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A
Brief
Description
of
the
Skull
of
Spheniscus
megellanicus,
the
Megellanic
penguin
By
Matthew
Borths
Department
of
Anatomical
Sciences
for
HBA
550:
Vertebrate
Evolution
Introduction
The
Megellanic
penguin
(Spheniscus
megellanicus)
is
found
on
the
South
Atlantic
coast
of
South
America
near
Tierra
del
Fuego
and
From
theanimalfiles.com
on
the
Pacific
coast
in
the
vicinity
of
the
Falkland
Islands.
In
the
winter
season
they
will
migrate
north
to
the
coasts
of
Brazil
or
Peru,
following
the
temperate
weather
they
prefer.
Megellanic
panguins
are
medium‐sized
for
penguins,
averaging
70
cm
in
length
and
weighing
between
3.8
kg
and
6.5
kg.
Like
all
penguins,
they
have
relatively
short
necks,
short
wedged
tails
and
rigid
wings
supported
by
fused
carpels
that
act
as
hydrofoils
rather
than
airfoils.
The
fused
wrist
prevents
penguins
from
folding
their
forelimbs
as
most
other
birds
do,
but
the
additional
rigidity
makes
the
wing
a
more
effective
paddle
for
moving
through
the
water.
They
are
countershaded
with
white
stomachs
and
black
backs,
likely
serving
the
dual
purpose
of
camouflaging
the
penguin
from
its
predators
and
prey
as
it
dives
through
the
water
with
other
group
members
in
pursuit
of
squid,
small
fish,
and
crustaceans.
The
Megellanic
penguin,
like
all
species
in
the
genus
Spheniscus,
has
a
distinctive
white
band
running
from
behind
the
eye,
around
the
mandible,
and
across
the
neck.
A
second
white
band
loops
around
the
stomach
and
thorax
(Wong
2001).
The
feathers
that
create
this
distinctive
pattern
are
water‐proof
and
insulate
the
animal
in
the
cool
depths
of
the
ocean
(Triche
2005).
Other
extant
species
in
the
genus
Spheniscus
(meaning
“wedge‐shaped)
include
other
“banded
penguins”
such
as
S.
humboldti
(The
Humboldt
Penguin,
native
to
Peru
and
Chile),
S.
mendiculus
(The
Galapagos
Penguin),
and
S.
demersus
(The
African,
or
Jackass
Penguin).
The
first
species
of
Spheniscus,
S.
muizoni,
occurs
in
the
latest
middle/earliest
late
Miocene
(11‐13
Ma)
of
Peru
(Göhlich,
2007),
suggesting
a
South
American
origin
for
the
genus.
S.
muizoni
is
also
the
oldest
occurrence
of
crown
penguins
(Ksepka
and
Clarke
2010).
http://www.theanimalfiles.com/birds/penguins/magel
lanic_penguin.html
2
The
monophyly
of
Sphenisciformes
(penguins)
is
well‐supported
by
molecular
and
morphological
evidence,
but
the
sister
group
to
Sphenisciformes
remains
controversial
with
a
recent
analysis
(Livezey
&
Zusi,
2007)
recovering
Gaviiformes
(loons)
and
Podicipediformes
(grebes)
as
sister
groups
to
Sphenisciformes,
though
the
authors
note
there
may
be
convergence
driving
this
placement
as
all
three
groups
are
characterized
by
aquatic
adaptations.
Molecular
and
fossil
evidence
suggests
penguins
diverged
from
their
last
common
ancestor
with
loons
or
grebes
in
the
earliest
Paleocene
or
the
Late
Cretaceous,
making
a
satisfactory
sister
relationship
difficult
to
recover.
For
further
detail
on
the
phylogeny
of
Sphenisciformes,
refer
to
Appendix
1.
General
Description
of
the
Avian
Skull
Note:
For
greater
detail
on
the
diversity
and
osteological
relationships
between
bones
of
the
avian
cranium,
refer
to
Zusi,
1993.
Anatomical
nomenclature
in
this
discussion
and
in
the
figures
comes
from
Zusi,
1993,
and
Bertelli
et
al.,
2010.
The
Avian
skull
is
a
highly
derived
structure
consisting
of
a
tightly
sutured
neurocranium
surrounding
a
relatively
large
brain
and
large
orbits
and
a
mobile
splanchnocranium
forming
the
jaws.
The
neurocranium
is
composed
of
frontal,
parietal,
occipital,
squamosal,
lacrimal,
pleurosphenoid,
parasphenoid,
ootic,
and
ethmoid
ossification
regions.
These
separate
portions
of
the
skull
are
usually
only
visible
in
juveniles.
