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

Nuchal Crest Temporal fossa

Cerebellar prominence

Postorbital Process

Foremen for the

external occipital vein

Squamosal

Posterior portion of the

jugal bar

Exoccipital
Parotic process Hypoglossal foramen (CN XII)
Foramen magnum
Occipital Condyle Palatine
Vagus foramen (CNX) Basilar tubercle


 9

 

Dorsal
View




 


Parotic process Postorbital process

Supraorbital fossa
Parietal Jugal
Nuchal crest
Palatines Nasals
Premaxilla
Nares

Supraoccipital

Saggital crest

Maxilla
Frontal

Quadratojugal


 10

 Ventral
View








 


Pterygoid
Quadratojugal

Tympanic Cavity Maxilla Premaxilla

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

Angular Prearticular Rostral Mandibular Splenial

Fenestra


 12

 

References



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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
quadrate,
two
important
bones
in
the
avian
skull
that
are
not
preserved

in
the
specimen
described
in
this
document.






 15