Flightless birds

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NATURAL
HISTORY
American
Museum
oj
Natural
History
Robert
G.
Coelet,
President
Thomas
D.
Nicholson,
Director
Vol.
92,
No.
9.
September
1983
Alan
Ternei.
Editor
Managing
Editor:
Erik
Eckholm
¡Judith
Friedman,
on
k'avfj
Thomas
Page,
Deaigner
Board
of
Editors:
Carol
Brestin,
Sally
Undsay.
Viltorio
Mae.<itro,
Rebecca
U.
Finnell,
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Weinberg
Florence
C
Edelstein,
Copy
Chief
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From
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The
articles
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follow
show
that
ornithology
•
one
of
the
oldest
sciences
and
one
particularly
open
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as
well
as
professionals
•
is
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and
fascinating
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ever.
This
special
issue
was
edited
by
senior
editor
Rebecca
B.
Finnell.
4
Authors
10
At
the
American
Museum
The
AOU's
Fledgling
Years
14
This
View
of
Dfe
Stephen
Jay
Gould
Darwin
at
Sea
22
A
Migratory
Bird's
Baedeker
Kenneth
R
Able
30
Fligiitless
Birds
Helen
F,
James
and
Starrs
L.
Olson
42
Marsh
Wrenditions
Donald
E.
Kroodsma
48
The
Bowerbird's
LalMir
of
Loie
Melinda
Pruett-Jones
and
Stephen
Pruett-Jones
56
Night
Owls
Are
Good
Listeners
Masakazu
Konlshi
60
Remembrance
of
Seeds
Stashed
Stephen
Vander
Wall
and
Russell
P.
Balda
66
Bird
Art
Roger
Tory
Peterson
76
The
Origin
of
a
Species
Peter
R.
Grant
and
Nicola
Grant
82
Gone
with
the
Trees
David
S.
Wllcove
and
Robert
F.
Whitcomb
94
Boolis
in
Review
Michael
Harwood
A
Birder's
Library
9$
Additional
Reading
102
A
Matter
of
Taste
Aajti/Moni/5o/co/ov
About
Eggs
Cover:
Armed
with
asymmetrical
ears,
a
face
that
functions
as
a
sound
collector,
and
a
mental
map,
the
nocturnal
barn
owl
uses
.wund
to
lixate
Its
prey.
Photograph
by
Rick
Mclntyre:
Tom
Stack.
Story
on
page
56.
Suhscriptioft
orders,
change
of
addrcis
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U76Z1

Authors
A
bird
watcher
since
childhood,
Ken-
neth
P.
Abie
was
impressed
early
on
by
the
dramatic
migration
of
birds
in
north-
ern
latitudes
every
spring
and
fall.
As
his
studies
of
birds
became
more
formul,
he
learned
that
just
how
birds
lind
their
way
during
these
migrations
is
far
from
completely
understood.
Since
that
rev-
elation,
his
research
has
centered
on
ori-
entation
and
navigation
mechanisms
in
animals,
especially
those
of
migratory
birds.
Able
is
associate
professor
of
biol-
ogy
at
the
Albany
campus
of
îhe
Slate
University
of
New
York.
Convinced
that
much
remains
to
be
learned
about
bird
song
by
looking
at
species
with
complex
song
systems,
Don-
ald
E.
Kroodsma
examined
different
populations
of
one
of
North
America's
most
voluble
songsters•the
mar.sh
wren.
Associate
professor
of
zoology
at
the
University
of
Massachusetts
in
Am-
herst,
Kroodsma
is
currently
focusing
on
species
of
warblers
that
use
different
songs
in
different
contexts.
He
hopes
to
determine
to
what
extent
the
birds
learn,
rather
than
know
innately,
which
.songs
are
appropriate
for
which
situations.
clearly
demonstrated
thai
bird
fossils,
often
thought
to
be
too
scarce
or
frag-
mentary
to
be
of
much
scientific
use,
can
contribute
significantly
to
our
under-
standing
of
avian
evolution.
In
addition
to
her
work
on
Hawaiian
fossils,
James
plans
to
study
5-mi
I
lion-year-old
bones
of
cormorants
and
parrots
from
South
Africa.
Olson's
projects
include
bird
fos-
sils
from
Bermuda,
the
North
Atlantic,
and
North
America.
Both
Helen
F.
James
and
Storrs
L.
Ol-
son
work
in
the
Division
of
Birds
of
the
Smithsonian
Institution's
National
Mu-
seum
of
Natural
History,
James
as
a
re-
search
assistant
and
Olson
as
curator.
