expectation of actual contact among large parts of the public. In turn
realization of that age-old dream: salvation from above, surrender to the
greater power of some wise navigators of the cosmos. 34
Vallee: Using a single probe the size of a beer keg in orbit a thousand
miles above the earth, human technology as it exists today would be able
to capture in a few weeks most of the important facts about the planets
geography, weather, vegetation, and culture. Given the earths abundant
emission of radio and television programs for the last fifty years, even
the deployment of such a space probe might be unnecessary. This is one
of the little-recognized facts of the UFO problem that any theory has yet
to explain. The theory of random visitation does not explain it. Either
the UFOs select their witnesses for psychological or sociological reasons,
or they are something entirely different from space vehicles. In either
case, their appearances are staged! 35
Vallee: I believe that when we speak of UFO sightings as instances of
space visitations we are looking at the phenomenon on the wrong level.
We are not dealing with successive waves of visitations from space. We
are dealing with a control system. 36
Vallee, speaking of the UFO phenomenon, said the UFOs cannot be
spacecraft in the ordinary nuts-and-bolts sense. The UFOs are physical
manifestations that cannot be understood apart from their psychic and
symbolic reality. What we see here is not an alien invasion. It is a
spiritual system that acts on humans and uses humans. 37
Vallee: I think UFOs are perpetrating a deception by presenting their
so-called occupants as being messengers from outer space 38

UFOs

Vallee: The UFO occupants are not extraterrestrials. They are the
denizens [inhabitants] of another reality. 39
Conclusions on UFOs from Some Other Top Experts
Astronomer Dr. J. Allen Hynek, who died in 1986, was considered one
of the top experts in the world on the subject of UFOs. Hynek was a
former skeptic who dismissed UFO sightings as something made up by
kooks and crackpots. 40 As he continued to study the issue, his view
changed. Dr. Hynek eventually started the Center for UFO Studies
(CUFOS) with Dr. Jacques Vallee. Hynek also headed the U.S. satellite
optical tracking program for many years. 41 He was the author of many
technical textbooks on astrophysics and several books on UFOs. When
Hynek investigated UFOs, he used the services of accomplished and
skilled scientists with a scientific approach to all the data. Hynek served
for over twenty years as the Air Forces scientific expert on UFOs. 42 He
studied thousands of reports, and interrogated hundreds of witnesses to
UFO experiences. 43 Hyneks conclusions on UFOs were accurately
summed up this way: Hynek submitted that perhaps UFOs were part of
a parallel reality, slipping in and out of sequence with our own. This was
a hypothesis that obviously pained him as an empirical scientist. Yet
after 30 years of interviewing witnesses and investigating sighting
reports no other hypothesis seemed to make sense to him. 44 Hynek
repeated this view when he was interviewed by Newsweek. 45
Speaking about UFOs, Dr. Hynek said the following: One of the most
intriguing aspects of UFO sightings is their apparent isolation in space
and time. Sightings are usually made in one place at a time and last for
short periods, sometimes appearing out of nowhere. This is much more
suggestive of a parallel reality than, lets say, visits from some far away
planet, in which case, we might expect them to stick around for a
while. 46
Dr. Hynek: Another peculiarity is the alleged ability of certain UFOs
to dematerialize... There are quite a few reported instances where two
distinctly different UFOs hovering in a clear sky will converge and
eventually merge into one object... 47
Dr. Hynek: Many UFO reports seem to pertain more to accounts of
poltergeists(cases where objects fly around the room and strange
sounds are heard) and other types of psychic manifestations than to
UFOs 5

actual solid items of nuts and bolts hardware. That is one of the reasons
why I cannot accept the obvious explanation of UFOs as visitors from
outer space. 48
Dr. Hynek and Dr. Vallee describe this strange behavior of UFOs in their
book, The Edge of Reality: If UFOs are, indeed, somebody elses nuts
and bolts hardware, then we must still explain how such tangible
hardware can change shape before our eyes, vanish seemingly melt
away in front of us, or apparently materialize mysteriously before us
without apparent detection by persons nearby or in neighboring towns.
We must wonder, too, where UFOs are hiding when not manifesting
themselves to human eyes. 49 Dr. Hynek concluded about UFOs: The
UFO phenomenon is the outstanding strange dilemma of our age. We
dont know what they are. 50
Another one of the top living experts in the world on UFOs is John
Keel. 51 Keel began to investigate UFOs on a full time basis in early
1966. During just one four-year period of time, Keel interviewed
thousands of people who claimed to see UFOs in more than twenty U.S.
states. He reviewed more than 2,000 books, in addition to thousands of
newspapers, magazines, and newsletters. 52 Initially, like Dr. Vallee,
Keel was hopeful that he could validate the ET visitation hypothesis, but
abandoned that hypothesis after about one year investigating UFOs. 53
Keel has authored many books on UFOs. He said: The statistical data
that I have extricated indicate that flying saucers [UFOs] are not stable
machines requiring fuel, maintenance and logistical support. They are,
in all probability, transmogrifications of energy and do not exist in the
same way that this book exists. They are not permanent constructions
of matter. 54
Keel: Sometimes they [UFOs] reportedly turn a brilliant red and vanish.
Other times they shift through all of the colors of the spectrum, turn
white, and fly off into the night sky until they look like just another
star. 55
Keel: There are countless sightings of objects that changed size and
shape in front of the viewers or split into several smaller objects, each
going off in a different direction. In some cases, this process was
reversed, with several small lights converging together to form a single
large one, which then went dashing off. Over and over again, witnesses
have told me in hushed tones, You know, I dont think that thing I saw
6

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was mechanical at all. I got the distinct impression that it was

alive. 56
Keel: The UFOs do not seem to exist as tangible, manufactured objects.
They do not conform to the accepted natural laws of our environment...
The UFO manifestations seem to be, by and large, merely
minor variations of the age-old demonological phenomenon. 57
Keel: Many flying saucers seem to be nothing more than a disguise for
some hidden phenomenon. They are like Trojan horses descending into
our forests and farm fields, promising salvation and offering us the
splendor of some great supercivilization in the sky. 58
Keel, speaking about the UFO flying aircraft: Our UFO catalog now
contains flying cubes, triangles, hexagons, doughnuts, spheres, objects
shaped like giant metal insects and transparent flying jellyfish. Weve
got UFOs with wheels, with wings, with antennas, with pointed domes,
flat domes, no domes at all. Weve got objects of every color of the
spectrum. There have been giant, multi-windowed cigars spitting blue
fire from their tails We have just about everything except a basic
assembly-line model Some tremendous unknown civilization is
exerting an all-out effort to manufacture thousands of different types of
UFOs and is sending all of them to our planet. 59
Keel, speaking about the descriptions made by people who have claimed
to have been abducted by aliens from the UFOs, said: The reported
UFO entities are also a mixed bag, not only in size but in general
appearance. There have been UFOnauts [aliens] with pupil-less eyes,
glowing eyes, eyes as big as saucers, tiny beady eyes, wraparound sliteyes, cat-like eyes, and even no eyes at all. There have been noseless
entities and mouthless entities. Ears have covered a wide range from
being pointed to being elephant-like. 60
Does this make any sense? If some alien civilization were visiting or
invading our world, their vehicles and personal appearance would look
basically the same. There wouldnt be vast differences among the
saucers and the appearances of the aliens.

UFOs 7

Could Aliens from Another Galaxy Travel to Earth?

According to Einsteins theory of special relativity, the speed of light is
the maximum speed possible. 61 If someone were travelling at the speed
of light, they would be travelling at the incredible speed of 186,000 miles
per second. Travelling at this speed you could circle the earth seven
times in just one second. If you took off from earth, you would pass the
moon in two seconds. 62
If aliens came from another galaxy, they would have to travel from a
galaxy at least two million light years away. Even if they travelled at the
speed of light (186,000 miles per second), it would take them two million
years to get to earth. But what if the UFOs travelled from some distant
star in our own galaxy? If these space travelers were to travel one
million miles per day from the closest star system (alpha centuria)
which is more than twenty-five trillion miles from the earth it would
still take them 70,000 years to arrive at earth.
Some estimates indicate that a space ship carrying ten people, and
travelling five light years to and from a nearby star system at almost the
speed of light, would use up 500,000 times the total amount of energy
consumed in the US in one year. 63 Also, an almost equal amount of
energy would be consumed just in slowing the aircraft from the
incredible speed at which it was travelling. For larger crafts, such as the
city-sized spaceships shown in the UFO movie Independence Day, the
required energy needed would be even more staggering. 64 Scientists like
Donald Menzel rejected ETs visiting earth because even the nearest stars
are too far away for interstellar travel to be possible. 65 But there are
other problems with space travel. It has been estimated that there are
100,000 dust particles per cubic kilometer of space. Travelling at the
speed of light (186,000 miles per second), an impact with even one of
these tiny dust particles would destroy a spaceship. At one-tenth the
speed of light, the impact would be equal to an explosion of almost ten
tons of TNT. 66 Based on these scientific facts alone, without even
considering all of the others about the true nature of UFOs, space travel
by some alien craft is extremely unlikely.

UFOs

Is Life Sustainable in Outer Space?

The conditions outside Earth are far more destructive than most people
think. For example, prolonged exposure to the cosmic rays in space can
be fatal. 67 There are also poisonous gases, deadly radiation, gigantic
explosions, extreme gravitational forces, and the lack of proper
atmospheres and chemical elements for life to exist. Just the temperature
extremes in outer space would make almost any form of life either too
incredibly hot or so cold it would be entirely rigid, brittle and dead. 68
Dr. Vallee held that, based upon the descriptions of many people who
claimed to have been abducted by aliens from UFOs, their bodies could
never sustain the rigors of space travel. Vallee stated: the humanoid
body structure of the alleged aliens is not likely to have originated on
another planet and is not biologically adapted to space travel. 69 Vallee
listed this as one of the main reasons that UFOs were not ETs. Also,
how do the aliens in UFOs breathe and live in our Earths air
conditions without the aid of respiratory devices? 70
The DVD Creation and Miracles, Past and Present 71 shows that there is
no scientific evidence for the theory of evolution. Nor is there any such
evidence in the Bible. But a large number of astronomers believe that
life evolved on earth, and therefore (according to them) it must also have
evolved on other planets in space. If this were true, many alien races on
other planets, which have been around a lot longer than we have, should
exist. Some of them would be much more technologically advanced than
we are. And if someone believed that somehow advanced aliens could
reach us with aircraft, these older and more advanced beings probably
would have already colonized our solar system, at least with robots. 72
While working at the Los Alamos, New Mexico nuclear facility in
1950, pioneer of atomic energy and Nobel Prize winner Enrico Fermi
asked the following question: Are we the only technologically advanced
civilization in the universe, and if we are not, then where are they?
Why havent we seen any traces of extraterrestrial life such as probes
or transmissions? 73 If some intelligent aircraft couldnt reach our solar
system with an advanced aircraft, they would at least send us an
intelligent radio signal.

UFOs 9

Has any Signal of Intelligent Life Come from Outer Space?

Around the globe, SETI (Search for Extra Terrestrial Intelligence)
researchers have searched outer space waiting for signals from intelligent
beings. They have listened on thousands of frequencies over the years.
Nothing indicating intelligent life has ever been heard. 74 In fact, all radio
astronomers will admit that they have never received any intelligent
signal from outer space. 75 After an interview with SETIs senior
astronomer, Seth Shostak, National Geographic magazine stated: He
and his colleagues have never found proof anyone or anything
up there is trying to make contact. 76
The Government of the U.S. has spent millions of tax dollars attempting
to contact aliens. In the 1980s and 1990s astronomers conducted the
largest search for ET life through the META and BETA programs. They
simultaneously scanned millions of radio frequencies for any signs of
intelligent communications, with no positive results. 77
Moreover, the official website for NASA states the following about
UFOs: The U.S. Air Force (USAF) and NASA have had intermittent,
independent investigations on the possibility of alien life on other
planets; however, none of these has produced factual evidence that life
exists on other planets, nor that UFOs are related to aliens. 78
Dr. Edward Condon was a physicist who led Colorado Universitys Air
Force-financed two-year study of UFOs. The final report of this study
was published in January of 1969. In a speech in April 1969, Condon
said: Some UFOs may be such [ET] visitors, it may be postulated, and
some writers go so far as to say that they actually are. To discover clear,
unambiguous evidence on this point would be a scientific discovery of
the first magnitude, one which I would be quite happy to make. We
found no such evidence, and so state in our report 79
A Few Major UFO Sightings Throughout History
One of the earliest descriptions of a UFO was by the ancient Egyptians
in approximately 1,500 BC. The Egyptians described seeing flying
discs. They also noted a foul odor that emitted from the spacecraft. 80
The deceased satanist Aleister Crowley, who described himself as the
beast 666, wrote how to guides on magic and contacting angels.
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Crowley was in contact with higher intelligences while writing The

Book of the Law in Cairo, Egypt in 1904 at the behest of a being called
Aiwass... It was Crowleys goal that through contact with non-human
intelligent beings [demons or aliens] to bring to fruition the New
Age... 81
Crowley claims to have contacted an alien intelligence and he made a
drawing of what it looked like in 1918. Crowley said the being was
named Lam. Crowley included the picture of Lam in his Dead Souls
Exhibition held in Greenwich Village, NY, in 1919. 82 Crowleys
drawing of Lam bears an incredible resemblance to the common visual
conception of the Gray ET. The Gray Aliens are the most common
alien claimed to have been seen by abductees of ETs. The Grays emit
a foul odor and sometimes appear naked to abductees. 83
On the night of October 25, 1973, fifteen people, including a young
farmer from Pennsylvania named Steven Pulaski, saw a bright red ball
hovering over a field nearby. Steven took his rifle and went out to
investigate. As the ball descended, its color changed from red to bright
white. Screaming sounds could be heard nearby. Two creatures then
appeared. They had greenish-yellow eyes and a burning rubber smell
was around them. Steven got the sense that the creatures were not
friendly, and fired a warning shot over their heads. The creatures
continued approaching, so Steven fired directly at them. The creatures
and the glowing object then instantly disappeared from the field.
Other witnesses also heard the strange noises from the woods and
noticed the creatures. A state trooper and UFO researchers were then
called in to investigate. Upon arrival, the people there said that Steven
had been violently flailing his arms and growling like an animal.
Stevens own dog ran toward him, and Steven attacked his dog. Steven
then made prophecies of doom in a mocking tone of voice, accompanied
by laughter. Steven then collapsed and, after a short time, came to his
senses. The group of people present during these events commented on
the nauseating sulfur-like odor that was present. 84 This odor is a
characteristic of many UFO incidents. It should be noted that cases of
demonic possession are also frequently characterized by horrible smells,
including sulfur-like odors.

