THE 1/CRM_1_1996...THE GALAPAGOS TORTOISES NOMENCLATURAL AND SURVIVAL STATUS PETER C.H. PRITCHARD...
82
Chelonia Institute 6 CHELONIAN RESEARCH MONOGRAPHS Number 1 — July 1996 Published by Chelonian Research Foundation in association with Conservation International and Chelonia Institute CONSERVATION INTERNATIONAL
Transcript of THE 1/CRM_1_1996...THE GALAPAGOS TORTOISES NOMENCLATURAL AND SURVIVAL STATUS PETER C.H. PRITCHARD...
TH
EG
AL
AP
AG
OS
TO
RT
OIS
ES
NO
ME
NC
LA
TU
RA
LA
ND
SU
RV
IVA
LS
TA
TU
S
PE
TE
RC
.H.
PR
ITC
HA
RD
Chelonia
Institute
6
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
AP
HS
Num
ber1
—July
1996
Published
byC
helonianR
esearchF
oundationin
associationw
ithC
onservationIn
ternatio
nal
andC
heloniaInstitute
CO
NS
ER
VA
TIO
NIN
TE
RN
AT
ION
AL
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
AP
HS
Contrib
utio
ns
inT
urtle
and
To
rtoise
Research
Series
Edito
rs
AN
DER
SG
.J.R
HO
DIN
Chelonian
Research
Foundation
168G
oodrichS
treetL
unenburg,M
assachusetts01462
USA
PET
ER
C.H
.PR
ITCH
AR
DF
loridaA
udubonSociety
1331P
almetto
Avenue,
Suite110
Winter
Park,
Florida
32789U
SA
Jow
iL
.B
EH
LE
RD
epartment
ofHerpetology
Wildlife
Consenation
SocietyB
ronx,N
ewY
ork10460
USA
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
AP
HS
(ISSN1088-7105)
isan
intern
ation
alpeer-rev
iewed
scientific
publicationseries
form
onograph-lengthm
anuscripts,collected
symposia,
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pilations,and
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The
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anyaspects
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research,w
itha
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conservationor
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anuscriptsdealing
with
conservationbiology,
systematic
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diversity,geo
graphicdistribution,naturalhistory,ecology,reproduction,m
orphologyand
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hu
man
exp
loitatio
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iblio
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erreference
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Pu
blish
edIssues
1.T
he
Galap
ago
sT
orto
ises:N
om
enclatu
ralan
dS
urv
ival
Statu
s.1996.
By
PETERC
.H.
PRITC
HA
RD
.85
pp.IS
BN
:0-9
65
35
40
-0-8
(hardcover);
0-9653540-1-6(soft
cover).
Issuesin
Pro
du
ction
2.T
he
Co
nserv
ation
Bio
logy
of
Fresh
water
Tu
rtles.P
ub
lication
pla
nnedfo
rlate
1996.E
dited
byPETER
C.H
.P
RIT
CH
AR
DA
ND
AN
DER
SG
.J.R
HO
DIN
.T
wo
vo
lum
es.
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
APH
Sare
availablefo
rpu
rchase
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helonianR
esearchF
oundation.No
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oodrichS
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ail:R
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Copyright
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oundation.
Date
ofissue:26
July1996
The
Galapagos
Tortoises:
Nom
enclatu
ralan
dS
urvivalS
tatus
PE
TE
RC
.H.
PR
ITC
HA
RD
I
The
Galapagos
Tortoises:
Nom
enclaturalan
dS
urvivalS
tatus
PE
TE
RC
.H.