The
coelurosaur
ancestors
of
birds
had
less
firmly
sutured
neurocrania
and
possessed
several
cranial
bones
that
have
been
lost
in
the
course
of
avian
evolution
which
are
highlighted
in
the
adapted
figure
below
from
Göhlich
&
Chiappe
2006.
These
include
the
postorbital
(po,
orange),
the
postorbital
process
of
the
jugal
(red),
the
squamosal
process
of
the
quadratojugal
(qj,
yellow),
the
ectopterygoid
(not
visible),
and
coronoid
of
the
mandible
(not
shown).
3
The
splanchnocranium,
including
the
premaxilla,
maxilla,
nasals,
jugal,
quadratojugal,
palatines,
pterygoids,
quatrate,
articular,
splenial,
supraangular,
angular,
articular
and
prearticular,
is
more
mobile
than
the
neurocranium.
The
splanchnocranium
supports
the
keratinized
skin
that
forms
the
beak
.
The
whole
upper
jaw
can
be
protracted
and
retracted
independently
of
the
neurocranium.
The
degree
of
cranial
kinesis
at
the
frontonasal
margin
is
variable,
but
all
birds
posses
some
degree
of
mobility
at
the
quadrate.
Neurocranium
The
anterior
portion
of
the
neurocranium
is
formed
by
the
roofing
frontals
that
arch
over
the
orbits.
Anteriorly,
the
frontals
meet
the
mesethmoid,
a
bone
that
contributes
to
the
anterior
portion
of
the
orbit.
Posterior
to
the
mesethmoid
is
the
interorbital
septum,
a
common
feature
in
birds
that
varies
in
extent
and
density.
The
ectethmoids
laterally
fuse
to
the
mesethmoid
and
form
a
portion
of
the
lacrimal
canal.
Lateral
to
the
ectethmoid
sits
the
lacrimal,
a
bone
that
rims
the
lacrimal
duct
and
contacts
the
jugal
bar.
The
lacrimal
has
also
been
called
the
preorbital
and
it
forms
the
posterior
border
of
the
antorbital
fenestra.
The
posterior
portion
of
the
frontal
contacts
the
parietals
and
squamosals.
Near
this
junction,
a
lateral
process
descends
behind
the
orbit
forming
the
postorbital
process
which
can
be
formed
from
the
pleurosphenoid,
squamosal,
or
frontal.
The
postorbital
ligament
continues
from
the
tip
of
the
process
and
inserts
on
the
mandible,
anterior
to
the
quadrate/articular
articulation.
The
lateral
portion
of
the
parietal
and
squamosal
can
have
a
shallow
fossa
and
rugosity
associated
with
the
origin
of
the
temporalis
muscle.
The
squamosal
also
has
a
condyle
that
articulates
with
the
quadrate
at
its
ventral‐most
extent.
The
ventral
portion
of
the
neurocranium
is
formed
by
the
basioccipital
at
the
posterior
margin
of
the
skull.
The
basiocciptial
may
form
a
portion
of
the
occipital
condyle
and
in
some
birds
such
as
the
rhea
it
forms
the
entire
occipital
condyle.
The
anterior
margin
of
the
basiocciptial
abuts
the
basitemporal
plate,
a
structure
derived
from
the
parasphenoid.
The
plate
supports
the
Eustachian
tube
which
opens
at
the
anterior
edge
of
the
plate
near
the
rostral
portion
of
the
parasphenoid,
one
of
the
posterior
supports
for
the
interorbital
septum.
4
The
Eustachian
tube
itself
is
closely
associated
with
the
tightly
fused
prootic,
opisthotic,
and
epiotic
bones
that
form
the
membranous
labyrinth.
The
prootic
has
a
beveled
articular
facet
where
the
quadrate
articulates
near
the
tympanic
membrane.
On
the
posterior
portion
of
the
skull,
four
occipital
bones
fuse
around
the
foramen
magnum:
the
supraocciptial
which
forms
the
superior
margin
of
the
occiput
and
contacts
the
parietals,
the
basioccipital
which
then
forms
the
ventral
portion
of
the
skull,
and
the
exoccipitals
which
usually
house
the
foramina
associated
with
many
of
the
cranial
nerves
including
the
hypoglossal
(XII),
the
accessory
(XI),
the
vagus
(X),
and
the
glossopharyngeal
(IX).
Splanchnocranium
Modern
birds
are
edentulous,
relying
on
specialized
keratinized
skin
to
cut,
crack,
and
spear
their
food.
The
beak
is
supported
internally
by
the
bones
of
the
maxilla
and
mandible.
In
ornithological
literature,
the
maxilla
is
the
tightly
sutured
complex
of
bones
associated
with
the
upper
jaw.