UFOs 11

In addition to the people who have claimed to see UFOs, there are
others who have claimed to have been abducted by the aliens that
they believe are flying in the UFOs.
Whitley Strieber
Whitley Strieber claims to have had real experiences of abduction by
alien beings on a non-physical level 85 on more than one occasion.
He wrote of some of his experiences in his bestselling books,
Communion and Transformation.
Whitley Strieber: I would say I was assaulted by something from the
unknown, rather than possessed by it. I hope I was never possessed by it.
Although there are those who would disagree with me... After what
happened to me (the communion experience) I decided it might be a
good idea to accept the idea of the devil just in case thats what I saw. 86
Strieber described this experience when he claims he was transported
back in time and described one of the beings as what seemed almost
to be a demon with a narrow face and dark, slanted eyes. 87
Strieber: Terrible fears raced through me: They were predators and they
were going to eat our souls; they were demons and they were going to
drag me off to hell. 88
Strieber: And before the great power of the visitors, what would their
counsel have gained me? I was alone with this and at that moment I
was in hell. 89
Strieber: I worried that they would come and steal me away. Or
perhaps they would kill me and eat my soul. 90
Strieber: Increasingly I felt as if I were entering a struggle that might
be even more than life or death. It might be a struggle for my soul, my
essence, or whatever part of me might have reference to the eternal.
There are worse things than death, I suspected... So far the word demon
had never been spoken among the scientists and doctors who were
working with me. And why should it have been? We were beyond such
things. We were a group of atheists and agnostics, far too sophisticated
to be concerned with such archaic ideas as demons and angels... Alone
at night I worried about the legendary cunning of demons... At the very
least, I was going stark, raving mad. 91
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Strieber: Why were my visitors so secretive, hiding themselves behind

my consciousness? I could only conclude that they were using me and
did not want me to know why... What if they were dangerous? Then I
was terribly dangerous because I was playing a role in acclimatizing
people to them. 92
Strieber: Of course they were demons. They had to be. And they were
here and I couldnt get away 93
Strieber: and [I] felt my soul being carried away by a triumphant
spiritual predator I wondered if I might not be in the grip of demons,
if they were not making me suffer for their own purposes, or simply for
their enjoyment. 94 Strieber said he could smell the presence of the
aliens and they smelled like sulfur. 95
Common Abduction Experiences & Medical Examinations
Conducted by Aliens on Abductees
Reports of alien abductions have skyrocketed since the late 1980s. Dr.
John Mack, Harvard psychiatrist, has stated that he thinks four to five
million Americans believe they have been abducted by aliens. 96
Here is a brief description of some of the things that have happened to
people who have claimed to have been abducted by aliens from UFOs:
burns, medical experiments, mutilations, mustard gas smells, grotesque
appearances, false prophecies, blatant lies, and occultic messages. 97
Other people who have claimed encounters with aliens have said they
experienced things like headaches, dizziness, nausea, confusion,
helplessness, despair, and various pains during the encounter. Some
abductees have even died or committed suicide after being contacted by
UFOs or their occupants. 98 Animals very frequently have given the
impression of being spooked just before and during UFO sightings. 99
Sometimes animals have been frightened to the extent of injuring
themselves in their fright. Inanimate objects, most commonly vehicles
and their engines, were disabled, and the headlights died or were
dimmed. 100 The radios would also cut out or exhibit uncommon static.
In such cases, the vehicles returned back to normal operation after the
UFO departed from the scene. 101

UFOs 13

These aliens are demons and bear the marks of demons. Here are more
of the things that happened to abductees after they had contact with these
aliens. The aliens altered the perception of their surroundings. Aliens
would control what the abductees were seeing. They appeared under
different forms and shapes. The aliens passed through walls and
windows, and sometimes materialized and dematerialized directly in
front of them. Aliens sometimes put marks or scars on the bodies of
abductees. These marks have included bruises, cuts, single and multiple
punctures, three and four-fingered claw marks, and triangles of every
sort.
Speaking about abductions, Joe Jordan, director for the mutual UFO
network, said: The similarity between the [UFO] abduction experience
and demonic possession is very, very close. 102
Abductees have also reported being scoffed at, jeered at, and threatened
by their alien captors. Some said they were subjected to different kinds
of medical experiments and examinations. Typically, special attention
was given to the genital area 103 and some claimed painful genital and
anal probes were performed. 104 Dr. John Miller reported that when
abductees asked the aliens why invasive and often humiliating medical
procedures were being performed, the aliens would often respond with
a statement like, we have a right to do this. 105 Abductees have been
threatened that very bad things would happen to them if they didnt
co-operate with their alien captors. 106 The medical examinations were
performed by the aliens against or irrespective of the will of the
abductee. These examinations often focused on sex and reproductive
biology. 107 Some abductees have claimed that the aliens have also
forced their human abductees to have sexual contact or intercourse. 108
Sometimes the aliens have spoken to the abductees. Generally, the
aliens questioned the abductees on their sexual practices or about life
on earth. 109
Science writer Jim Schnabel has tied modern-day alien abduction story
narratives to stories of 16th-17th century demonic possession and
witchcraft cases, some current third-world spirit possession syndromes,
and Satanic Ritual Abuse and sexual abuse claims that mesmerized many
American psychiatrists in the 1980s and 1990s. 110
Gwen Dean, a California-based therapist, noted forty-four parallels
between Satanic Ritual Abuse (SRA) and alien abductions. Both
14

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became widespread in the late 1970s and early 1980s. Victims of both
use hypnosis to recover suppressed or lost memory. Events and
narratives explained by SRA victims and alien abductees feature many
similarities. These similarities include how the alien examination table
is similar to the satanic altar, and how both phenomena focus on rape,
sexuality, and breeding. 111
Astronomer Carl Sagan also pointed out that the modern alien abduction
experience is remarkably similar to common stories of demonic
possession throughout history. Speaking of the similarities, Sagan said:
most of the central elements of the alien abduction account are
present, including sexually obsessive non-humans who live in the sky,
walk through walls, communicate telepathically, and perform breeding
experiments on the human species. 112
If the alleged aliens have super-advanced technology, why would they
need to repeatedly perform crude medical experiments on person after
person to discover what they need to find out?
Dr. Jacques Vallee: The medical examination to which abductees are
said to be subjected, often accompanied by sadistic sexual manipulation,
is reminiscent of the medieval tales of encounters with demons. It makes
no sense in a sophisticated or technical or biological framework: any
intelligent being equipped with the scientific marvels that UFOs possess
would be in a position to achieve any of these alleged scientific
objectives in a shorter time and with fewer risks. 113
Vallee: abductees of both sexes have described having intercourse
with the aliens. 114
Vallee: I pointed out in Invisible College that the structure of abduction
stories was identical to that of occult initiation rituals. 115
Vallee: An equally impressive parallel could be made between UFO
occupants and the popular conception of demons 116
Vallee: The belief pattern that has been created around the UFO
abductions is reminiscent of medieval theories of abduction by demons,
pacts with Satan complete with the Mark of the Devil on the body of
the witch. As French sociologist Pierre Lagrange has pointed out to me,
UFOs 15

the only missing element is the familiar the black cat or the owl that
used to accompany the witches! 117
Scientist Dr. Pierre Guerin of the Paris Astrophysical Institute 118 said:
UFO behavior is more akin to magic than to physics as we know it...
the modern UFOnauts [aliens] and the demons of past days are probably
identical. 119
Psychologist Elizabeth Hillstrom has pointed out that a growing number
of academics support the conclusion that aliens are synonymous with
historical demons. 120
Speaking of the ancient study of demonic possession, John Keel said:
Thousands of books have been written on the subject [demonic
possession] The manifestations and occurrences described in this
imposing literature are similar, if not entirely identical, to the UFO
phenomenon itself. Victims of demonomania (possession) suffer the
very same medical and emotional symptoms as the UFO contactees. 121
Hypnosis Brings Out the Detailed Stories of UFO Abductions
Most stories of UFO abductions are only recalled under hypnosis.
Hypnosis puts a person into an altered state of consciousness. It puts a
person in a highly suggestible state. For example, the hypnotist Kreskin
proved this in an experiment with a group of fourteen people. After he
put them under hypnosis, they were told (after being taken outside) that
a UFO was in the sky, and everyone in the group claimed to see a UFO.
The hypnotist then took them out of the hypnosis, got them in front of a
TV camera, and asked them: Have you ever seen a UFO? They
answered: No. Yet, they just claimed they had seen one moments
before. 122 This proves that a person can be made to believe that he saw
a UFO if he has been hypnotized. Numerous studies have demonstrated
that hypnotic regression can cause memories of events that never
happened; 123 these aid in exaggerations that are stated as facts. 124
Dr. Martin Orne is one of the leading experts on hypnosis and is a former
president of the International Society of Hypnosis. Orne has cited
experiments conducted as far back as 1961 which demonstrated that it is
possible for even deeply hypnotized subjects to willfully lie and it is
possible for an individual to feign hypnosis and deceive even highly
experienced hypnotists We should keep in mind that psychologists and
16

UFOs

psychiatrists are not particularly adept at recognizing deception... the

average hotel credit manager is considerably more adept at recognizing
deception than we are. 125 Orne went on to say that the clinicians
belief in the historical accuracy of the memories brought forth under
hypnosis is likely to be erroneous. 126
Hypnosis is practiced by witchdoctors and yogis and is certainly an
unscientific process. 127 Under hypnosis a person can also come under
demonic activity or possession. TM (Transcendental Meditation) is a
practice in which a person is put into a hypnotic trance. Ufologist Don
Elkens said: I have found that some people can achieve the [UFO]
contact phenomenon simply by being hypnotized 128
Considering the examples above, the fact that most UFO abduction
experiences are usually never fully revealed by UFO abductees until
the abductees are put under hypnosis is important. As has been proven,
just because a story is told under hypnosis doesnt mean at all that the
story is true and that the experiences were real. For example, there was
a criminal case... in which a man was convicted of murdering his wife on
the basis of testimony given by his son under hypnosis. Some time then
passed, and the wife turned up alive and healthy. 129
The Famous Betty & Barney Hill Abduction
One night Betty Hill (a believer in UFOs) was driving home, and she
believed that she and her husband were being followed by a bright, starlike object. Sometime after this event, Betty had a nightmare for four
nights in which she and her husband were taken aboard a flying
saucer. 130 Betty was convinced she had been abducted by aliens, but
Barney wasnt so sure. The Hills decided to undergo hypnosis so that
they would better recall what might have happened or what this meant.
Under hypnosis, the Hills gave details concerning what they believed had
happened to them. Betty and Barney disagreed on the appearance of the
aliens. 131 Betty said that the aliens spoke to her in English with a foreign
accent, while Barneys recollection contradicted that assertion. 132 Under
hypnosis, Bettys description of the short men, whom she claimed to see
in the UFO encounter, was different from how she described them from
her dreams. Barneys hypnotic sessions were often quite emotional,
punctuated with angry outbursts, expressions of fear, and episodes of
hysterical crying. Betty later claimed to see UFOs a number of times
UFOs 17

after her original abduction and became a celebrity in the UFO

community. 133
The Abduction Experience
Most of the abductees have reported being taken from their bedroom
right before falling asleep. Typically, at the beginning of the abduction
experience, the abductee reports paralysis, the freezing or slowing of
time, or seeing a bright light, and the appearance of humanoid figures. 134
This experience is not an abduction, but is in reality a visitation by a
demon.
Bud Hopkins has written many books on UFOs. He is probably the
foremost expert in the world on UFO abduction cases. He stated:
Now we have come to the conclusion that [alien] abduction their
involvement with human beings is the center of the entire UFO
phenomenon... Actually, when people have UFO sightings, it is very
possible that what they are seeing is something [an alien] on its way
to or from an abduction of somebody. They are not just flying around
up there; they are interested very profoundly in human beings... These
experiences sound crazy; it sounds impossible. It is impossible,
according to our physics. But we often use hypnosis to flesh out all
the recollections. 135
Bud Hopkins: One thing that has given credence to the phenomenon of
UFO Abductions, aka Close Encounters of the Third Kind, is the fact
that many of the victims dont remember their experiences until they are
hypnotically regressed to the time when the incident occurred. That is,
the victim only consciously remembers, at most, a UFO sighting. But he
or she has symptoms, often including time that cannot be accounted for,
which cause the victim to seek professional help, resulting in a hypnosis
session. The hypnotist puts the victim in a trance and tells him or her to
return to the time of the event. Then the hypnotist asks the victim
questions about what happened, and the victim tells a story about an
abduction. 136 Some abductees under hypnosis will relate their event
calmly, while others may cry in apparent horror, shout angrily, tremble
with fear, or beg pathetically for the event to stop. 137 Bud Hopkins has
also claimed that a person could be a UFO abduction victim without even
remembering having seen a UFO or having been in a flying saucer. 138