PR
ITC
HA
RD
Florida
Audubon
Society,1331
Palm
ettoAvenue,
Suite110,
Winter
Park,
Florida
32789U
SA
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
AP
HS
Num
ber1
July1996
Chelonian
Research
Foundatio
n
I
Fro
nt
Co
ver:
The
cover
pictu
reof
Onan,
aceleb
ratedold
male
torto
isefrom
Du
ncan
Island
(Pin
zón),
was
pain
tedby
Naoko
Izuhara,
anartist
residen
tin
Win
terP
ark,F
lorid
a.T
hep
aintin
gis
based
upona
colo
rp
ho
tog
raph
taken
bythe
auth
or
inA
ug
ust
1986.T
hestory
ofO
nan,a
nearly
blin
dbut
agg
ressive
andsp
iritedanim
alth
ou
gh
tto
bew
ellover
acen
tury
inage,
istold
byS
witak
(1995).T
henam
eO
nan,given
tothe
torto
iseby
Craig
MacF
arland
inthe
early1970s,
deriv
esfrom
theB
iblical
Onan
(Gen
esis38:9),
who
incu
rredthe
wrath
ofG
odand
was
slainfor
spillin
ghis
seedupon
theground.
The
chelo
nian
Onan
toohas
beenobserv
edattem
ptin
gto
copulate
with
inan
imate
objects,
inclu
din
gro
cks.
Sadly
butin
evitab
ly,
Onan
was
found
deadw
ithin
hiscu
stom
aryh
aun
tsby
Gayle
Davis
inM
ay1990.
His
remain
sw
ereb
uried
onD
uncan.
Fro
ntisp
iece:T
hispain
ting
of
bu
ccaneers
andG
alapag
os
torto
isesby
Charles
M.
Sh
eldon
appeared
inP
ioneers
inT
rop
icalAm
ericaby
Sir
Harry
Joh
nsto
n(u
ndated
,ca.
1912).T
heveg
etation
issom
ewhat
fancifu
l,in
that
theO
puntia
andJasm
ino
cereus
seemto
bem
iniatu
rized,
andA
loeand
Melo
cactus
arenot
known
fromthe
Galap
ago
s.T
hegeo
grap
hic
features
arerem
iniscen
tof
thevalley
leadin
gup
fromT
agusC
oveto
ward
sV
olcan
Darw
inon
Alb
emarle
Island
(Isabela).
Ho
wev
er,the
torto
isein
thefo
regro
un
d,
with
itssad
dleb
acked
shelland
up
righ
t,th
ickneck,
isa
goodrep
resentatio
nof
theto
rtoise
of
Abin
gdo
nIslan
d(P
inta).
The
po
sture,
theshell
mo
rpholo
gy,
andeven
theg
roo
ve
alongthe
undersid
eof
then
eck(an
artifactof
thepreserv
ation
andm
ountin
gp
rocess,
unknown
inliv
ing
torto
ises),are
virtu
allyid
entical
toone
ofthe
two
mo
un
tedsp
ecimen
sof
Ab
ing
do
nto
rtoises
inthe
British
Museum
ofN
atural
Histo
ry(B
MN
H76.6.21.38—
39),co
llectedby
Co
mm
ander
J.C
ooksonin
1875(see
Fig.
23).
Back
Cover
(Upper):
This
nin
eteenth
centu
ryen
grav
ing
of
unknown
pro
ven
ance
po
rtrays
wh
aleboats
closeto
theshore
of
Alb
emarle
Island
(Isabela).
The
precip
itous,
rawn
ature
of
theterrain
resemb
lesV
olcan
Wo
lfin
thenorth
;but
Cerro
Azul
inthe
sou
thw
estis
nearly
assteep
,and
theru
gg
edness
po
rtrayed
may
havebeen
enh
anced
bythe
artist.
Back
Cover
(Low
er):T
heposterio
rend
of
thebony
carapace
of
alarg
em
aleto
rtoise
fromthe
To
rtoise
Reserv
eon
Indefatig
able
Island
(San
taC
ruz).T
helast
word,
asit
were
—the
end.
Contribution
No.
199of
theC
harlesD
arwin
Foundation
forthe
Galapagos
Islands
The
Galapagos
Tortoises:
Nom
enclaturaland
Survival
Status.B
yP
ET
ER
C.H
.P
RIT
CH
AR
D.