It
is
composed
of
the
elongate
premaxilla
which
forms
most
of
the
dorsal
and
lateral
portions
of
the
bill,
the
nasal
which
begin
posterior
to
the
external
nares
and
forms
the
proximal‐lateral
portion
of
the
bill,
and
the
maxillary
which
limited
to
the
ventral
margin
of
the
maxilla
and
forms
the
distal‐most
portion
of
the
jugal
bar.
The
jugal
bar
is
a
structure
found
in
all
birds.
The
anterior
portion
of
the
bar
is
formed
by
the
maxilla
which
sutures
with
the
jugal.
The
jugal
forms
the
middle
portion
of
the
bar,
then
fuses
with
the
quadratojugal
which
in
turn
contacts
the
mobile
quadrate.
The
articulation
of
the
quadrate
to
the
jugal
bar
allows
mobilization
of
the
upper
jaw
at
the
craniofacial
hinge
which
is
frequently
demarcated
by
a
transverse
line
at
the
juncture
of
the
nasals,
premaxillae
and
the
frontals.
This
is
called
prokinesis.
The
loss
of
the
coelurosaur
cranial
bones
may
be
related
to
an
increased
degree
of
prokinesis
in
the
course
of
avian
evolution,
freeing
the
jugal
bar
to
transmit
force
from
the
quadrate
to
the
maxilla.
Some
birds
posses
a
secondary
hinge
at
the
intersection
of
the
premaxilla
and
the
nasals
(rhynchokinesis)
that
allows
even
greater
independent
movement
of
the
upper
jaw.
5
Along
with
a
jugal
process,
the
maxillaries
have
a
palatine
process
that
may
fuse
to
create
the
bony
desmognathous
secondary
palate
of
some
neognathous
birds
such
as
ducks
and
geese.
However,
most
birds
have
a
keratinized
patent
cleft
palate
(schizognathous
palate).
The
palatines
contact
the
maxillopalatine
processes
anteriorly
and
proceed
posteriorly.
At
the
posterior
edge
of
their
contact
with
the
superior
vomers,
ventral
processes
protrude
from
the
elongate
palatines.
At
the
posterior
margin
of
the
palatines
the
flared
bones
articulate
with
the
pterygoids
which
vary
in
form
from
thin,
strut‐like
structures
(i.e.
Corvus
corax)
to
wide,
flaring
bones
that
earn
their
name
(i.e.
Spheniscus
megellanicus).
The
posterior
portion
of
the
pterygoid
forms
a
mobile
articulation
with
the
quadrate.
The
quadrate
then
has
an
articulation
with
the
palate,
the
jugal
bar,
the
prootic,
and
the
articular
bone.
This
final
articulation
leads
to
the
mandible,
a
rigid
structure
composed
of
six
separate
bones.
Posterior
to
the
mandibular
joint
is
the
retroarticular
process
and
medial
to
the
joint
is
often
a
medial
process.
Anterior
and
inferior
to
the
articular
is
the
angular
bone
which
contacts
the
superior
prearticular.
Superior
and
anterior
to
the
prearticular
is
the
supraangular.
The
supraangular
is
perforated
laterally
by
the
caudal
mandibular
fenestra.
Anteriorly
the
supraangular
contacts
the
dentary
and
forms
the
superior
border
of
the
rostral
mandibular
fenestra.
The
medial,
rostral
portion
of
the
mandible
is
supported
by
the
splenial,
a
bone
not
visible
in
lateral
view.
The
splenial
is
tightly
fused
to
the
dentary
which
forms
the
rest
of
the
distal
mandible.
At
its
tip,
the
dentary
makes
a
rigid
symphysis
with
the
opposing
dentary.
Some
birds
posses
streptognathic
mandibles
that
can
flex
laterally
at
the
intersection
of
the
anterior
splenial
and
dentary
with
the
posterior
supraangular,
prearticular,
and
angular
near
the
rostral
mandibular
fenestra.
Streptognathism
is
particularly
well
developed
in
birds
that
feed
their
chicks
by
letting
the
juveniles
stick
their
heads
into
the
adult’s
mouth
(i.e.
Larus,
the
seagull),
and
in
insect‐eating
birds
that
capture
insects
by
using
their
mouths
as
a
kind
of
net
(i.e.
Nyctibius).
This
kind
of
lateral
flexibility
requires
a
less‐rigid
mandibular
symphysis
at
the
dentary.
6
The
Penguin
Skull
Note:
The
specimen
described
and
illustrated
here
is
missing
both
the
lacrimal
bones
which
sit
between
the
frontal
and
ectethmoids
near
the
junction
of
the
beak
and
neurocranium.
Anterior
to
the
lacrimal
bone
would
be
the
antorbital
fenestra.
This
particular
specimen
is
also
missing
the
quadrate
bone.