18

UFOs

Dr. Leo Spinkle is a scientist who conducted a five-year study of 82

people who reported UFO experiences. Most of them claimed to be
able to re-live the experiences they had through hypnosis, and all
suffered severe psychological disturbances afterwards. 139
UFO expert John Keel stated: Millions of people have been affected at
least temporarily by UFO contact; thousands have gone insane and ended
up in mental institutions after their experiences with these things
began. 140
Surveys have also confirmed that a vast majority of abductees have
shown an interest in the New Age movement and Eastern religions.
Many abductees have also claimed to have past involvement with
astrology, astral projection, witchcraft, Ouija boards, and
channeling. 141 This supports the argument that UFO phenomena are
a result of demonic activity.
Most people who have claimed to have been abducted by aliens are
almost always totally changed after the abduction. Their view of the
world is changed and many develop a deep interest in the occult and
paranormal or mystical activities. 142 UFO researcher Trevor James
said: A working knowledge of occult science is indispensable to
UFO investigations. 143
Many believers in aliens think that by allowing themselves to be put into
a hypnotic trance they can be used as a channeling device through which
the alien can speak through them or to them. Many times the person who
is used as a channeling device is called a walk-in. 144
Ruth Montgomery is a New Ager and a famous automatic writer who
has claimed to receive some messages from aliens. Ruth Montgomery
says this about walk-ins: A walk-in is a high-minded entity that is
permitted to take over the body of another human being who wishes to
depart. 145 Walk-ins are actually what the Bible teaches is the reality of
demonic possession. Many UFO believers have invited alien entities to
take over their bodies. They say their goal is to usher in a new age. It is
claimed by many New Agers that these walk-ins are very highly evolved
entities that are seeking embodiment inside of a human being. 146 Also,
interviews done with ex-Satanists and Wiccans indicate a high level of
contact with various alien types, especially during coven meetings and
holiday rituals. 147 1 Timothy 4:1 says: The spirit clearly says that in
UFOs 19

the last times some will abandon the faith and follow deceiving spirits
and things taught by demons.
In 1969, the United States Printing Office published a 400-page
document authored by Lynn Catoe called UFOs and Related Subjects, an
Annotated Bibliography. Catoe was senior bibliographer for the Library
of Congress. Catoe read over one thousand books, articles, and other
literature during her investigation of UFOs. 148 Catoe gave a general
conclusion of her findings in the preface of her bibliography: A large
part of the available UFO literature is closely linked with mysticism and
the metaphysical. It deals with subjects like mental telepathy, automatic
writing and invisible entities as well as phenomena like poltergeist
manifestations and possession. Many of the UFO reports now being
published in the popular press recount alleged incidents that are
strikingly similar to demonic possession and psychic phenomenon that
have long been known to theologians and parapsychologists. 149 Catoes
document was compiled for the United States Air Force and is currently
in the Library of Congress.
The Aliens Messages to UFO Abductees
If you went to a bookstore to find a book on UFOs, you would probably
find the book in the New Age/Occult section, not the science section. 150
The belief in UFO contact through visits and communication with aliens
is a part of the New Age philosophy.
Most of the abductions happen at night time, and the person then wakes
up. In their messages, the aliens have said the Bible is outdated or false,
and that Jesus Christ is not God. The aliens say that we need to unite
into a global religion and government. 151 These aliens have also made
false prophecies, promoted a false Jesus, and attempted to explain that
sin doesnt exist. In their messages, the aliens say that they are our
creators, but we can also evolve into gods. In some messages they say
that everyone is God. 152 This is simply the original temptation from the
Devil in the Garden of Eden. For God doth know that in the day ye eat
thereof, then your eyes shall be opened, and ye shall be as gods,
knowing good and evil (Genesis 3:5).
Also interesting is that certain aliens have warned of a time of tribulation
and disasters which will occur in the near future. These predictions bear
a close resemblance to certain predictions in the Bible. Their goal is to
20

UFOs

increase the credibility of their own message by incorporating elements

of well known and true prophecies.
Robert Prentiss (a former California policeman) and his wife claimed to
see a multitude of UFOs. The UFOs appeared in a variety of different
colors and shapes over a period of weeks. Prentiss said: During the
weeks of tantalizing sightings, I became totally obsessed with the UFOs,
convinced that something great was about to happen. I abandoned my
daily Bible reading and turned my back on God. 153
One reason that many people are so interested in UFOs is that they hope
for an alien salvation. They hope that we might soon make contact with
some advanced race of people out there which has discovered the secrets
of how to escape death. For many, following UFOs has become a
substitute religion. Whitley Strieber remarked: The only thing now
needed to make the UFO myth a new religion of remarkable scope and
force is a single undeniable sighting. 154
Common Sense Questions About UFOs
If the aliens are all around us, why is it that no one has ever seen them
getting out of their spacecrafts to eat or drink anything? We dont see
them landing to take some food and water back to their spacecraft. If the
aliens really wanted to make contact with us, why would they land in
remote areas and make personal contact with just a few people? Why
wouldnt the aliens land in a major city and appear in front of huge
crowds of people? How come no one has found even one UFO landing
area or base when there are so many in the air flying around?
Why havent they landed and taken us over? If their intelligence and
technology are more advanced than ours, they would have probably
taken us over a long time ago. Or why would aliens have to endlessly
observe our activity? If these aliens are as advanced as some people
think, for them to just keep observing us for centuries or decades would
be a complete waste of time and utterly stupid. Besides, the aliens would
have detection devices that would have told them everything they needed
to know a long time ago. If these super-advanced aliens are material
beings, how could they instantly disappear and pass through walls and
the ground, as they are said to do?

UFOs 21

Numerous people have claimed to have been contacted by aliens and

even taken aboard their spacecrafts. Yet no one has ever recovered a
spacecraft, or the undisputed fragments of a spacecraft. A UFO skeptic
who has spent 28 years investigating UFO cases put it this way: why
is it that not one person who claims to have been aboard a flying saucer
has ever brought back a paper clip or cigarette lighter or some other
souvenir? 155
No one has ever recovered even one alien, or even one piece of wreckage
from an alien craft. Another skeptic of aliens said: One would think
that if aliens actually desired to make contact, that we would have at least
one rock-solid, irrefutable close encounter of the third kind, complete
with credible eyewitnesses and proof to substantiate it beyond question,
before the military could seal off the area and deny everything. 156 The
fact is that there has never been even one undeniable encounter with an
alien that has ever been even reasonably verified. The reason for this is
that there are no aliens. Its all a highly sophisticated demonic deception.
All these facts prove that UFOs and aliens are a spiritual phenomenon,
not a material phenomenon. They come from another dimension, the
spirit world right around us. UFOs are indeed false apparitions produced
by Satan, the prince of the power of the air (Ephesians 2:2).
Is It Theologically Possible for Other Alien Races to Exist in
Addition to the Human Race?
The Bible doesnt mention ETs. In Romans 5:12, it states: through one
man [Adam] sin entered the world. The original Greek word for world
is kosmos, which is usually translated to mean the entire universe. Also,
Romans 8:22 says: the whole creation groans and suffers as a result of
Adams sin. This would mean that alien beings in outer space which
didnt descend from Adam, would still suffer because of his original sin,
which makes no sense. Jesus died once on planet Earth to save the sons
of Adam, and He will not die again (Romans 6:9-10; 1 Peter 3:18).
Biblical Verses on the Powers of Angels & Fallen Angels
In the Bible, we see many supernatural objects described as wheels,
pillars, thrones, chariots, lights and fire, balls of fire, flaming chariots,
spinning objects, dark objects and shining clouds. These descriptions
bear a striking resemblance to modern UFOs. These objects in the Bible
22

UFOs

have been described doing things such as flying, hovering, ascending,

and descending.
In Exodus chapters 7 & 8, we see that God allowed the Devil to imitate
some of Gods wonders in order to deceive people. The Devil was
allowed to cause physical changes to material things.
Angels are very powerful spiritual beings that can:
1) Interact with people and even kill them
(Genesis 16:9; Genesis 19:15; 1 Chronicles 21:15; John 20:12)
2) Appear as normal human beings, undetected by people
(Genesis 19:1-17; Hebrews 13:2)
3) Appear as bright, shining beings that terrify those who see them
(Matthew 28:2-5)
4) Appear in peoples dreams (Matthew 1:20-24)
5) Appear to animals, but invisible to humans (Numbers 22:21-27)
Angels can take on many different forms and shapes. It would make
sense, therefore, that if demons are fallen angels, they possess many of
the same abilities which the faithful and good angels described in the
Bible possess. Many people who say they have encountered aliens have
described a reptilian likeness to their appearance. Its very interesting
that when Satan appeared to Eve in the Garden of Eden, he appeared in
the form of a snake (a reptile). A snake may be the first thing that comes
to a persons mind when you hear the word reptile.
End of sample. Copyright 2008: Bro. Michael Dimond of Most Holy Family Monastery. All rights
reserved.
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UFOs 23

mars
REPTOID PREDATION OF HUMANOIDS FOUND ON MARS
IN TWO PHOTOGRAPHS FROM THE NASA ROVER SPIRIT
SEATTLE, WA (MARS) April 12, 2009 Four months after establishing that MARS is an inhabited
planet, the lawyer from Washington State who discovered life on Mars has found evidence in two NASA
photographs that reptoid predation of humanoids is taking place on the surface of the Red Planet.

12/12/2008: The day that life on Mars was proved

th

On December 12 of last year, Andrew D.

Basiago, 47, founder and president of the Mars
Anomaly Research Society, published his
landmark paper The Discovery of Life on Mars.
The paper is an analysis of a single NASA
photograph, PIA10214, which is a westward view
taken by the rover Spirit from the western edge of
the Home Plate Plateau in the Columbia Basin of
the Gusev Crater of Mars in November 2007 that
was beamed back to Earth in January 2008. It
was in PIA10214 that Basiago found the first
dispositive evidence of life on Mars.

Andrew D. Basiago

Since then, Basiago, who is leading the campaign to have the US government release its secret
documents pertaining to Mars, has carefully studied other photographs taken by Spirit from different parts
of the Home Plate Plateau. These photographs include PIA11049, which is a southward view of the
Columbia Basin taken by Spirit from the southern edge of the plateau during the first six months of 2008.

NASA image PIA10214

NASA image PIA11049

According to Basiago, both of these NASA images reveal a pattern of predation by reptoid species of
humanoid species on Mars. It shocks our sensibilities as human beings on this planet, he said. But the
data being retrieved by Spirit shows that on Mars, humanoids are not at the top of the food chain.

Mars Anomaly Research Society | P.O. Box 2311 | Vancouver, WA 98668 | U.S.A.

mars
Reptoid predation of humanoids on Mars, page 2

Reptoid predation in NASA image PIA10214

Basiago isolated the first evidence of reptoid predation of humanoids on Mars from PIA10214. It is
located near the site identified in his paper The Discovery of Life on Mars as The Rock Garden.
In the photograph below, The Rock Garden, with its defining feature The Grotesque Humanoid Skull, can
be seen occupying the lower right corner of the picture. In the upper left corner of this image, a reptile,
resembling a turtle without a shell, can be seen consuming a humanoid being, legs first.

A reptoid consumes a humanoid near The Rock Garden in the Columbia Basin of Mars

In this close-up (right). the reptoid, with a set of

long tentacles, can be seen chewing on the
remains of one humanoid, as the lower
extremities of a second humanoid being,
consisting of a pair of legs and a bloody stump
where the rest of its body should be, lie to the
right of the first humanoid being. This finding
will test the limits of the Prime Directive, the
discoverer of life on Mars observed. Should we
let humanoids perish as the food for reptoids on
Mars to preserve the balance of nature there, or
should we intervene to save humanoids from
predation because it is the ethical thing to do?

Close-up: A reptoid consumes a humanoid

Mars Anomaly Research Society | P.O. Box 2311 | Vancouver, WA 98668 | U.S.A.

mars
Reptoid predation of humanoids on Mars, page 3

Reptoid predation in NASA image PIA11049

Corroborating evidence of reptoid predation of humanoid beings on Mars was found by Basiago in NASA
image PIA11049. There, at the far southern reaches of the Columbia Basin, the head of a green reptile
lurking behind its human quarry can be seen. The humanoid was dressed in turquoise blue clothing at
the time that he became the reptoids next meal. Shortly before he became lunch, the lower parts of his
legs were bitten off. Apparently, to the reptoids on Mars, the feet of humanoids are considered a delicacy
or perhaps this technique is used by the reptiles to immobilize human prey so they can be consumed.

Reptoid predator lurks behind its prey

Close-up: Human prey with feet bitten off

So plesiosaurus a carnivore
This predation of humanoid beings by reptile species on Mars might include the plesiosaurs that Basiago
discovered on Mars and wrote about in The Discovery of Life on Mars. Plesiosaurs are ubiquitous on
Mars but they did not survive the K-T extinction on Earth, where they were carnivorous by nature.

Plesiosaurs on Mars
Mars Anomaly Research Society | P.O. Box 2311 | Vancouver, WA 98668 | U.S.A.

mars
Reptoid predation of humanoids on Mars, page 4

Below, we see two men, half like beetles, pitch forward to confront a plesiosaur (left), as serpents slither
across the ridge (center) and tiny humanoids clamber onto a rock seeking safety (right). These scenes
show that on Mars, dire conditions make predation between competing life forms very intense.