An
interest
in
paleontology
and
the
evo-
lution
of
birds
has
taken
them
several
times
to
the
Hawaiian
Islands,
where
a
number
of
exciting
bird
fossil
finds
have
been
made
in
the
last
decade.
Their
re-
search
in
the
Pacific
and
elsewhere
has
When
Melinda
Pruett-Jones
arrived
in
Papua
New
Guinea
in
1980
with
her
husband
and
coauthor,
Stephen,
she
found
Macgregor's
bowerbirds
in
the
forest
near
the
Wau
Ecology
Institute
where
they
planned
to
work.
"There
was
no
doubt
in
my
mind
what
I
was
going
to
study,"
she
says.
Ever
since
she
was
an
undergraduate
at
the
University
of
Washington,
she
has
been
aware
of
the
unique
behavior
of
bowerbirds,
which
build
and
decorate
terrestrial
display
sites
for
courtship,
Melinda
had
come
to
New
Guinea
to
conduct
research
for
her
M.S.
degree
at
Brigham
Young
Univer-
sity;
Stephen
to
initiate
his
Ph.D.
re-
search
in
association
with
the
University
of
California
at
Berkeley.
His
disserta-
tion
will
focus
on
the
evolution
of
social
behavior
in
Lawes's
six-wired
bird
of
paradise.
The
couple
work
on
indepen-
dent
projects,
but
the
underlying
ques-
tions
in
their
studies
are
the
same
and
they
collaborate
in
much
of
their
work.
Their
scholarly
interests
are
evolution-
ary
biology
and
promiscuous
mating
pat-
terns
in
birds.
When
not
in
the
field,
they
enjoy
sailing,
the
ballet,
good
Italian
res-
taurants,
and
gourmet
ice
cream•none
of
which
can
be
found
in
Wau.
4
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HrsTORV
9/83

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Flightiess
Birds
Recení
fossil
finds
in
Hawaii
suggest
that
in
prehistoric
times,
islands
in
the
Pacific
supported
a
rich
assortment
of
flightless
birds
by
Helen
F.
James
and
Storrs
L.
Olson
Hundreds
of
years
ago,
a
group
of
hard-
pressed,
sun-baked
navigators
sailed
a
double
canoe
through
unfamiliar
waters
for
day
after
weary
day,
until
the
sudden
appearance
of
a
string
of
majestic
moun-
tains,
rising
out
of
the
middle
of
the
Pa-
cific
Ocean,
revived
their
withered
spirits.
With
fearful
anticipation,
they
paddled
ashore
and
became
the
first
human
beings
ever
privileged
to
walk
on
a
Hawaiian
beach.
This
is
a
scene
that
we
have
played
over
and
over
again
in
our
minds
with
un-
suppressed
envy
in
recent
years.
Our
re-
search
on
fossil
bird
bones
of
the
Hawai-
ian
Islands
has
given
us
a
special
insight
into
what
these
Polynesian
discoverers
must
have
found,
and
we
now
know
that
it
was
very
different
from
what
anyone
had
previously
imagined.
We
had
long
known
that
the
islands
were
once
clothed
in
virgin
forests
com-
posed
mostly
of
plants
found
nowhere
else
in
the
world,
but
only
in
the
last
twelve
years
have
we
begun
to
realiz.c
the
extent
to
which
the
forests
were
home
to
a
re-
markable
variety
of
endemic
species
of
birds,
most
of
them
doomed
to
rapid
ex-
tinction
at
the
hands
of
humans.
More
than
half
of
Hawaii's
endemic
species
of
land
birds
would
never
be
seen
by
western-
ers.
Forty-five
or
more
species•including
ibises,
geese,
rails,
owls,
a
hawk,
an
eagle,
ravens,
and
a
plethora
of
songbirds•^van-
ished
in
the
prehistoric
Polynesian
period.
Curiously,
a
large
number
of
these
birds,
no
fewer
than
seventeen,
were
incapable
of
flight.
The
existence
of
these
unusual
birds
was
Hawaii's
best
kept
secret
up
to
the
last
decade.
Partly
because
of
two
unsubstantiated
bi!t
prevalent
bits
of
wisdom-
-that
volca-
nic
islands
are
unlikely
spots
for
fossil
col-
lecting
and
that
fossils
of
birds
are
rare
in
any
case•no
paleontologist
or
biologist
seriously
considered
looking
for
fos.sil
birds
in
the
Hawaiian
Islands
until
1971.
in
that
year,
Joan
Aidem,
an
ardent
natu-
ralist
from
Molokai
Island,
managed
to
ig-
nite
some
interest
with
her
discovery
of
the
complete
skeleton
of
an
extraordinary
flightless
goose
weathering
out
of
a
wind-
blown
sand
dune
on
Molokai.