CH
EL
ON
IAN
RE
SE
AR
CH
MO
NO
GR
AP
HS
,Num
ber1.
Edited
byA
ND
ER
SG
.J.R
HO
DIN
.
ISS
N(m
onographseries):
1088-7105IS
BN
(thisvolum
e):0-9653540-0-8
(hardcover);
0-9653540-1-6(soft
cover)
Published
andC
opyright©
i996by
Chelonian
Research
Foundation,L
unenburg,Massachusetts,U
SA
.Allrights
reserved.N
opart
ofthis
bookm
aybe
usedin
anym
annerw
ithoutwritten
permission
exceptin
thecase
ofbriefquotations
embodied
inarticles
andreview
s.P
rintedby
MT
CP
rinting,Inc.,
Leom
inster,M
assachusetts,U
SA
.
TA
BL
EO
FC
ON
TE
NT
S
ED
ITO
RIA
LP
RO
LO
GU
E7
Tortoises
inthe
Mist.
By
AN
DE
RS
G.J.
RH
OD
IN9
The
Darw
inC
onnection11
AB
ST
RA
CT
13
AC
KN
OW
LE
DG
ME
NT
S15
INT
RO
DU
CT
ION
17
HIS
TO
RIC
AL
OV
ER
VIE
W21
Discovery
Phase
21T
axonomic
Phase
21R
escueP
hase23
InSitu
Phase
24
NO
ME
NC
LA
TU
RA
LS
TA
TU
S27
Garm
an’5
The
Galapagos
Tortoises
27C
urrentN
omenclature
34S
umm
aryof
Proposed
Nom
enclature49
Synonym
yof
Geochelone
(Chelonoidis)
nigra49
SU
RV
IVA
LS
TA
TU
S52
Historical
Review
ofG
alapagosT
ortoiseP
opulations52
Abingdon
(Pinta)
52D
uncan(P
inzón)55
Hood
(Española)
58C
hatham(S
anC
ristóbal)59
Indefatigable(S
antaC
ruz)60
Narborough
(Fernandina)
63C
harles(S
antaM
ariaor
Floreana)
63Jervis
(Rábida)
65B
arrington(S
antaFe)
65Jam
es(San
Salvador
orS
antiago)66
Albem
arle(Isabela):
Volcan
Wolf
67A
lbemarle
(Isabela):V
olcanD
arwin
69A
lbemarle
(Isabela):V
olcanA
lcedo71
Albem
arle(Isabela):
Sierra
Negra
andC
erroA
zul74
CO
NC
LU
SIO
NS
79N
ornenclaturalS
tatus79
Survival
Status
79
LIT
ER
AT
UR
EC
ITE
D81
ED
ITO
RIA
LP
RO
LO
GU
E
Itake
great pleasurein
presentingthe
following
mono-
graphicw
orkby
Peter
C.H
.Pritchard.P
eterand
I havebeen
goodfriends
andprofessional
colleaguessince
them
iddle1970s
andduring
thistim
ew
ehave
hadthe
marvelous
fortuneto
experiencethe
magic
ofthe
Galapagos
Islandstogether
ontw
ooccasions.
The
firstofthosevisits,in
1982,w
asform
ethe
kindofexperience
thatcanonly
bedescribed
asone
oflife’s
definingm
oments.
Touring
thearchipelago
firstw
itha
groupof
ecotourists
organizedby
Florida
Audubon
Society
andled
byP
etergavem
ean
overviewand
achance
toappreciate
thefull
experienceof
theG
alapagos.T
heincredible
andvaried
topography,the
teeming
seas,theprofusion
ofbirdlife,
andeveryw
herethe
evidenceof
what
nature’sm
agnificenceoffers
inits
variedflora
andfauna
impressed
me
beyondw
ords.O
fgreatest
interesttom
e,of
course,w
asthe
herp
etologicalfauna.T
heland
iguanas,marine
iguanas,and
lavalizards
allheldincredible
fascinationand
Ispent along
time
marveling
attheirprofusion
anddiversity.