Two men confront a plesiosaur

Snakes slither across ridge

Humanoids clamber onto rock

It may be that reptoid predation is the cause of the thousands of humanoid bodies littering the valley seen
in PIA10214, as if they are in a mass grave or killing field, where they are now fructifying in a sticky paste
(left). These corpses can also be seen on the Home Plate Plateau itself, deposited in great slag heaps
(center). In some parts of PIA10214, serpents can be seen slithering across the bodies (right).

Humanoid bodies

A great slag heap of corpses

Snake slithers across bodies

This evidence does not exist in a vacuum. Rather, these humanoid corpses are deposited by the
thousands in the valley bottom immediately below the statue that Basiago has called The Beings on the
Edge. He has shown that this statue does not consist merely of a female figure in a blue dress but is part
of a statuary complex that includes a male figure and an elaborate rock carving of a reptilian form biting
the neck of a white male human being (below), perhaps indicating that it is an altar of human sacrifice.

Mars Anomaly Research Society | P.O. Box 2311 | Vancouver, WA 98668 | U.S.A.

mars
Reptoid predation of humanoids on Mars, page 5

Reptoid predation on Mars is serious business.

When one considers the evidence of a reptilian
scenario on Mars that PIA10214 contains
archaeological evidence of a possible altar of
reptoid sacrifice of humanoids on Mars (right);
that forensic evidence has been found in the
form of thousands of humanoid bodies littering
the western littoral of the Home Plate Plateau;
and that biological evidence now exists that
reptoid predation of humanoid beings is quite
commonplace on Mars the data Basiago has
collected from NASAs images may confirm that
there is a reptilian agenda to take over not just
Earth, as posited by David Icke, but the entire
solar system. Thus, Basiagos Mars findings
may be of interplanetary security significance.

A statue with a reptile biting a human

Mars Anomaly Research Society | P.O. Box 2311 | Vancouver, WA 98668 | U.S.A.

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/012345/0203)6271839/

HMG

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/012345/0203)6271839/

HMI

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/012345/0203)6271839/

HMK

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/012345/0203)6271839/

HMM

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Tokita et al. SpringerPlus 2013, 2:336

http://www.springerplus.com/content/2/1/336

a SpringerOpen Journal

RESEARCH

Open Access

Skeletal gene expression in the temporal region

of the reptilian embryos: implications for the
evolution of reptilian skull morphology
Masayoshi Tokita1,3*, Win Chaeychomsri2 and Jindawan Siruntawineti2

Abstract
Reptiles have achieved highly diverse morphological and physiological traits that allow them to exploit various
ecological niches and resources. Morphology of the temporal region of the reptilian skull is highly diverse and
historically it has been treated as an important character for classifying reptiles and has helped us understand the
ecology and physiology of each species. However, the developmental mechanism that generates diversity of
reptilian skull morphology is poorly understood. We reveal a potential developmental basis that generates
morphological diversity in the temporal region of the reptilian skull by performing a comparative analysis of gene
expression in the embryos of reptile species with different skull morphology. By investigating genes known to
regulate early osteoblast development, we find dorsoventrally broadened unique expression of the early osteoblast
marker, Runx2, in the temporal region of the head of turtle embryos that do not form temporal fenestrae. We also
observe that Msx2 is also uniquely expressed in the mesenchymal cells distributed at the temporal region of the
head of turtle embryos. Furthermore, through comparison of gene expression pattern in the embryos of turtle,
crocodile, and snake species, we find a possible correlation between the spatial patterns of Runx2 and Msx2
expression in cranial mesenchymal cells and skull morphology of each reptilian lineage. Regulatory modifications of
Runx2 and Msx2 expression in osteogenic mesenchymal precursor cells are likely involved in generating
morphological diversity in the temporal region of the reptilian skull.
Keywords: Reptiles; Skull; Morphology; Development; Osteogenesis; Heterotopy

Introduction
Amniotes (Amniota) consist of two large groups of
tetrapod vertebrates, Synapsida and Reptilia, that
diverged from one another over 300 million years
ago (Ma) (Carroll, 1988, Modesto & Anderson, 2004,
Benton, 2005). The synapsids are represented today
by mammals while reptiles by extant turtles, tuatara,
lizards, snakes, crocodiles, birds, and their extinct
relatives, including dinosaurs and pterosaurs. Over
time, reptiles have evolved highly diverse morphological and physiological traits that allow them to exploit various ecological niches and resources on the
land, in water, and in the air.

* Correspondence: tokky@biol.tsukuba.ac.jp
1
Graduate School of Life and Environmental Sciences, University of Tsukuba,
Tenno-dai 1-1-1, Tsukuba, Ibaraki 305-8572, Japan
Full list of author information is available at the end of the article

Morphology of the skull of reptiles, especially the temporal region is highly diverse (Figure 1). This morphological diversity observed in the temporal region is broadly
categorized into three architectural patterns. The anapsid
condition of the skull is observed in basal amniotes such
as Scutosaurus and Captorhinus, and in turtles. In these
animals, the temporal region of the skull is completely
roofed by bones, without temporal openings (fenestrae).
The synapsid condition of the skull is recognized in ancestral lineages of extant mammals and is characterized by the
presence of a temporal fenestra at lower position of either
side of the skull. The diapsid condition of the skull is seen
in "non-turtle" extant reptiles and in their extinct relatives.
In these animals, two temporal fenestrae exist on either
side of the skull. In reptiles with fully diapsid skulls, the
upper temporal fenestra is dorsally bordered by the parietal
bone, anteriorly by the postorbital bone, and posteriorly by
the squamosal bone. The lower temporal fenestra is

2013 Tokita et al.; licensee Springer. This is an Open Access article distributed under the terms of the Creative Commons
Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction
in any medium, provided the original work is properly cited.

Tokita et al. SpringerPlus 2013, 2:336

http://www.springerplus.com/content/2/1/336

dorsally surrounded by both the postorbital and squamosal

bones, and ventrally by both the jugal and quadratojugal
bones. These temporal fenestrae are thought to have
evolved to allow space for accommodating the enlarged
jaw-closing muscles that enable powerful biting, or to
minimize stresses exerted by the contraction of jaw muscles on the skull, or to reduce the weight of the skull
itself (Frazzetta, 1968, Carroll, 1982, Rieppel, 1993a, Benton,
2005). During the course of diapsid evolution skull morphology has been rearranged repeatedly, resulting in a variety
of modified patterns (Rieppel, 1993a, Rieppel & Gronowski,
1981, Mller, 2003, Moazen et al., 2009). Among extant
reptiles, fully diapsid skull is only seen in tuatara and crocodiles. Because the lower temporal bar that encloses the
lower temporal fenestra ventrally is regarded to be lost once
in the common ancestor of lepidosaurs (lizards, snakes,
tuatara) and archosaurs (crocodiles and birds) (Rieppel,
1993a, Mller, 2003), these reptilian lineages possibly acquired the lower temporal bar secondarily (Mller, 2003,
Moazen et al., 2009). Both snakes and birds have lost the
upper temporal bar so that their temporal region is free
from any bony frames (Pough et al., 2005).
In reptiles, phylogenetic position of turtles is highly
controversial. Traditionally, turtles have been regarded
as the only surviving clade of stem reptiles based on the
pattern of their skull morphology: an anapsid skull
whose temporal region is completely roofed with bones
(Williston, 1917, Gregory, 1946, Romer, 1968, Gaffney,
1980, Reisz & Laurin, 1991, Lee, 1993, Laurin & Reisz,
1995, Lee, 1996, 1997, Reisz, 1997, Lee, 2001). However,
recent comprehensive analysis of morphological traits
(Rieppel & deBraga, 1996, deBraga & Rieppel, 1997,
Rieppel, 2000, Hill, 2005, Li et al., 2008, but see Lyson
et al., 2010, 2013 for opposed conclusion) and molecular
phylogenetic studies (Hedges & Poling, 1999, Kumazawa
& Nishida, 1999, Iwabe et al., 2005, Hugall et al., 2007,
Shedlock et al., 2007, Shen et al., 2011, Tzika et al., 2011,
Chiari et al., 2012, Crawford et al., 2012, Fong et al.,
2012, Lyson et al., 2012, Wang et al., 2013) suggest that
there is a close relationship of turtles to diapsid reptiles,
implying that the temporal fenestrae were secondarily
closed in turtles. In this study, we employ the hypothesis
that turtles are descendent of diapsid reptiles.
Although skull morphology has been regarded as an important character in classification of reptiles and in understanding the ecological and physiological aspects of each
reptilian species, the developmental mechanism underlying
diversification of reptilian skull morphology is poorly
understood (Rieppel, 1993a, Evans, 2008). As a consequence, a general genetic and developmental model of reptile skull diversity does not yet exist. In this paper, we test
the hypothesis that changes of skeletal gene expression patterns cause diversification of reptilian skull morphology
through comparative analyses of gene expression in the

Page 2 of 20

Figure 1 Simplified phylogeny of the Reptilia highlighting

diversity of their skull morphology. Paleontological evidence
suggests that all reptiles, including extant lizards, snakes, tuatara,
crocodiles, birds, and turtles, were derived from ancestor whose
temporal region was completely roofed by bone. Earliest diapsid reptiles
such as Petrolacosaurus (Araeoscelidia) acquired two temporal openings
(fenestrae) on either side of the skull (red vertical bar). Recent molecular
phylogenies indicate that turtles (Testudines) were derived from diapsid
ancestor, which would require secondary closure of temporal fenestrae
(green vertical bar). The bone surrounding anteroventral border of the
upper temporal fenestra and anterodorsal border of the lower temporal
fenestra in diapsids: the postorbital was colored in pink. The bone
surrounding posteroventral border of the upper temporal fenestra and
posterodorsal border of the lower temporal fenestra in diapsids: the
squamosal was colored in blue. The bone surrounding the anteroventral
margin of the lower temporal fenestra in diapsids: the jugal was colored
in yellow. Lizards do not have the lower temporal bar. Both upper and
lower temporal bars are absent in snakes and birds. Other extinct
diapsid lineages such as Ichthyosauria and Sauropterygia were not
included in phylogeny for simplicity.

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embryos of representative reptilian species and reveal a potential developmental basis underlying reptilian skull evolution. First, we describe the pattern of early phases of
cranial morphogenesis in a crocodile species with both
upper and lower temporal bars surrounding temporal fenestrae, using molecular markers specific for musculoskeletal tissue precursors. Then, we compare these data with
cranial morphogenesis in a turtle species. We found a
broader expression of the early osteogenic genes, Runx2
and Msx2 in the mesenchymal cells at the temporal region
of turtle embryos, compared to that in crocodile embryos.
Finally, to obtain a broader picture of reptilian skull
morphogenesis, we examined expression patterns of
Runx2 and Msx2 in cranial morphogenesis of a snake
species without temporal bars on the skull and compared with the patterns in crocodile and turtle embryos.
Our findings suggest that there is a possible correlation
between the expression patterns of Runx2 and Msx2
and the architectural pattern seen in the temporal
region of the reptilian skull.

Results
In previous studies in which cranial osteogenesis of reptilian embryos was described, whole-mount clearing and
staining with Alizarin red was used to detect mineralization of intramembranous bones that comprise the
dermatocranium (Kamal et al., 1970, Haluska & Alberch,
1983, Rieppel, 1993b, Rieppel, 1993c, Kuratani, 1999,
Rieppel & Zaher, 2001, Sheil, 2003, Sheil, 2005, Boughner
et al., 2007, Vickaryous & Hall, 2008, Snchez-Villagra
et al., 2009, Werneburg et al., 2009). However, this method
is unable to identify the distribution of the precursor cells
of bones: osteoblasts, as reported by others (Kerney et al.,
2010). To overcome this, we conducted section in situ
hybridization analysis, which labels tissues located deep
inside of the embryonic body and is effective for detecting
tissue-specific domains of expression. We used a probe to
Runx2, which is a molecular marker for osteogenic mesenchymal precursor cells (Ducy et al., 1997, Bobola et al.,
2003, Abzhanov et al., 2007, Han et al., 2007, Kerney et al.,
2010) and described its expression pattern in the temporal
region of reptilian embryos where mineralization of bones
has not been initiated. Furthermore, to describe distribution pattern of "non-osteoblast" cell lineages relative to
that of osteoblasts in the cranial tissue of the embryos, we
also examined expression of other tissue-specific markers:
MyoD for skeletal muscle precursors (Hacker & Guthrie,
1998, Noden et al., 1999), Sox9 for cartilage precursors
(Wright et al., 1995, Bell et al., 1997), Scleraxis (Scx) and
Six2 for precursor of connective tissues, including ligaments and tendons (Oliver et al., 1995, Schweitzer, 2001,
Dreyer et al., 2004, Edom-Vovard & Duprez, 2004,
Schweitzer et al., 2010).