Since
then,
fossil
sites
have
been
discovered
on
five
of
Hawaii's
eight
main
islands,
in
settings
as
diverse
as
limestone
sinkholes,
a
sea
clifl",
collapsed
lava
tubes
(tunnellike
caves
formed
after
rivers
of
molten
lava
drain
away),
and
a
flooded
cavern
where
skele-
tons
were
found
under
fifteen
feet
of
fresh
water.
Archeological
sites
containing
the
re-
mains
of
prehistoric
meals
have
proved
to
be
another
good
source
of
bones
of
extinct
birds.
These
are
doubly
interesting
be-
cause,
in
combination
with
stratigraphie
and
radiometric
dating
studies,
they
con-
firm
that
many
extinct
species
siill
sur-
vived
when
the
Polynesians
first
arrived,
about
1,500
years
ago.
The
fossils
from
all
these
sources,
now
gathered
at
the
Smith.sonian's
National
Museum
of
Natural
History,
number
in
the
tens
of
thousands.
The
tasks
of
identi-
fying
and
cataloging
the
bones,
isolating
new
species,
writing
scientific
descriptions
of
ihem,
and
tracing
the
evolutionary
rela-
tionships
among
them
arc
complex
and
time
con-suming.
The
work
is
sometimes
tedious,
bul
it
is
amply
rewarded
by
occa-
sional
breakthroughs,
each
augmenting
our
knowledge
of
evolution
on
islands.
The
flightless
species
of
the
Hawaiian
Islands
were
not
representatives
of
some
obscure,
long-extinct
lineage
but
were
de-
scendants
of
such
familiar
kinds
of
birds
as
ducks,
geese,
rails,
and
ibises.
Their
an-
cestors
had
to
have
been
excellent
fliers;
otherwise
they
never
could
have
crossed
the
more
than
two
thousand
miles
of
ocean
between
the
Hawaiian
Islands
and
the
nearest
significant
land
mass.
Once
ar-
rived
in
the
Islands,
many
species
of
birds
responded
to
the
changed
conditions
of
life
by
losing
the
ability
to
fly.
Evolution-
30
NATURAL
HISTORY
9/83

ariiy
speaking,
the
process
may
have
hap-
pened
fairly
rapidly:
the
oldest
of
the
main
islands,
Kauai,
is
a
mere
six
million
years
old,
and
the
youngest,
the
island
of
Ha-
waii,
may
be
less
than
a
million
years
old.
Less
time
than
this
was
available
for
the
evolution
of
flight
lessness
because
the
an-
cestors
of
the
flightless
birds
could
not
col-
onize
a
particular
island
until
after
enough
soil
had
formed
and
plants
arrived
to
cre-
ate
an
environment
suitable
for
terrestrial
birds.
The
genetic
mechanism
for
the
evolu-
tion
of
a
flightlcs,s
bird
from
a
llying
one
is
actually
quite
simple.
All
birds
are
flight-
less
when
they
are
small
chicks,
and
the
young
of
flying
birds
have
the
same
fea-
tures
that
characterize
the
adults
of
flight-
less
birds•proportionately
large
legs,
re-
duced
wings,
and
a
small
or
no
keel
on
the
sternum,
or
breastbone,
(¡n
flying
birds,
the
bony
keel
is
where
ihe
large
flight
muscles
are
attached.)
Merely
by
retain-
ing
the
skeletomuscular
structure
of
in-
fancy
into
adulthood•probably
by
the
alteration
of
a
few
regulatory
genes•al-
most
any
bird
species
could
become
flight-
less.
This
process
of
retention
of
juvenile
characters,
known
as
neotcny,
is
a
fairly
common
evolutionary
phenomenon.
But
why
become
flightless?
To
must
people,
birds
symbolize
flight,
and
most
of
the
evolutionarj-
history
of
the
class
Aves
has
indeed
revolved
around
the
opportuni-
ties
created
by
flight,
on
the
one
hand,
and
the
physical
constraints
of
flight,
on
the
other.
Yet
the
Hawaiian
Islands
are
not
unique;
flightless
birds
occur
on
islands
throughout
the
world's
oceans.
What
great
selective
pressure
spurs
the
develop-
ment
of
flightlessness
on
islands?
Beginning
with
Darwin,
some
scientists
suggested
that
flying
organisms
on
islands
might
be
blown
away
by
winds
and
perish,
thus
leading
to
the
evolution
of
flightless-
ness.