This
is
not
surprising
as
the
quadrate
does
not
rigidly
fuse
with
any
portion
of
the
skull.
Refer
to
Appendix
2
for
a
CT‐scanned
Jackass
Penguin
skull
(Sheniscus
demersus)
from
Digimorph
that
preserves
both
these
bones
and
shows
their
articulation
with
the
rest
of
the
penguin’s
cranium.
Also
on
Digimorph
is
a
variety
of
other
penguin
skulls
and
a
wide
diversity
of
other
bird
skulls.
Relative
to
other
birds,
the
penguin’s
skull
is
robust
(Zusi
1993)
with
limited
cranial
kinesis.
One
of
the
most
distinctive
aspects
of
the
penguin’s
skull
is
the
large
supraorbital
fossa.
This
rugose
depression
in
the
frontal
bone
arches
over
the
orbit,
stopping
short
of
the
postorbital
process.
This
fossa
houses
the
supraorbital
gland,
also
called
the
salt‐gland.
This
parasympathetically
controlled
gland
is
a
common
structure
in
marine
birds
that
ingest
salt‐water
when
feeding
on
small
prey
such
as
krill.
The
extreme
salt
accumulation
in
the
blood
would
overwhelm
the
kidneys,
and
the
supraorbital
gland
acts
as
an
accessory
kidney,
filtering
salt
from
the
bird’s
blood
stream.
The
concentrated
salt‐solution
is
then
drained
from
the
gland
through
ducts
that
open
into
the
vestibular
concha
and
finally
empty
from
the
external
nares.
In
terrestrial
birds
this
gland
is
present,
but
marine
species
have
supraorbital
glands
10
to
100
times
larger
than
their
terrestrial
counterparts
(Schmidt‐Nielsen,
K.
1960).
Enlargement
of
the
supraorbital
gland
has
occurred
in
multiple
avian
lineages
that
contain
species
that
exploit
marine
resources
including
Laridae
(seagulls),
Pelecanidae
(pelicans)
and
even
the
toothed
Cretaceous
diving
bird
Hesperornis.
Spheniscus
is
also
notable
for
the
deep
temporal
fossae
that
embay
the
parietal
bones,
raising
a
high
sagittal
crest
that
meets
the
nuchal
crest
at
a
perpendicular
angle.
The
temporal
muscle
that
originates
at
the
tall
sagittal
crest
inserts
on
the
long,
narrow
retroarticular
process
of
the
mandible.
Spheniscus
penguins
can
also
be
readily
recognized
by
their
deep,
triangular
beaks
that
lack
a
ventral
curve
such
as
is
seen
in
Aptenodytes
forsteri
(The
Emperor
Penguin).
7
Lateral
View
Olfactory nerve
CN IV
Supraorbital fossa for salt-gland CN I Mesoethmoid
Parietal
CN II Frontal
Nasal Premaxilla
Lacrimal duct Nares
Postorbital Process
Temporal Fossa
Maxilla
Jugal
Parotic process
Squamosal
Quadratojugal Palatine
Tympanic cavity
Interorbital Septum
8
Posterior
View
Parietal
Supraoccipital
Cerebellar prominence
Postorbital Process
Squamosal
Exoccipital
Parotic process Hypoglossal foramen (CN XII)
Foramen magnum
Occipital Condyle Palatine
Vagus foramen (CNX) Basilar tubercle
9
Dorsal
View
Supraorbital fossa
Parietal Jugal
Nuchal crest
Palatines Nasals
Premaxilla
Nares
Supraoccipital
Saggital crest
Maxilla
Frontal
Quadratojugal
10
Ventral
View
Pterygoid
Quadratojugal
Basilar tubercle
Occipital Condyle
Foramen magnum
Open schizognathus palate
Basioccipital Vomer
Opening of the auditory tube
(Eustachian tube)
Palatine
Parotic process
Rostral portion of the
parasphenoid Postorbital process
11
The
Mandible
Lateral View
Supraangular Dentary
Coronoid Process
Articular Angular
Rostral Mandibular
Caudal Mandibular Fenestra
Retroarticular Process Fenestra
Medial View
Caudal Mandibular
Retroarticular Process Articular Fenestra Supraangular Dentary
Mandibular symphysis
12
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13
Appendix
1.
A
phylogeny
of
Sphenisciformes
calibrated
to
the
stratigraphic
record
from
Ksepka
and
Clarke
2010.
14
Appendix
2.
A
left
lateral
view
of
Spheniscus
demersus
modified
from
http://digimorph.org/specimens/Spheniscus_demersus/.
Note
the
presence
of
the
lacrimal
bone
and
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important
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in
the
avian
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that
are
not
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in
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15