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Differential expression of early osteoblast marker, Runx2,

in the head of crocodile and turtle embryos

Osteogenic mesenchymal precursor cells that express

Runx2 are first detected in the temporal region of crocodile
embryos at stage 14 (Additional file 1) and an almost
identical pattern of Runx2 expression was observed in a
subsequent embryonic stage (stage 15; Figure 2A). These
Runx2-positive cells were localized at the domain dorsal to
the oral cavity where the ventral part of the braincase and
future palatine and pterygoid bones develop, as well as in a
limited domain dorsolateral to the orbit where the future
dorsal projection of the postorbital bone forms (Figure 2B).
We also detected a population of Runx2-positive cells at
the domain ventrolateral to the orbit where future jugal
bone and ventral projection of postorbital bone are formed.
At this stage, the precursor of the jugal and ventral projection of the postorbital were dorsoventrally continued as a
layer of cells but it was thin mediolaterally, especially at the
middle part. In the posterior part of the head, we observe a
population of Runx2-positive mesenchymal cells that later
differentiate into the main body of the postorbital bone
(Figures 2E and 2H). In these stages of crocodile embryogenesis, jaw muscle precursors that were derived from cranial mesoderm migrated to the first pharyngeal arch and
expressed MyoD was clearly detected at the central domain
of the jaw primordia (Figures 2C and 2F; Additional file 1).
Expression of Sox9 was detected at cartilage precursors
that later differentiate into quadrate and Meckel's cartilages
at the domain ventral to jaw muscle precursor, as well as in
chondrocytes that form the future braincase (Figures 2D
and 2G). Expression of Scx was detected in tendon precursor cells that are distributed within the primordia of the
jaw muscles and in the connective tissue within the eye
muscles (Additional file 1). Expression of Six2 was somewhat broader than that of other markers, expressed in
mesenchymal cells surrounding the eyes, cartilaginous precursors of the braincase, quadrate, and Meckel's, as well as
in the mesenchyme at the interface between muscle precursors and the skeletal tissues to which the muscles attach
and in a population of the mesenchyme that dorsally surrounds the brain (Additional file 1).
In crocodile embryos at stage 17 where none of the
dermatocranial elements were positive for Alizarin red in
previous studies (Rieppel, 1993b, Vickaryous & Hall, 2008)
(Figure 3A), we could detect Runx2 expression in the cell
populations that were localized to the area where the future dermatocranium differentiates (Figure 3B). Although
the domain where Runx2-positive cells were populated
was almost identical to that in previous stages, the boundary of each precursor of the dermatocranial elements became clearer. Although differentiation of the parietal bone
is delayed compared to other dermatocranial elements as
described previously (Rieppel 1993b, Vickaryous & Hall,
2008), a pair of precursors of the parietal were recognized

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Figure 2 Expression of musculoskeletal tissue marker genes in the head of crocodile embryos at stage 15. (A) Lateral view of the head
of a crocodile embryo at stage 15. (B-D) Frontal sections prepared around the plane indicated by the red line in (A). (E-G) Frontal sections
prepared around the plane indicated by the blue line in (A). (H) Frontal sections prepared around the plane indicated by the green line in (A).
(B, E, and H) Runx2-positive mesenchymal cells are observed at the location where future dermatocranial elements are developed. (C and F)
Expression of MyoD is detected at the cranial muscular tissues. (D and G) Cartilaginous tissues, including the braincase, quadrate, and Meckel's,
are clearly labeled by Sox9 probe. The red outlined domains in (B, D, E, and G) indicate the location of the pseudotemporal muscle (ptm)
deduced from adjacent sections where muscular tissues are labeled by MyoD probe. Scale bar in (A) is 1 mm. Scale bars in (B-H) are 0.5 mm.

as Runx2-positive cell aggregation at the domain dorsolateral to the cartilaginous braincase (Figures 3B and 3E).
We could detect Runx2-negative cell populations between
the nascent parietal precursor and the morphologically
more developed postorbital precursor and also between
precursors of the postorbital and quadratojugal located
lateral to Sox9-positive quadrate cartilage (Figure 3E). In
this stage, MyoD was expressed in differentiated jaw and
eye muscles (Figures 3C and 3F) and Sox9 was expressed
in differentiated chondrocranium and splanchnocranium
components, including the braincase, quadrate, and
Meckel's (Figures 3D and 3G). Expression of Scx was
detected in tendinous tissues accompanying MyoD-positive muscles as in previous stages (Figure 3H). Expression

of Six2 was detected in the mesenchyme localized around

the jaw articulation between quadrate and Meckel's, as
well as in adjacent mesenchyme of the braincase, postorbital bone, and jaw muscles (Figure 3I).
Next, we examined cranial morphogenesis of turtles that
have an anapsid skull, using the same method to identify
the distribution pattern of precursors of each tissue that
constitutes the cranial musculoskeletal system. In turtle
embryos at stage 14 (Additional file 2) that correspond to
crocodile embryos at stage 14 in external morphology, and
in turtle embryos at stage 15 (Figure 4A) that are comparable
to crocodile embryos at stage 15, we observed almost identical patterns of expression for each gene. We detected MyoD
expression specifically in the primordia of jaw adductor

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Figure 3 Expression of musculoskeletal tissue marker genes in the head of crocodile embryos at stage 17. (A) Lateral view of the head
of a crocodile embryo at stage 17. (B-D) Frontal sections prepared around the plane indicated by the red line in (A). (E-I) Frontal sections
prepared around the plane indicated by the blue line in (A). (B and E) Expression of Runx2 is more concentrated to the precursors of
dermatocranial elements, compared to the previous stages. (C, D, F, and G) Cranial muscular and cartilaginous tissues are clearly labeled by
MyoD and Sox9 probes, respectively. (H and I) Expression domains of Scx and Six2 are indicated by arrowheads. The former is expressed in
tendinous tissues accompanying cranial muscles and the latter is expressed mainly in connective tissue cells associated with cartilages of the jaw
and the braincase. The red outlined domains in (B, D, E, G, H, and I) indicate the location of the pseudotemporal muscle deduced from
adjacent sections where muscular tissues are labeled by MyoD probe. Green line in (A) indicates the plane where sections given in Figure 7D, E
and F were prepared. Scale bar in (A) is 1 mm. Scale bars in (B-I) are 0.5 mm.

and eye muscles (Figures 4C and 4H; Additional file 2)

and Sox9 expression in precursor cells of the braincase,
quadrate, and Meckel's cartilages (Figures 4D and 4I;
Additional file 2) as in stage-matched crocodile embryos.
Expression of Scx was first detected in a layer of mesenchymal cells that was located at the periphery of the jaw
adductor and eye muscle precursors in stage 15 turtle
embryos (Figure 4E). Expression of Six2 was observed
in the mesenchyme surrounding the eye and adjacent

mesenchyme of the braincase and jaw cartilages, as well as

in some mesenchymal cells within jaw muscle precursors
(Figure 4F; see Additional file 2), as in stage-matched
crocodile embryos. Interestingly, we observed expression
of the early osteoblast marker, Runx2, in a broader domain
at the temporal region of the head of turtle embryos, compared to that in stage-matched crocodile embryos. In
stage14 turtle embryos, Runx2 expression was detected in
a population of cells medial to the precursor of the jaw

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Figure 4 Expression of musculoskeletal tissue marker genes in the head of turtle embryos at stage 15. (A) Lateral view of the head of a
turtle embryo at stage 15. (B-F) Frontal sections prepared around the plane indicated by the red line in (A). (G-I) Frontal sections prepared
around the plane indicated by the blue line in (A). (B, G) Runx2-positive mesenchymal cells (arrows) are broadly distributed at lateral portion of
the head, from the top of the head to the ventral margin of the first pharyngeal arch. (C and H) Cranial muscular tissues are clearly labeled by
MyoD probe. (D and I) Cartilaginous tissues, including the braincase and quadrate, are clearly labeled by Sox9 probe. (E) Scx is expressed in
tendon primordia accompanying cranial muscles (arrowheads). (F) Six2 is expressed mainly in connective tissue cells associated with the
braincase cartilage and jaw adductor muscle (arrowheads). Note that the anlage of jaw adductor muscle (red outlined domain) is covered by a
thick layer of Runx2-positive mesenchyme laterally. Scale bar in (A) is 1 mm. Scale bars in (B-I) are 0.5 mm.

adductor muscles and the mesenchyme localized at

the domain dorsolateral and ventrolateral to the orbit
(Additional file 2). The domain of Runx2 expression became further expanded in the head of turtle embryos at
stage 15. A thick layer of the mesenchymal cells that express Runx2 completely covered the brain and the precursor of jaw adductor muscle laterally (Figures 4B and 4G).
In turtle embryos at stage 17 (Figure 5A) that correspond to crocodile embryos at stage 17 in overall morphology, we observed MyoD expression in differentiating
cranial muscles, including external adductor muscles
(Figure 5C) and Sox9 expression in the cartilaginous
tissues that constitute the braincase, quadrate, and
Meckel's (Figure 5D). Scx was specifically expressed in
tendinous tissues at the periphery of jaw adductor muscles, as well as in the precursor of the bodenaponeurosis

(central tendon of external adductor) just appeared within

the jaw adductor muscular tissue (Figure 5E). The expression domain of Six2 was broader in the temporal region of
the head compared to that of Scx, diffusively expressed in
the mesenchymal cells surrounding jaw adductor muscles,
braincase, and jaw cartilages (Figure 5F). A thick layer of
Runx2-positive mesenchymal cells that surrounds the
braincase and jaw adductor muscle laterally was observed
(Figure 5B). Runx2-expressing mesenchyme was also distributed around the quadrate cartilage and the ventral part
of the braincase.
Expression of potential upstream osteogenic regulatory
genes in the head of crocodile and turtle embryos

Through comparative analysis of expression patterns of

tissue-specific marker genes, we noticed a difference in

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Figure 5 Expression of musculoskeletal tissue marker genes in the head of turtle embryos at stage 17. (A) Lateral view of the head of a
turtle embryo at stage 17. (B-F) Frontal sections prepared around the plane indicated by the blue line in (A). (B) Runx2-positive mesenchymal cells
(arrows) are broadly distributed at the lateral portion of the head, from the top of the head to the ventral margin of the first pharyngeal arch derivative.
(C) Cranial muscular tissues are clearly labeled by MyoD probe. (D) Cartilaginous tissues, including the braincase and quadrate, are clearly labeled by
Sox9 probe. (E) Scx is expressed in tendinous tissues at the periphery of external adductor muscle (ame) (arrowheads) and in the bodenaponeurosis
(boap. central tendon of jaw adductor muscle). (F) Six2 is expressed mainly in connective tissue cells associated with cartilages of the braincase and
quadrate, as well as in connective tissue cells within jaw muscles (arrowheads). Note that the external adductor muscle (red outlined domain) is
covered by a thick layer of Runx2-positive mesenchyme laterally. Scale bar in (A) is 1 mm. Scale bars in (B-F) are 0.5 mm.

the spatial pattern of expression of the early osteoblast

marker, Runx2 in the head of crocodile and turtle embryos. To reveal potential mechanisms that account for
such differential distribution of osteogenic mesenchymal
precursor cells between two reptilian lineages with or
without temporal fenestrae, we next examined expression patterns of some candidate genes that are known to
regulate cranial osteogenesis. In the present study, we
focused on Bmp4, Msx1, and Msx2. Bmp4 is a signaling
molecule and plays a key role in the Bmp signaling pathway. Because exogenous Bmp4 increases tissue volume
in calvarial bone tissue culture, this protein is considered
to be involved in calvarial bone growth (Kim et al., 1998,
Rice et al., 2003). Both Msx1 and Msx2 are members of
the muscle segment homeobox (msh) gene family of
transcription factors and both loss-of- and gain-of-function analyses of these genes suggest their essential roles
in vertebrate cranial osteogenesis (Satokata & Maas,
1994, Satokata et al., 2000).
In the present analysis, we found that Bmp4 and Msx1
showed almost identical expression patterns through
cranial osteogenesis in crocodile and turtle embryos. In
crocodile embryos we examined (through stage 14 to
stage 17), Bmp4 was strongly expressed in the epithelium of cochlear canal, the mesenchyme surrounding the
eye, the mesenchyme distributed in the medial part of
jaw primordia, the precursors of the palatine bones, and
a population of mesenchymal cells that covered the
brain dorsally (Figure 6A; Figures 7A and 7D). In turtle
embryos we examined (through stage 14 to stage 17),
Bmp4 was expressed in a spatially limited domain:
the epitthelium of cochlear canal, the mesenchyme

dorsolateral and ventrolateral to the eye and a limited

population of the mesenchyme in close proximity of the
jaw articulation (Figure 6G; Figure 7G). We observed
Msx1 expression in the epithelium of the cochlear canal,
the mesenchyme that occupies the domain close to the
jaw articulation and lateral to the quadrate and Meckel's
cartilages, and a thin layer of mesenchymal cells that
covers the brain dorsally in crocodile embryos examined
(Figure 6B; Figures 7B and 7E). In turtle embryos, Msx1
was expressed in the epithelium of the cochlear canal,
the mesenchyme distributed around the jaw articulation
and lateral to quadrate and Meckel's cartilages, as well
as in the mesenchyme that populates the domain dorsal
to the eye (Figure 6H; Figure 7H).
In contrast to Bmp4 and Msx1, we detected differential expression patterns of Msx2 in the head of crocodile and turtle embryos. In crocodile embryos at stage
14 and 15, Msx2 was expressed in a thin layer of mesenchymal cells surrounding the dorsal aspect of the brain
(Figures 6C and 6E). In the posterior part of the head, the
ventral edge of this Msx2-expressing cell population is located dorsal to the eye. In these crocodilian embryos,
Msx2 expression was also observed in a population of the
mesenchyme that occupied the domain between the ventrolateral part of quadrate cartilage and surface epidermis
(Figures 6C and 6E). These mesenchymal cells expressed
Msx1 as well (Figure 6B) and appeared to differentiate into
the quadratojugal bone later. In crocodile embryos at stage
17, specific expression of Msx2 was detected at a population
of mesenchymal cells in close proximity of Runx2-expressing
precursors of postorbital and quadratojugal bones, as well
as in a thin layer of the mesenchyme surrounding the