Many
flightless
birds,
however,
are
found
on
New
Zealand
and
other
large
is-
Stand
in
awe
of
pre-Colombian
rui
or
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^^
Flyingbirds.
like
this
modern
goose,
have
well-developed
scapulae
and
coracoids,
which
mee!
at
a
fairly
sharp
angle
and
help
to
brace
the
wing
bones.
They
also
have
a
sizable
keel,
for
the
attachment
of
flight
muscles.
Douglas
Cranner
Flying
Goose
Coracoid
lands,
where
there
is
ample
protection
from
winds.
Moreover,
tiny
islands•Lay-
san
and
Nihoa
in
Hawaii,
for
example•
are
home
lo
small
birds
that
retain
the
ability
to
fly.
The
wind
theory
therefore
is
not
a
satisfactory
one.
The
real
evolutionary
advantage
of
flightlessness
on
islands
may
revolve
around
more
ellicient
use
of
energy.
Mainly
because
of
the
difficulty
terres-
trial
mammais
have
crossing
large
bodies
of
water,
predators•other
than
birds
of
prey•are
relatively
scarce
on
oceanic
is-
lands.
When
neither
escape
from
preda-
tors
nor
seasonal
migration
is
necessary,
individual
birds
with
a
reduced
ability
to
fly
could
have
an
important
advantage.
With
no
large
wings
or
massive
flight
mus-
cles
to
support,
they
would
need
less
food
to
survive
and
reproduce.
Loss
of
the
abil-
ity
to
fly
would
not
be
an
advantage
for
all
types
of
birds,
however.
Species
that
must
fly
lo
obtain
food,
such
as
birds
of
prey
and
small
arboreal
species
that
forage
in
treetops,
do
not
become
flightless,
while
those
that
feed
on
or
near
the
ground
fre-
quently
do.
Among
the
most
con.spicuous
species
that
must
have
greeted
the
first
Hawai-
ians
were
large,
ungainly
gooselike
crea-
tures.
Flightlessness
in
these
birds
had
proceeded
to
the
extreme
stage
in
which
the
sternum
no
longer
bore
any
îrace
of
a
keel.
The
rest
of
the
skeleton
had
taken
on
a
somewhat
comical
aspect,
with
tiny,
dis-
torted,
and
useless
wings
contrasting
with
a
massive
pelvic
girdle
and
large,
stout
leg
bones.
These
were
not
fieet-footed
birds.
The
early
Hawaüans
doubtless
had
little
trouble
catching
them
for
dinner.
The
evolutionary
process
had
so
pro-
foundly
altered
the
skeleton
of
these
Ha-
waiian
geese
that
for
years
we
despaired
of
ever
being
able
to
pinpoint
their
closest
32
NATURAL
HISTORY
9/83

GRAB
YOUR
CAMERA...
TAKE
A
KLR
FAMILY
SAFARI
THIS
CHRISTMAS
December
18-January
1
we
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escorted
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Lynne
Leakey
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•
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^
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For
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visiting
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eminent
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1984.
For
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and
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write
to
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Tours,
Central
Park
West
at
79
St.,
NY
NY
10024,
or
call
{212)
873-1440.
relatives.
Then
in
1982,
an
¡mportanl
cäue
turned
up
in
the
form
of
a
fossil
trachea!
bulla
ttiat
we
received
in
a
collection
of
goose
bones
found
by
naturalist
Michael
Severns
in
a
lava
tube
on
Maui.
The
tra-
chéal
bulla
is
a
hollow,
bony
expansion
of
the
windpipe
thaï
occurs
in
males
of
most
groupsofducksbut
not
in
true
geese
(gen-
era
Anser
and
Brama)
or
swans,
which
can
therefore
be
eliminated
as
possible
an-
cestors
of
the
odd
Hawaiian
birds.
Further
osteological
comparisons
have
narrowed
the
field
to
sheldrakes
and
the
typical
ducks,
such
as
mallards.
Many
different
species
of
these
geese
have
been
found
in
fossil
deposits
on
vari-
ous
Hawaiian
islands.
They
probably
evolved
in
response
to
the
absence
of
na-
tive
herbivorous
mammals
in
the
islands.
Several
species
belong
in
the
peculiar
ge-
nus
Thambetochen,
meaning
"astonishing
goose,"
and
arc
characterized
by
bony,
toothlike
projections
on
the
jaws.
These
birds
may
have
fed
by
browsing
or
grazing
on
coarse
vegetation.
Another
genus
had
a
very
broad,
flat
lower
jaw,
almost
like
that
of
a
turtle,
and
an
extremely
heavy
upper
jaw
higher
than
it
was
long,
giving
it
a
most
unusual
appearance.