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Figure 6 Expression of Bmp4, Msx1, and Msx2 in crocodile and turtle embryos at stage 15. (A-D) Frontal sections prepared around the
plane indicated by the red line in Figure 2A. (E, F) Frontal sections prepared around the plane indicated by the blue line in Figure 2A. (G-J)
Frontal sections prepared around the plane indicated by the red line in Figure 4A. (K, L) Frontal sections prepared around the plane indicated by
the blue line in Figure 4A. (A and G) In both crocodile and turtle embryos, expression of Bmp4 is detected at the mesenchyme distributed in
medial part of the jaw primordia (arrows). (B and H) Expression of Msx1 is detected at the mesenchyme that occupies the domain close to jaw
articulation and lateral to quadrate and Meckel's cartilages (arrows). (C and E) In crocodile embryos, Msx2 is expressed in a thin layer of
mesenchymal cells surrounding dorsal aspect of the brain and in a population of the mesenchyme that occupies the domain between
ventrolateral part of quadrate cartilage and surface epidermis (arrows). (I and K) In turtle embryos, Msx2 is expressed in mesenchymal cells that
populate lateral aspect of the head (arrows). In contrast to the condition in crocodile embryos, the ventral edge of Msx2-expressing mesenchymal
layer is terminated ventral to the eye in turtle embryos and these cells cover MyoD-expressing jaw adductor muscle precursor (J and L) laterally.
The red outlined domains in (A-C, and E) indicate the location of the anlagen of the pseudotemporal muscle deduced from adjacent sections
where muscular tissues are labeled by MyoD probe. The red outlined domains in (G-I, and K) indicate the location of the anlagen of jaw
adductor muscle. Scale bars are 0.5 mm.

brain dorsally where future parietal bones were developed

(Figures 7C and 7F). We observed that the space adjacent
to Msx2-positive precursors of these dermatocranial elements was filled with Msx2-negative mesenchymal cells.
Interestingly, we observed broader expression of Msx2 in
turtle embryos, compared to that in stage-matched crocodile embryos. In turtle embryos examined, Msx2 was
expressed in mesenchymal cells that populate lateral aspect
of the head of embryos (Figures 6I and 6K; Figures 7I and
7J). The ventral edge of the Msx2-expressing mesenchymal
layer was terminated ventral to the eye and these cells

covered MyoD-expressing external adductor muscle laterally. Showing its dorsoventrally broadened expression pattern, the domain of Msx2 expression largely overlapped
with that of Runx2 in turtle embryos (Figures 4B and 4G;
Figure 5B).
Expression of Runx2 and Msx2 in the head of snake embryos

To verify a correlation between the expression patterns

of Runx2 and Msx2 and reptilian skull morphology, we
finally examined expression patterns of these genes, as
well as of marker genes for muscular and cartilaginous

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Figure 7 Expression of Bmp4, Msx1, and Msx2 in crocodile and turtle embryos at stage 17. (A-C) Frontal sections prepared around the
plane indicated by the blue line in Figure 3A. (D-F) Sections prepared around the plane indicated as the green line in Figure 3A. (G-I) Sections
prepared around the plane indicated as the red line in Figure 5A. (J) Section prepared around the plane indicated as the blue line in Figure 5A.
(A, D, and G) In both crocodile and turtle embryos, expression of Bmp4 is detected in the epithelium of the cochlear canal, the mesenchyme
surrounding the eye, and the mesenchyme distributed in medial part of the jaw (arrows). (B, E, and H) Expression of Msx1 is detected at the
mesenchymal cells that later differentiates into quadratojugal bone (qj) and in a thin layer of mesenchymal cells that covers brain dorsally (arrows
at the top of E and H), as well as in the epithelium of the cochlear canal. (C and F) In crocodile embryos, expression of Msx2 is detected at a
population of mesenchymal cells in close proximity of postorbital and quadratojugal bones, as well as in a layer of the mesenchyme surrounding
the brain dorsally where future a pair of parietal bones are developed. (I and J) In turtle embryos, Msx2 was expressed in a thick layer of
mesenchymal cells that populate lateral aspect of the head (arrows). The Msx2-positive mesenchymal cells cover the external adductor muscle
precursors (red outlined domains in G-J) laterally. The red outlined domains in (A-F) indicate the location of the pseudotemporal muscle
deduced from adjacent sections where muscular tissues are labeled by MyoD probe. The blue outlined domains in (A-J) indicate the location of
quadrate cartilage deduced from adjacent sections where cartilaginous tissues are labeled by Sox9 probe. Scale bars are 0.5 mm.

tissues, in cranial morphogenesis of a snake species

where their temporal fenestrae are not encircled by the
temporal bars. In snake embryos at stage 26 (Figure 8A)
that morphologically correspond to crocodile and turtle
embryos at stage 14 or 15, MyoD was expressed in the
primordia of the first arch muscles (Figures 8C and 8G).
Sox9 was strongly expressed in the precursors of quadrate and Meckel's cartilages and the base of the braincase, as well as in a layer of mesenchyme surrounding
the brain laterally (Figures 8D and 8H). In these snake
embryos, early osteoblast marker, Runx2 was expressed

in the mesenchyme that occupied the space medial to

the Sox9-positive quadrate precursor and in a limited
population of mesenchymal cells ventral to the orbit
(Figure 8B). We also detected Runx2 expression in a
layer of mesenchymal cells that surround the brain laterally. In the posterior temporal region, Runx2 was only
faintly expressed in the adjacent mesenchyme of the
Sox9-positive quadrate precursor (Figure 8F). Expression
of Msx2 was detected in the mesenchyme medial to the
precursor of the quadrate (Figure 8E). Its expression domain was spatially overlapped with the domain where

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Figure 8 (See legend on next page.)

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(See figure on previous page.)

Figure 8 Expression of the genes that regulate the development of cranial musculoskeletal tissues in snake embryos. (A) Lateral view of
the head of a snake embryo at stage 26. (B-E) Frontal sections prepared around the plane indicated by the red line in (A). (F-I) Frontal sections
around the blue line in (A). (J) The head of a snake embryo at stage 29. (K-N) Frontal sections around the red line in (J). (O-R) Frontal sections
around the blue line in (J). (S) The head of a snake embryo at stage 31. (T-W) Frontal sections around the red line in (S). (B, F, K, O, and T)
Expression of Runx2. (C, G, L, P, and U) Expression of MyoD. (D, H, M, Q, and V) Expression of Sox9. (E, I, N, R, and W) Expression of Msx2. In
snake embryos, expression of Runx2 and Msx2 are detected in the precursor cells of dermatocranial elements (arrows), as in crocodile and turtle
embryos. However, these mesenchymal cells are not seen in the temporal region lateral to the jaw adductor muscles. Instead, these cells are
distributed in the vicinity of the brain, laterally covering it, and differentiate into the parietal bone in older embryos. Open arrowheads in
(F, K, N, O, R, T, and W) indicate the mesenchyme around Sox9-positive jaw cartilages, where the expression of Runx2 and Msx2 is detected. The
red outlined domains in (O, Q, R, T, V, and W) indicate the location of the external adductor muscle deduced from adjacent sections where
muscular tissues are labeled by MyoD probe. Scale bars in (A), (J), and (S) are 1 mm. Scale bars in other pictures are 0.5 mm.

Runx2-positive cells were distributed, but the former was

narrower. We also detected Msx2 expression in a mesenchymal layer that surrounded the brain dorsally (Figures 8E
and 8I). In snake embryos at stage 29 (Figure 8J), which
morphologically correspond to crocodile and turtle embryos at stage 17, well-differentiated cranial muscular and
cartilaginous tissues were specifically labeled by expression
of MyoD (Figures 8L and 8P) and Sox9 (Figures 8M and
8Q), respectively. We observed spatially overlapped expression of Runx2 and Msx2 in these embryos: both genes
were expressed in the precursors of palatine and pterygoid
bones medial to Sox9-positive jaw cartilages and in mesenchymal cells accompanying jaw cartilages, as well as in a
layer of loose mesenchyme that later forms a precursor of
parietal bones that cover the dorsal part of the brain
(Figures 8K, 8N, 8O, and 8R). However, only Runx2 was
expressed in the mesenchyme that populated the domain
ventral to the orbit, which possibly corresponds to precursors of the maxilla bones of the upper jaw. No Runx2 and/
or Msx2 expressing osteogenic mesenchymal precursor
cells populated the domain lateral to jaw adductor muscles.
In more developed snake embryos (at stage 31; Figure 8S),
both Runx2 and Msx2 were specifically expressed in the
precursors of the dermatocranial elements, including the
parietal that encases the brain dorsally (Figures 8T and
8W). As in previous stages, expression of the former was
more expanded. The lateral aspect of the external adductor
muscles was never covered by the skeletal tissues that express Runx2 and/or Msx2. The results on expression domains of each gene analyzed in crocodile, turtle, and snake
embryos are summarized in Table 1.

Discussion
Potential developmental basis of anapsid skull in turtles

Skull morphology, especially the osteological configuration of the temporal region, has historically been
treated as the most important character in the classification of major lineages of reptiles. Based on their anapsid
skull, turtles have been regarded as a sole descendent of
stem reptiles (Williston, 1917, Gregory, 1946, Romer,
1968, Gaffney, 1980, Reisz & Laurin, 1991, Lee, 1993;
Laurin & Reisz, 1995, Lee, 1996, 1997, Reisz, 1997, Lee,

2001) despite the contrary argument that turtles were

derived from an ancestor with a diapsid skull (Lakjer,
1926, Goodrich, 1930). Recent phylogenetic studies
where the interrelationships of both extant and extinct
reptiles were surveyed through comprehensive analysis
of multiple osteological traits concluded that turtles
were closely related to lepidosaurian diapsids (Rieppel &
deBraga, 1996, deBraga & Rieppel 1997, Rieppel 2000,
Hill, 2005, Li et al., 2008). Furthermore, results of molecular phylogenetic studies have strongly suggested
diapsid affinity of turtles (Hedges & Poling, 1999,
Kumazawa & Nishida, 1999, Iwabe et al., 2005, Hugall
et al., 2007, Shedlock et al., 2007, Shen et al. 2011, Tzika
et al., 2011, Chiari et al., 2012, Crawford et al., 2012,
Fong et al., 2012, Lyson et al., 2012, Wang et al., 2013).
If turtles were derived from a diapsid ancestor, then the
anapsid skull of turtles evolved independently from that
of ancestral lineages of reptiles by secondary closure of
the temporal fenestrae. However, although the phylogenetic position of turtles within amniotes still remains inconclusive (Lyson et al., 2010, 2013, Kuratani et al.,
2011), there has been no study in which the process of
development of their anapsid skull is described with molecular markers for labeling precursor cells of the
dermatocranium. In the present study, we examined
early cranial morphogenesis of representative reptilian
species through comparative analysis of gene expression
patterns and found unique expression patterns of Runx2
and Msx2 in turtle embryos that are not observed in
crocodile and snake embryos.
Runx2 is widely known as a transcription factor that
plays a fundamental role in osteoblast differentiation in
vertebrate embryos (Ducy et al., 1997, Komori et al.,
1997, Mundlos et al., 1997, Nakashima et al., 2002, Ishii
et al., 2003, Dobreva et al., 2006, Kerney et al., 2010) and
its transcript has been used as a molecular marker for
preosteoblasts or osteoblast progenitors (Ducy et al.,
1997, Karsenty, 2001, Bobola et al., 2003, Ishii et al.,
2003, Abzhanov et al., 2007, Han et al., 2007, Tokita &
Schneider 2009). In turtle embryos where mineralization
of dermal bones of the skull has not yet occurred, mesenchymal cells that express Runx2 were broadly

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Table 1 Expression domains of the genes in the head of crocodile, turtle, and snake embryos
Genes Crocodile
Runx2

Turtle

Snake

St.14/15: The domain dorsal to the oral

cavity where the ventral part of the braincase
and future palatine and pterygoid bones
develop; a domain dorsolateral to the orbit
where the future dorsal projection of the
postorbital bone forms; the domain
ventrolateral to the orbit where future jugal
and postorbital bones form; the mesenchyme
that later differentiates into the main body of
the postorbital bone.

St.14/15: A population of cells medial to the

precursor of the jaw adductor muscles; the
mesenchyme localized at the domain
dorsolateral and ventrolateral to the orbit; a
thick layer of the mesenchymal cells that
completely covers the brain and the
precursor of jaw adductor muscle laterally.

St.26: The mesenchyme occupying the space

medial to the quadrate cartilage precursor;
the mesenchyme ventral to the orbit; a layer
of mesenchymal cells surrounding the brain
laterally.

St.17: The cell populations localized to the

area where the future dermatocranium
differentiates (palatine, parietal, postorbital,
pterygoid, quadratojugal bones).

St.17: A thick layer of mesenchymal cells

surrounding the braincase and jaw adductor
muscle laterally; the mesenchyme associated
with the quadrate cartilage and the ventral
part of the braincase.

St.29: The precursors of palatine and

pterygoid bones; the mesenchyme
accompanying jaw cartilages; a layer of loose
mesenchyme that later forms a precursor of
parietal bones; the precursors of the maxilla
bones.
St.31: The precursors of the dermatocranial
elements, including the parietal.

MyoD

Sox9

Scx

Six2

St.14/15: Precursor cells of each jaw muscle

in the first pharyngeal arch; eye muscle
precursors.

St.14/15: The primordia of jaw and eye

muscles.

St.26: The primordia of jaw and eye muscles.

St.17: Differentiated jaw and eye muscles.

St.17: Differentiated jaw and eye muscles.

St.29-31: Differentiated jaw and eye muscles.

St.14/15: Cartilage precursors that later

differentiate into the quadrate, Meckel's
cartilage, and the braincase.

St.14/15: Precursor cells of the braincase,

quadrate, and Meckel's cartilages.