This
bill
form
may
also
have
been
an
adaptation
for
browsing
or
grazing.
Intriguing
as
these
goosclikc
birds
are,
they
had
diverged
so
much
from
the
an-
cestral
species
that
they
are
of
little
use
in
our
efforts
to
track
down
the
very
t>egin-
nings
of
flightlessness.
Hawaii
does
boast
one
true
goose,
however,
which
may
prove
helpful
in
this
respect.
The
living
nene,
or
Hawaiian
goose
{Bratita
sandvicensis),
is
the
only
endemic
Hawaiian
goose
to
sur-
vive
into
historical
times,
probably
be-
cause
it
is
perfectly
able
to
fly.
whereas
the
other
species
of
geese
were
not.
Threatened
with
extinction
during
this
century,
the
species
was
rescued
by
a
cap-
tive
breeding
program
and
survives
in
the
upland
regions
of
Hawaii
and
Maui.
Unlike
the
big,
flightless
forms
of
Thambetochen,
the
nene
still
closely
re-
sembles
its
nearest
continental
relative,
the
Canada
goose
(B.
canadensis).
In
ad-
dition,
fossil
finds
have
turned
up
some
in-
teresting
relatives
of
the
nene.
Remains
have
been
found
on
many
Hawaiian
is-
34
NATURAL
HISTORY
9/83

lands,
including
ones
where
no
nene
has
ever
been
recorded
alive.
Some
of
the
fos-
sil
skeletons
resemble
those
of
typical
modern
nenes;
others
belonged
to
bigger-
bodied
birds
with
shorter
wings,
birds
that
were
barely
capable
of
night,
if
at
all.
We
hope
thai
comparisons
of
the
various
fos-
sils
will
elucidate
patterns
of
speciation
during
the
early
stages
in
the
evolution
of
flightiessness.
Among
the
other
flightless
birds
found
on
the
islands
of
Hawaii
are
rails
and
ibises,
the
latter
being
familiar
to
most
people
as
long-legged
water
birds
with
long,
curved,
curlewlike
bills
and
excel-
lent
powers
of
flight.
Before
the
discovery
on
Molokai
and
Maui
of
the
remains
of
twoorthreespcciesofibisesthat
were
ab-
solutely
unable
to
fly,
no
one
had
given
the
first
thought
to
the
possibility
that
a
flight-
less
ibis
might
have
existed.
Yet
within
a
year
after
the
first
Hawaiian
ibises
had
been
described,
bones
of
another,
even
stranger,
flightless
ibis
from
Jamaica
were
found
in
the
American
Museum
of
Natu-
ral
History's
Department
of
Mammalogy,
scattered
among
drawers
of
fossil
mam-
mal
scraps.
The
leg
bones
of
our
Hawaiian
ibises
are
much
shorter
and
stoutcr.ihan
those
of
typical
ibises,
so
much
so
that
we
were
not
confident
they
belonged
to
an
ibis
until
most
of
an
associated
skeleton
was
found
in
a
lava
tube
on
Maui.
With
such
short
legs,
these
ibises
were
certainly
no
longer
water
birds
but
had
shifted
to
foraging
on
land.
They
must
have
made
their
living
fossicking
about
in
litter
much
like
the
New
Zealand
kiwis.
If
there
is
one
group
of
birds
that
pa-
leontologists
can
almost
always
expect
to
find
represented
by
flightless
species
on
unexplored
islands,
it
is
the
rails
(family
Rallidae).
Rails
are
well
known
for
their
ability
to
colonize
distant
islands,
and
on
many
of
these
islands,
flightless
species
have
evolved.
Most
flightless
rails
are
ex-
tinct,
but
at
one
lime
they
wcrea
common
feature
on
oceanic
islands.
At
¡east
two
species
survived
into
modern
times
in
the
Hawaiian
Islands:
one
on
the
island
of
Ha-
waii
and
one
on
the
tiny
northwest
atoll
of
Laysan.
Both
are
now
extinct.
The
fossil
record
shows
that
there
was
an
additional
Discover
the
uncoitimon
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E
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20
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December
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Our
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Arabia
Compared
with
a
flying
goose,
the
extinct
ftighüess
Thambetochen.
below,
had
stout
hind
¡imbs
and
a
massive
pelvis,
small
and
unscuiptured
wing
bones,
anda
small
sternum
with
no
keel.
The
angle
between
the
coracoid
and
scapula
is
wide,
a
trait
common
amongflightless
birds
but
never
found
in
adullßying
birds.
Some
of
the
siieletalfeatures
of
adult
flightless
birds
result
from
the
retention
of
juvenile
characters.or
neoteny.
This
hatchling
of
a
flying
goose,
bottom,
is
too
young
to
fly.
Notice
the
open
angle
between
coracoid
and
scapula,
reduced
wings,
and
absence
of
a
keel.
OOU^S
CrHTWF
Extinct
Flightless
Thambetochen
Scapula
Coracoid
Hind
Limb
Hatctiiing
of
/"""^
Ftying
Goose
f
^
species
on
Hawaii,
and
at
least
one
fiight-
less
rail
on
Kauai,
two
on
Oahu,
three
on
Maui,
and
one
on
Molokai,
a
species
that
was
smaller
than
any
other
rail
ever
found.
Why
so
many
species?
Archipelagoes
have
certain
characteristics
that
promote
speciation.
A
colonizing
species
that
in-
vades
an
archipelago
usually
spreads
to
most
or
all
of
the
islands
present.
Each
population
may
become
sedentary,
mini-
mizing
gene
flow
between
islands
and
al-
lowing
the
various
island
populations
to
drift
apart
genetically,
which
may
lead
to
speciation.
36
NATURAL
HISTORY
9/83

LEARN
BIRD
PHOTOGRAPHY
AT
HOME
Now
..
with
an
C3ttiiirtj¡
rtfw
ti)ur*i-
iiffLTin^
fmni
Corrt-ll
L^nivcrsit^'s
1^'^^ld-fa^lolJ^
tent«
ftïT
rhr
^utty
of
hirdlife
..
.
you
tan
team
rht
st-tTcLs
crrsiif;(-f:VK(ul
hirtl
|>hotuf(raphi-.
Siud>-;ii
horm.-.
ji
your
(m-n
pixx.
antí
n-tcii-f
iht
miwt
comprehensive
swurtt
tif
in/c^rmaTton
í>n
bird
ptiiHof^raphy
a^^tlahlr,
p\\iy
pcrv>njl
[u[<}ririg
of
your
wiirt
ihmü)(h
ll)f
awii^nmcnts
j«u
suhmJi.
V^nir
»tudy
iTiat(Tj»l?i
incliKif
J
tomplrtt
tcxi
with
ovvr
iwo
hun<3rctl
pti<)Co|;raphs
and
Jrawiii^f».
rvía^Ti
h^'
ntjittl
binJ
phi}t{)t^phL'r^
amJ
^Httitholti^isiü.
and
For
more
iníonTnaii{>n
i>n
hovk'
viui
can
cnn}Ei,
nil
out
ilur
c^upiin
and
wnd
il
hKÍay
lo
PhíHO
rA>Ljrsc.
DçTK.
\H.
Laburatorv
orOrniíholiit^y.
Japsuckcr
WOIKIï,
Ithaca.
IV^'
L-iK^
YES.
Picase
send
mr
Jnforniatlon
on
Ule
Bird
Fhocognphy
Cour».
NH-'>'KÍ
KiíW
.
(jly
./tp.
GeoTße
Gay
lord
Simpson
Penßuins
"One
of
tht
best
tKM)ks
on
the
penguin
that
I
have
read....
Delightful,
amusing,
and
iniormativc."
•
Gerald
Durrcll,
Harper's
•^A
book
for
every
animal
lover."
•
Roger
Caras,
CBS
35
b/w
+
6
color
illus.
S6,9S
pb.
Past
and
Present,
Hen
and
Then
Splendid
The
Curious
History
ISOlfftion
AfJlart
Mammals
"A
welcome
contriburion
from
the
field's
leading
aiithorit)'."
•
Larry
G.
Marshall,
Science
SS.9S
pb.
Send
orders
to
Dept,
SMS
Yale
University
Press
New
Hawn.CT
06520
The
South
American
rhea
belongs
to
an
assortment
of
flightless
birds,
known
as
ratites,
thai
are
distinguished
by
peculiarities
of
the
palate
and
often
considered
to
be
an
ancient
group
descended
from
a
common
ancestor.
The
new
fossil
evidence
from
Hawaii,
however,
suggests
that
many
of
the
ratites
may
have
evolved
independently
of
one
another.
In
time,
flying
birds
in
an
archipelago
can
recolonize
neighboring
islands,
so
that
a
single
island
could
eventually
be
host
to
numerous
species,
all
evolved
From
the
original
colonizers.
In
contrast,
popula-
tions
that
become
flightless
are
efí'ectively
held
captive
on
their
home
island.
The
only
easy
way
for
them
to
spread
to
nearby
islands
is
via
land
bridges,
which
rarely
form
between
oceanic
islands.