St.26: The precursors of quadrate and

Meckel's cartilages and the braincase; a layer
of mesenchyme surrounding the brain
laterally.

St.17: Differentiated chondrocranium and

splanchnocranium components (the
braincase, quadrate, and Meckel's).

St.17: Differentiated chondrocranium and

splanchnocranium components (the
braincase, quadrate, and Meckel's).

St.29-31: Differentiated chondrocranium and

splanchnocranium components (the
braincase, quadrate, and Meckel's).

St.14/15: Tendon precursor cells within jaw

muscle primordia; connective tissue within
eye muscles.

St.14/15: A layer of mesenchymal cells

located at the periphery of the jaw adductor
and eye muscle precursors.

St.17: Tendinous tissues accompanying jaw

muscles; connective tissue associated with
eye muscles.

St.17: Tendinous tissues at the periphery of

jaw adductor muscles; the precursor of the
bodenaponeurosis (central tendon of external
adductor) within the jaw adductor muscular
tissue.

St.14/15: The mesenchyme surrounding the

eyes and cartilaginous precursors of the
braincase, quadrate, and Meckel's; the
mesenchyme between jaw muscle precursors
and the skeletal tissues to which the muscles
attach; the mesenchyme that dorsally
surrounds the brain.

St.14/15: The mesenchyme surrounding the

eye; the mesenchyme associated with the
braincase and jaw cartilages; the
mesenchyme within the jaw muscle
precursors.

St.17: The mesenchyme localized around the St.17: The mesenchyme surrounding jaw
jaw articulation between quadrate and
adductor muscles, braincase, and jaw
Meckel's; the mesenchyme associated with
cartilages.
the braincase, postorbital bone, and jaw
muscles.
Bmp4

St.14-17: The epithelium of cochlearcanal;

the mesenchyme surrounding the eye; the
mesenchyme distributed in the medial part
of jaw primordia; the precursors of the
palatine bones; a population of mesenchymal
cells covering the brain dorsally.

St.14-17: The epithelium if cochlear canal;

The mesenchyme dorsolateral and
ventrolateral to the eye; a limited population
of the mesenchyme in close proximity of the
jaw articulation.

Msx1

St.14-17: The epithelium of the cochlear

canal; the mesenchyme adjacent to the jaw
articulation; the mesenchyme lateral to the
quadrate and Meckel's cartilages; a thin layer
of mesenchymal cells covering the brain
dorsally.

St.14-17: The epithelium of the cochlear

canal; the mesenchyme adjacent to the jaw
articulation; the mesenchyme lateral to
quadrate and Meckel's cartilages; the
mesenchyme that populates the domain
dorsal to the eye.

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Table 1 Expression domains of the genes in the head of crocodile, turtle, and snake embryos (Continued)
Msx2

St.14/15: A thin layer of mesenchymal cells

surrounding the dorsal aspect of the brain;
the mesenchyme located dorsal to the eye;
the mesenchyme occupying the domain
between the ventrolateral part of quadrate
cartilage and surface epidermis.

St.14-17: Mesenchymal cells populating

lateral aspect of the head (lateral to external
adductor muscle).

St.17: A population of mesenchymal cells in

close proximity of postorbital and
quadratojugal bone precursors; a thin layer of
the mesenchyme surrounding the brain
dorsally where future parietal bones form.

St.26: The mesenchyme medial to the

quadrate precursor; a mesenchymal layer
surrounding the brain dorsally.

St.29: The precursors of palatine and

pterygoid bones; the mesenchyme
accompanying jaw cartilages; a layer of loose
mesenchyme that later forms a precursor of
parietal bones.
St.31: The precursors of the dermatocranial
elements, including the parietal.

distributed in the lateral domain of the head, from the

top of the head to the ventral margin of the jaw (Figure 9). This means that turtle embryos have wider
distribution of the cells that have a potential to differentiate into bones in the temporal region of the
head, compared to other lineages of reptiles. The reduction of the amount of Runx2 mRNA causes developmental defects in calvarial bones, called
cleidocranial dysplasia, in mouse embryos (Lou et al.,
2009). In contrast, early onset of Runx2 expression
that eventually results in an increase of the amount
of its mRNA in the cranial mesenchyme accelerates
the timing of mineralization of cranial dermal bones
in mouse embryos and brings about craniosynostosis
characterized by overgrowth of bones (Maeno et al.,
2011). Similar result has been obtained from experiments using avian embryos (Merrill et al., 2008). We
speculate that heterotopy in the preosteoblast distribution observed in early stage turtle embryos may
lead to the increase of the amount of Runx2 expression that results in the increase of the level of ossification in the temporal region of their skull and this
would be a primary factor to build the anapsid skull
where the temporal region is completely covered
with bone.
In this study, we focused on several candidate molecules that potentially regulate Runx2 expression and examined their expression patterns in reptilian embryos.
Bmp4 is known to be involved in osteogenesis of vertebrates where it regulates expression of other osteogenic
regulatory genes, including Msx1, Msx2, and Runx2
(Marazzi et al., 1997, Kim et al., 1998, Hollnagel et al.,
1999, Tribulo et al., 2003, Zhang et al., 2003, Brugger
et al., 2004). Msx1 is a transcription factor known to
regulate growth and patterning of calvarial bones in
mouse embryos (Satokata & Maas, 1994, Han et al.,
2007, Roybal et al., 2010). Although, as previously
reported in mouse embryos (Rice et al., 2003, Han et al.,
2007), both Bmp4 and Msx1 are expressed in limited

populations of cranial mesenchyme in embryos of crocodiles and turtles, we could not detect any substantial differences in their expression domains between the two
species. On the other hand, we observed spatially different expression patterns of Msx2 in the head of embryos
of all reptilian species we examined. Expression of Msx2
was detected in cranial mesenchyme and dermal bone
precursors as reported in mouse embryos (Jabs et al.,
1993, Ishii et al., 2003, Rice et al., 2003, Han et al., 2007,
Roybal et al., 2010). Furthermore, its expression spatially
overlapped with that of Runx2 in reptilian embryos, as
in mouse embryos (Ishii et al., 2003, Rice et al., 2003,
Han et al., 2007). In turtle embryos, expression domain
of Msx2 in the mesenchyme distributed in the temporal
region of the head was broad in a dorsal-ventral direction, showing similar pattern with Runx2 in the mesenchyme. A mutation in the homeobox of Msx2 gene
causes craniosynostosis in human and mouse (Jabs et al.,
1993, Liu et al., 1999). Similarly, overexpression of Msx2
promotes osteogenesis (Cheng et al., 2003, Ichida et al.,
2004) and causes overgrowth of dermal bones of the
skull by increasing the number of proliferative osteoblasts (Dodig et al., 1999, Liu et al., 1999). In contrast,
loss-of-function of Msx2 results in defects of skull ossification in mammals (Satokata et al., 2000, Wilkie et al.,
2000, Ishii et al., 2003, Antonopoulou et al., 2004, Han
et al., 2007). Furthermore, Msx2 is known to positively
regulate downstream Runx2 expression (Ishii et al.,
2003, Han et al., 2007, Watanabe et al., 2008). Considering the evidence provided by previous studies, regulatory
changes in Msx2 expression in turtle embryos may
influence expression patterns of downstream Runx2,
which regulate osteoblast differentiation. Dorsoventrally
broadened distribution of osteogenic mesenchymal precursor cells in the temporal region of the head owing to
the regulatory alteration of these osteogenic genes may
allow this reptilian lineage to reacquire the anapsid
skull. Although the precise mechanism underlying regulatory change of Msx2 expression in the head of turtle

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Figure 9 Potential developmental basis that generates morphological diversity in the temporal region of the reptilian skull. All extant
reptilian lineages are considered to be derived from ancestor with diapsid skull. In crocodiles that have both upper and lower temporal bars like
the stem Diapsida (e.g., Petrolacosaurus), osteogenic mesenchymal precursor cells which express Runx2 and/or Msx2 are distributed at the domain
where future temporal bars are formed in the head of early stage embryo (top of the middle column). Through ontogeny (black arrow), these
osteoblast precursors may differentiate into the dermatocranial elements including upper and lower temporal bars (bottom of the middle
column). Between these bony bars, both upper and lower temporal fenestrae are clearly recognized. In turtles (left white arrow), distribution of
osteogenic mesenchymal precursor cells is broadened in a dorsal-ventral direction, filling the whole lateral portion of the head of early embryos
(top of the left column). Through ontogeny, these osteoblast precursors may differentiate into the dermatocranial elements roofing the temporal
region of the head (bottom of the left column). In snakes that have modified diapsid skull where temporal bars are absent, osteogenic
mesenchymal precursor cells do not fill lateral domain of the head of embryo. Rather, these cells are mainly distributed in the vicinity of the brain
(top of the right column). Through ontogeny, these osteoblast precursors may differentiate into the dermatocranial elements accompanying the
braincase, without forming bony temporal bars (bottom of the right column). A condensed mesenchymal layer that differentiates into the
braincase in later stages is highlighted by dotted line in the head of embryos.

embryos has not been identified, recent findings that

early stage arrest of Msx2 expression in neural crestderived odontoblasts may account for the absence of
teeth in turtles (Tokita et al., 2012) supports the
hypothesis that this transcription factor may play a
pivotal role in the development of their unique cranial
morphology.

The development of the dermatocranium occurs in

multiple steps (Ishii et al., 2003). The first phase includes
the genesis, migration, and initial specification of osteogenic mesenchymal precursor cells. The second phase
consists of the differentiation of the mesenchyme into
osteoblasts. And the last phase includes deposition of
osteogenic extracellular matrix around the osteoblasts

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and mineralization of the matrix. The dermatocranium

of vertebrates is formed from cranial mesenchyme derived from two distinct embryonic sources: neural crest
and mesoderm (Jiang et al., 2002, Gross & Hanken
2005, Noden & Trainor, 2005). Unfortunately, fate
mapping studies of each dermatocranial element as
performed in avian and mammalian embryos (Le Livre,
1978, Noden, 1978, 1983, Couly et al., 1993, Kntges &
Lumsden, 1996, Jiang et al., 2002) have not been done in
non-avian reptiles. Interestingly, the pattern of migration
and distribution of cranial neural crest cells from which
some cranial dermal bones should form is almost identical
in early stage embryos of crocodiles and turtles (Meier &
Packard, 1984, Hou & Takeuchi, 1994, Kundrt, 2008).
Such data may support that differentiation or maturation
processes of osteogenic mesenchyme are more responsible
for producing diversity of reptilian skull morphology. We
speculate that the developmental program, which determines cranial mesenchymal populations where earlyphase osteogenic transcription factors Msx2 and Runx2
are activated, may be important in the patterning of reptilian skull morphology.
There exists substantial diversity in the skull morphology within turtles and most living turtle species do not
have fully anapsid skulls and instead possess varying degrees of dorsal and/or ventral emargination on their
skull (Jones et al., 2012, Werneburg, 2012a). In the
present study, we could not sample and analyze the embryos of turtle species with fully anapsid skull, such as
marine turtles (Kuratani, 1999, Jones et al., 2012), alligator
snapping turtle (Macrochelys temminckii) (Sheil, 2005),
and big-headed turtle (Platysternon megacephalum),
owing to difficulty in the access to the materials. Instead,
we analyzed the embryos of a soft-shelled turtle species
with highly emarginated skull. In fact, soft-shelled turtles
have only a narrow bar of bone across the temporal region
lateral to the external adductor muscles due to large scale
emargination from the dorsal and ventral margins of the
cheek (Ogushi, 1911, Sheil, 2003). In normal development
of soft-shelled turtles, the postorbital bone does not grow
in a posterior direction significantly, keeping its relatively
small size within the dermatocranium (Sheil, 2003,
Snchez-Villagra et al., 2009). Therefore, the small postorbital bone of soft-shelled turtles does not largely contribute to the formation of a bony roof at the temporal
region of the skull.
It is interesting that we observed dorsoventrally broadened distribution of the mesenchymal cells that express
Runx2 at the temporal region of the embryos of a softshelled turtle species with highly emarginated skull. Dermal
bone development occurs through a multi-step molecular
pathway regulated by different transcription factors (Zhang,
2010). As an initial step, Runx2 is required for the differentiation of mesenchymal cells into preosteoblasts. In

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subsequent stage where these preosteoblasts differentiate

into mature osteoblasts, Osx, a downstream gene of Runx2,
is necessary (Nakashima et al., 2002, Nishio et al., 2006).
Furthermore, in the later stages where the osteoblasts produce osteogenic extracellular matrix and the mineralization
of these extracellular matrix is occurred, many additional
molecules such as bone sialoprotein, osteopontin, and
osteocalcin are involved (Zhang, 2010). We speculate that
in soft-shelled turtles only a limited population of cells
within Runx2-positive preosteoblasts distributed in the
temporal region of the head is allowed to differentiate into
mature osteoblasts and eventually osteocytes through
regulation of expression of down stream genes (e.g. Osx),
to form a pair of relatively small postorbital bones.
Although the regulatory mechanism of Osx expression in
osteogenic mesenchyme is not fully understood, both
Runx2-dependent and -independent pathways have been
suggested (Lee et al., 2003, Celil & Campbell, 2005,
Maehata et al., 2006, Xing et al., 2007, Zhang, 2010). Histological analysis reveals that late stage soft-shelled turtle embryos have a layer of (non-muscular) fibrous connective
tissue lateral to the external adductor muscles (Additional
file 3). Judging from its position, the connective tissue layer
appears to be derived from Runx2-positive preosteoblasts
and have a potential to ossify themselves as other connective tissues represented by tendons and ligaments (Okawa
et al., 1998; Tokita et al., 2007). Interestingly, similar type
of connective tissue layer is absent in the temporal region
of crocodile and snake embryos (Additional file 3). Those
histological observations support the above hypothesis that
later processes of cranial osteogenesis may largely contribute to the construction of the main body of each
dermatocranial element from the osteogenic mesenchymal
progenitor pool.
The dorsoventrally broadened distribution of preosteoblasts observed in turtle embryos might be a developmental synapomorphy re-acquired by the common ancestor of turtles. In the course of chelonian evolution, each
chelonian lineage may develop the temporal dermal bones
(e.g. postorbital, parietal, jugal) with various sizes and
shapes, through regulatory changes of the osteogenic down
stream molecules. Future studies should investigate expression pattern of Runx2 and Msx2 in the head of embryos of
turtle species with fully anapsid skull, as well as expression
pattern of downstream genes that regulate differentiation of
mature osteoblasts and osteocytes in turtle embryos, to verify a correlation between the gene expression pattern and
their skull morphology.
Heterotopy in distribution of osteogenic mesenchymal
precursor cells and diversification of reptilian skull
morphology