In
the
Ha-
waiian
Islands,
such
bridges
did
occur
be-
tween
the
islands
of
Maui,
Molokai,
and
Lanai
during
the
last
ice
age.
At
that
time,
so
much
water
from
the
world's
oceans
was
frozen
in
glaciers
and
the
polar
icecaps
that
sea
levels
worldwide
were
about
three
hundred
feet
lower
than
they
are
today.
The
shallow
channels
that
now
separate
Matii
from
Lanai
and
Lanai
from
Molokai
were
then
exposed
as
dry
land,
forming
a
single
large
island
called
Maui
Nui
(Big
Maui)
and
permitting
overland
dispersal
among
three
formerly
separate
islands.
While
there
can
be
little
doubt
that
such
dispersal
occurred,
estab-
lishing
precisely
how
the
joining
and
sub-
sequent
fractionation
of
Maui
Nui
has
in-
fluenced
the
distribution
of
a
particular
species
or
group
of
species
remains
frus-
tratingty
out
of
our
reach.
Since
land
bridges
are
rare,
mcst
flight-
less
species
never
had
the
opportunity
to
colonize
a
neighboring
island.
This
means,
significantly,
that
flightlessness
has
arisen
independently
again
and
again
in
the
Ha-
waiian
islands
and
an
untold
number
of
times
on
oceanic
islands
the
world
over.
It
is
a
common
evolutionary
response
to
the
conditions
of
life
on
oceanic
islands
of
all
sorts,
from
tiny
atolis
to
huge,
mountain-
ous
land
masses.
This
repeated
evolution
of
flightlessness
is
a
testimony
to
the
tre-
38
NATURAL
HISTORY
9/83

mendous
potential
of
organisms
to
inde-
pendently
evolve
convergent
or
parallel
adaptations
in
response
to
similar
selective
pressures.
Flightless
birds
are
extremely
vulner-
able
to
introduced
predators
and
other
hu-
man
disturbances,
and
an
untold
number
of
ihem
must
have
vanished
as
humans
discovered
and
settled
islands
everywhere.
Why,
with
ail
the
islands
in
the
tropical
Pacific,
some
of
them
larger
and
older
than
the
Hawaiian
Islands,
have
truly
bi-
zarre
flightless
birds
been
found
only
in
Hawaii,
New
Zealand,
and
New
Caledo-
nia
(home
of
the
kagu)?
In
our
minds,
the
lack
of
examples
reflects,
not
reality,
but
our
ignorance.
Without
doubt,
flightless
birds
did
exist
on
most
major
islands
in
the
Pacific.
The
early
Hawaiians,
through
habitat
destruction,
prédation,
and
the
in-
troduction
of
foreign
plants
and
animals,
drove
at
least
seventeen
species
of
flight-
less
birds,
and
no
fewer
than
twenty-eight
flying
species,
to
extinction.
There
is
no
reason
to
believe
that
prehistoric
human
settlement
did
not
also
bring
about
great
changes
in
the
avifauna
of
other
Pacific
is-
lands.
The
coming
years
may
prove
to
be
a
golden
age
for
paleontologists
exploring
the
Pacific
for
bird
bones.
From
our
studies,
we
can
envision
the
Hawaiian
Islands
at
a
time
in
the
past,
when
eagles
soared
over
unbroken
forests
that
swarmed
with
an
undreamed-of
vari-
ety
of
small
birds,
when
troops
of
un-
gainly,
waddling
flightless
geese
grazed
on
plants
no
botanist
has
ever
seen,
and
when
flightless
ibises
probed
among
litter
for
insect
species
that
never
felt
the
thrust
of
an
entomologist's
pin.
The
material
evi-
dence
for
this
former
glory
now
consists
mainly
of
tray
upon
tray
of
bare
bones.
But
we
are
the
lucky
ones.
The
botanists
and
entomologists
will
unfortunately
never
know
what
they
are
missing,
D
The
flightless
birds
of
Hawaii,
particu-
larly
the
geese
and
ibises,
raise
some
per-
plexing
questions
about
the
evolution
of
another
assortment
of
flightless
birds,
the
so-called
ratitcs.
These
include
running
birds,
such
as
ostriches,
rhca.i,
and
emus;
the
giant,
lumbering
extinct
moas
of
New
Zealand;
and
the
little,
long-billed
kiwis
of
shoe
polish
fame.
Many
scientists
con-
sider
ratites
to
be
an
ancient
group,
de-
scended
from
a
single
common
ancestor,
that
is
separable
from
all
other
birds
by
the
palcognathous
palate,
a
peculiar
con-
figuration
of
the
bones
in
the
roof
of
the
mouth.