The frame-like skulls possessed by diapsid reptiles

evolved in response to functional forces (Rieppel, 1993a,

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Moazen et al., 2009, Herrel et al., 2007, Curtis et al.,

2011) and several studies have suggested heterochrony
as a driving force for producing this morphological diversity (Rieppel, 1993a, Whiteside, 1986, Irish, 1989). The
ancestral lineage of diapsid reptiles possessed upper and
lower temporal bars that encircle temporal fenestrae
(Mller, 2003, Moazen et al., 2009). The lower temporal
bar that encloses lower temporal fenestra ventrally was
probably lost once in the common ancestor of
lepidosaurs and archosaurs, possibly as the outcome of
paedomorphosis: incomplete ossification of a quadratomaxillary ligament between jugal and quadratojugal
bones (Rieppel, 1993a, Mller, 2003). If this is true, the
lower temporal bar that possibly results from
peramorphosis (hypermorphosis): complete ossification
of a quadrato-maxillary ligament was independently reacquired in the lineages of tuatara and crocodiles, as
well as in several extinct reptilian lineages (Rieppel,
1993a, Mller, 2003). Furthermore, disappearance of
upper temporal bar, which is regarded as an extreme
condition of reduction of the dermatocranium in reptiles, may have independently evolved in the skull of
geckos (Gekkonidae), miniaturized fossorial lizards (e.g.,
Typhlosaurus, Dibamus), amphisbaenian, and snakes, as
the outcome of paedomorphosis represented by the retardation of ossification (Rieppel, 1993a, Irish, 1989,
Cundall & Irish, 2008). In the present study, we revealed
a possible correlation between distribution pattern of
Runx2 and/or Msx2-expressing osteogenic mesenchymal precursor cells and the skull morphology of each
reptilian lineage (Figure 9). In early stage crocodile embryos, we observed focal distribution of osteogenic mesenchyme around the domain where future temporal
bars are formed. In early stage snake embryos, osteogenic mesenchymal cells were primarily found adjacent
to the primordium of the braincase and the spatial pattern presaged the absence of bony temporal bars in the
temporal region of adult animal.

Conclusions
Regulatory modifications of Runx2 and Msx2 expression in osteogenic mesenchymal precursor cells are
likely involved in generating morphological diversity
in the temporal region of the reptilian skull, including
secondary closure of the temporal fenestrae in turtles.
Our findings demonstrate that not only heterochrony
in ossification of the dermatocranium that has been
traditionally regarded as the major factor producing
diversity of reptilian cranial morphology but also
heterotopy in distribution of the osteogenic precursor
cells may play a fundamental role in this process and
it should be further investigated in future studies of
reptilian cranial development and evolution.

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Materials and methods

Sample collection and staging of embryos

Fertilized eggs of Chinese soft-shelled turtle, Pelodiscus

sinensis, were purchased commercially from a local breeder
in Japan. Fertilized eggs of Siamese crocodile, Crocodylus
siamensis, were provided by a local breeder in Thailand.
Fertilized eggs of corn snake, Pantherophis guttatus, were
obtained by the first author after mating several pairs of the
reproductively mature adults in the laboratory. Staging of P.
sinensis embryos was performed after Tokita and Kuratani
(2001). Because there is no embryonic staging system for C.
siamensis at present, we used the system for Alligator
mississippiensis embryos (Ferguson, 1985) where each stage
was determined based on external morphology of the
embryos, for staging of this species. Staging of P. guttatus
embryos was performed on the basis of staging table of
Thamnophis sirtalis (Zehr, 1962). Interspecific comparisons
of gene expression pattern were performed in the embryos
that are comparable to each other in terms of overall external morphology. Because snake embryos are limbless,
we mainly employed external features of the head of
the embryos as primary criteria for determining the
stages for comparison. All animal experiments were
approved by the University of Tsukuba Committee for
Animal Care (No.10-034).
Molecular cloning

Total RNA was extracted from embryos using ISOGEN

reagent (NIPPON GENE CO., LTD).
RTPCR was performed to amplify fragments of P.
sinensis Runx2, Six2 and C. siamensis Bmp4, Msx2, MyoD,
Runx2, Scleraxis (Scx), Six2, Sox9 and P. guttatus Msx2,
Runx2, Sox9 messenger RNA. Primer sequences used for
isolation of the fragments of these genes are available upon
request. Because Bmp4, Msx1, Msx2, MyoD, Scx, Sox9 of
Pelodiscus and MyoD of Pantherophis were already sequenced and sequence data were deposited in the database
by other researchers, we isolated the orthologous fragments by RTPCR with primers constructed by referring
to the reported sequence data. The fragments were isolated
using the pGEM T-easy vector systems (Promega) or
TOPO TA cloning kit (Invitrogen) and sequenced using
an ABI 3130 sequencer (Applied Biosystems). To identify
the orthologous genes of the isolated fragments, comparable sequence data were surveyed using a BLAST search,
and phylogenetic trees with neighbor joining method
were constructed after sequence alignment using the
CLUSTALX software. All new DNA sequence data were
deposited in the DDBJ database (AB811933-AB811944).
Gene expression analysis

Embryos were fixed in 4% PFA, dehydrated using an

methanol series, placed in xylene, embedded in paraffin,

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and sliced with a microtome. Serial sections were hybridized with digoxigenin-labeled RNA riboprobes as
described in Neubser et al. (1995) with slight modifications. To identify the expression domain of Msx1 in crocodile tissues, chicken Msx1antisense riboprobe was
hybridized. Generally, hetero-specific RNA probes easily
hybridize among reptilian lineages (Harris et al., 2006,
Tokita et al., 2012). In this study, we only analyzed reptilian
embryos at the ontogenetic stages where early cranial
osteogenesis occurs. To confirm the expression pattern of
each gene in the cranial tissues, two to five individuals
representing each embryonic stage were sampled for analysis. The consistency of the gene expression patterns
among all individual embryos at the same stage was confirmed. Multiple sections representing several longitudinal
(anterior-posterior) planes prepared from the same individual were hybridized with the probes and the sections prepared at corresponding longitudinal planes were compared
between different individuals. Corresponding longitudinal
planes between different reptilian species were determined
based on overall histological configuration of the head of
the embryos. For visualization of each cranial tissue and
interspecific comparison of general histology of the head,
Miligan's Trichrome staining was performed following
standard protocols. To identify each anatomical structure in
cranial musculoskeletal tissues of the embryos, we took the
results of other's researches into account: (Schumacher,
1973, Rieppel, 1993b, Vickaryous & Hall, 2008, Bona &
Desojo, 2011) for crocodile, (Schumacher, 1973, Rieppel,
1990, Rieppel, 1993c, Snchez-Villagra et al., 2009,
Werneburg, 2012a, 2012b) for turtle, and (Kamal et al., 1970,
Haas, 1973, Zaher, 1994, Buchtov et al., 2007) for snake.

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including the braincase and quadrate, are labeled by Sox9 probe. (E) Six2
is expressed mainly in the mesenchyme around Sox9-positive cartilage
precursors and MyoD-positive cranial muscle anlagen. Scale bar in (A) is 1 mm.
Scale bars in (B-E) are 0.5 mm.
Additional file 3: A layer of fibrous connective tissue lateral to the
external adductor muscle is found in late stage soft-shelled turtle
embryo. (A) Lateral view of the head of a turtle embryo at stage 22.
(B, C) Frontal sections of the head prepared in the planes indicated in
(A). Note a clear layer of fibrous connective tissue lateral to the external
adductor muscle (red arrowheads). (D) Lateral view of the head of a
crocodile embryo at stage 20. (E, F) Frontal sections of the head
prepared in the planes indicated in (D). (G) Lateral view of the head of a
snake embryo at stage 31. (H, I) Frontal sections of the head prepared in
the planes indicated in (G). A layer of fibrous connective tissue is not
seen in the domain lateral to the external adductor muscle in crocodile
and snake embryos. Rather, in crocodile and snake embryos, the domain
is occupied by mesenchymal cells in low density or by acellular cavities.
Scale bars are 1 mm.

Abbreviations
Am: Jaw adductor muscle or the anlage of jaw adductor muscle;
Ame: External adductor muscle; Bc: Braincase; Bp: Basicranial plate cartilage;
Br: Brain; C: precursor of jaw cartilages (quadrate and Meckel's); Co: Cochlear
canal; E: Eye; Em: Eye muscles or the anlagen of eye muscles;
Itm: Intramandibular muscle; J: Jugal bone; Jm: Anlage of jaw muscle
complex; Mc: Meckel's cartilage; Mx: Primordium of maxilla bone; Oc: Oral
cavity; P: Parietal bone or precursor of parietal bone; Pc: Parachordal
cartilage; Pg: Primordium of pterygoid bone; Pgm: Pterygoid muscle;
Pl: Primordium of palatine bone; Po: Postorbital bone or primordium of
postorbital bone; Ptm: Pseudotemporal muscle; Q: Quadrate cartilage or
precursor of quadrate cartilage; Qj: Quadratojugal bone or primordium of
quadratojugal bone; T: Trabecular cartilage; Tr: Ganglion of trigeminal nerve;
1a: First pharyngeal arch; 2a: Second pharyngeal arch.
Competing interests
The authors declare that they have no competing interests.
Authors contributions
MT conceived and designed the study. MT, WC, and JS contributed to
collection and preparation of biological materials. MT performed
experiments and analyses and wrote the paper. All authors read, discussed
and approved the final manuscript.

Additional files
Additional file 1: Expression of musculoskeletal tissue marker
genes in the head of crocodile embryos at stage 14. (A) Lateral view
of the embryo. (B-D, and F) Frontal sections prepared around the plane
indicated by the blue line in (A). (E) Frontal section prepared around the
plane indicated by the red line in (A). (B) Expression of Runx2 is faintly
detected at the mesenchymal cells above and below the eye, as well as in
the mesenchyme distributed medial to the precursor of quadrate cartilage
and in the mesenchyme surrounding the braincase (arrows). (C) Cranial
muscular tissues are clearly labeled by MyoD probe. (D) Cartilaginous tissues,
including the braincase and the quadrate (q), are labeled by Sox9 probe. (E)
Scx is expressed in tendon precursor cells in close proximity of MyoDpositive jaw and eye muscle anlagen (arrowheads). (F) Six2 is expressed
mainly in the mesenchyme around Sox9-positive cartilage precursors,
including the quadrate and the braincase, as well as in the mesenchyme
around MyoD-positive cranial muscle anlagen. The red outlined domains
indicate the location of the anlagen of the jaw muscle complex. Scale bar in
(A) is 1 mm. Scale bars in (B-F) are 0.5 mm.
Additional file 2: Expression of musculoskeletal tissue marker
genes in the head of turtle embryos at stage 14. (A) Lateral view of
the embryo. (B-E) Frontal sections prepared around the plane indicated
by the red line in (A). (B) Runx2-positive mesenchymal cells are
distributed above and below the eye, as well as in the domain medial to
the anlagen of jaw adductor muscle (black arrows). (C) Cranial muscular
tissues are clearly labeled by MyoD probe. (D) Cartilaginous tissues,

Acknowledgments
We appreciate Sriracha Crocodile Farm & Product Co., LTD. and the company
staffs, especially Nussara Thongprasert in collection of fertilized eggs of
Crocodiles. We also thank Manasaree Klomtun, Punnapa Pinweha, and Ekawit
Threenet for their kind help in collection of Crocodile eggs. MT thanks
Hiroshi Wada who allowed the use of facilities for experiments and analyses,
Hiroki Ono who kindly gifted chick Msx1 probe, Matthew Brandley and
Richard Schneider for critical reading and editing of a draft of the
manuscript, and Johannes Mller for helpful comments to early version of
the manuscript. This study was partially supported by Grants-in-Aid from the
Ministry of Education, Culture, Sports, Science and Technology of Japan to
M.T. (22770077).
Author details
Graduate School of Life and Environmental Sciences, University of Tsukuba,
Tenno-dai 1-1-1, Tsukuba, Ibaraki 305-8572, Japan. 2Department of Zoology,
Kasetsart University, 50 Ngam Wong Wan Road, Chatuchak, Bangkok 10900,
Thailand. 3Department of Organismic and Evolutionary Biology, Harvard
University, 16 Divinity Avenue, Cambridge, MA 02138, USA.
1

Received: 25 June 2013 Accepted: 8 July 2013

Published: 23 July 2013
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Cite this article as: Tokita et al.: Skeletal gene expression in the temporal
region of the reptilian embryos: implications for the evolution of reptilian
skull morphology. SpringerPlus 2013 2:336.

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