The
Hyvi-aiian
fassils,
however,
seem
to
suggest
that
at
least
some
of
the
ratites
may
have
originated
more
recently
from
more
typical
flying
birds.
The
term
raiite,
from
the
Latin
ralis
for
raft,
was
originally
applied
to
these
birds
because
their
sternum
lacks
a
keel.
But
[he
keel
may
also
be
greatly
reduced
or
even
absent
in
flightless
birds
that
are
clearly
not
ratites.
in
some
of
the
Hawai-
ian
geese,
the
sternum
has
no
trace
of
a
keel
and
has
lost
all
resemblance
to
the
sternum
in
typical
ducks
and
geese;
in-
stead,
it
is
rounded
and
bowllike,
more
like
the
sternum
of
an
ostrich
or
an
emu.
In
some
of
the
ratites•
moas,
for
exam-
ple•the
bones
in
the
leg
are
relatively
shorter
and
much
stouter
than
in
running
ratites.
Again,
wcsec
very
similar
adapta-
tions
in
the
flightless
birds
of
Hawaii.
Some
of
the
geese
had
ponderous
legs
that
contrasted
greatly
with
their
little
wings.
The
fiightlc.ss
Hawaiian
ibises
had
leg
bones,so
much
stockier
than
those
of
their
long-legged
wading
ancestors
that
they
Lessons
from
a
Flightless
Ibis
are
remarkably
similar
in
proportion
to
those
in
the
New
Zealand
kiwis.
We
find
that
almost
all
the
features
in
the
body
plan
of
ratites
have
evolved
re-
peatedly
in
a
variety
of
nonratites
through
the
retention
of
juvenile
traits
(neoteny).
Might
the
paleognathous
palate
likewise
be
a
result
of
neoteny,
and
consequently
something
that
could
theoretically
de-
velop
in
many
kinds
of
birds?
Thi.s
is
the
subject
of
current
investigation,
for
if
all
of
the
characteristics
of
the
paleognath-
ous
palate
are
present
in
the
embryonic
condition,
as
some
of
them
are
known
to
be,
the
various
kinds
of
ratites
might
have
evolved
independently
of
one
another
by
retaining
such
a
palate
into
adulthood.
With
this
possibility
in
mind,
the
strik-
ing
parallels
between
the
flightless
birds
of
the
Hawaiian
Islands
and
those
of
New
Zealand
become
of
even
greater
interest.
Hawaii
had
an
extensive
radiation
of
gooselike
birds,
derived
from
at
least
two
dilfcrcnt
ancestors,
that
were
flightless
Great
spotted
kiwi
of
New
Zealand
Brian
Ëntinç.
pnotc
fíoi^stctvrí
and
had
widely
divergent
bill
shapes,
evi-
dently
for
feeding
on
diJTerent
kinds
of
vegetation.
Similarly,
in
New
Zealand,
the
moas
were
large,
flightless,
herbivo-
rous
birds
with
a
variety
of
bill
shapes.
Taxonomists
have
at
times
divided
the
moas
into
two
different
families
that
might
actually
represent
separate
coloni-
zations
of
New
Zealand
by
flying
ances-
tors.
The
resemblance
between
living
ki-
wis
and
the
extinct
Hawaiian
ibises
are
not
restricted
to
leg
bones;
both
also
have
long,
probing
bills
quite
unlike
those
of
geese
or
moas.
The
flightless
birds
in
the
main
Hawai-
ian
Islands
lost
the
ability
to
fly
in
con-
siderably
less
than
6
million
years.
The
flightless
ibises,
for
example,
which
were
restricted
to
{^e
islands
of
Maui
Nui,
could
not
have
colonized
those
islands
more
than
1.8
million
years
ago,
the
age
oí
the
oldest
rocks
found
on
Motokai,
the
oldest
of
the
Maui
Nui
group.
But
New
Zealand
has
been
available
for
coloniza-
tion
by
vagrant
birds
for
tens
of
millions
of
years.
If
an
¡bis
could
become
almost
un-
recognizable
in
less
than
1.8
million
years,
what
would
it
look
like
if
it
had
continued
to
evolve
for
another
10
or
20
million
years?
Are
the
moas
and
kiwis
of
New
Zealand
merely
the
descendants
of
geese,
ducks,
and
ibi.ses
that
had
much
more
time
to
diverge?
Such
intriguing
ques-
tions
would
probably
never
have
been
raised
had
the
Hawaiian
fossils
gone
un-
discovered.
Perhaps
more
work
with
fossil
birds
from
other
Pacific
islands
will
help
us
find
the
answers.
Storrs
L.
Olson
40
NATURAL
HISTORY
9/83