Butas
expected,and
without
adoubt,
itwas
thegianttortoises
thatcaptured
my
imagination
andpropelled
me
intothe
kindof
reveriefrom
which
onenever
trulyrecovers.
Afterthe
formaltourofthe
islands,Peterand
Iwere
ableto
visitseveraltortoiseareas,w
hereIw
asable
tofocus
fullyon
thechelonian
experience.Itwas
transcendenttosee
theseliving
reminders
ofancient
times,
tosit
with
themin
quietsolitude,
andto
sharew
iththem
them
editativestate
ofan
unhurriedexistence
inan
everm
oreintrusive
world.
We
firstvisitedthe
Tortoise
Reserve
southwestofS
antaR
osaon
IndefatigableIsland
(Santa
Cruz).
Here
theab
un
danthugedom
e-shelledtortoises
livein
lush,verdantforestsand
asI
sharedtheir
world
with
themfor
ashortw
hileI
feltas
ifI
hadbeen
transportedin
atim
ecapsule
backto
some
Cretaceous
landscape.Asoftrain
fellfromtim
eto
time,the
luxuriantvegetation
andtortoise
carapacesglistened,
andI
feltprivileged
toexperience
them
oment.
We
nextvisited
Duncan
Island(P
inzón).T
hisisland
istotally
different,w
itha
hot,xeric
habitatof
sunbakedcacti
andsparse
brush.T
hetortoises
hereare
small
andsaddlebacked,
unlikethe
giantson
Indefatigable.A
ndthey
arehard
tofind.
We
spenta
longtim
ehiking
intothe
highareas
where
thenative
tortoiseslive,and
thensearched
longand
hardbefore
we
founda
fewold
animals
hidingunder
rocksor
brush.T
heirancient
wrinkled
facesand
smooth
worn
shellsspoke
ofa
lifetime
ofstruggle
forexistence
intheir
harshenvironm
ent.A
sI
saton
thehigh
hillsidein
theirpresence
andgazed
outpastthe
dryveg
etation
andrough
landscapeto
theblue
waters
ofthesurround-
ingocean
belowIfelta
senseofpeace
andcontentm
ent.This
was
am
agicalplace
andits
chelonianinhabitants
imparted
asense
oftim
elessnessto
itsstark
beauty.A
placew
edid
notvisittogether,butthatIhad
hopedto,
was
thecaldera
andcrater
rimof
Volcan
Alcedo
onA
lbemarle
Island(Isabela).
Here
thepopulation
ofto
rtoises
was
them
ostintact
—the
most
primeval
—in
theentire
archipelago.D
ifficultto
reachfrom
thecoast
andthereby
protectedfrom
slaughterby
nineteenthcentury
whalers,
thepopulation
was
alsoisolated
fromsouthern
Albem
arleby
thelava
flows
ofP
erryIsthm
us,w
hichhad
preventedthe
influxof
feralgoatsto
theregion.
Recently
andom
inously,how
ever,goats
havesucceeded
incross-
ingthis
barrierand
arenow
seriouslythreatening
thetortoise
populationthrough
widespread
habitatd
estruc
tionas
theirnum
bersm
ultiplyastronom
ically.A
cam-
paignto
savethe
tortoisesof
Alcedo
hasrecently
beeninitiated
bythe
Charles
Darw
inF
oundation.W
eallhope
thattheir
effortsare
successfulin
savingthese
tortoisesso
thatour
childrenand
theirdescendants
will
deriveas
much
joyand
inspirationfrom
theirsplendor
asw
ehave.
Our
passionfor
thesem
agnificentcreatures
must
driveour
continuingefforts
tobetter
understandand
protectthem
.M
ayour
legacyto
thefuture
bethat
we
succeeded.Peter
C.H
.Pritchard
with
alarge
male
gianttortoise
(140cm
carapacelength
overthecurve)
inthe
Tortoise
Reserve
southwest
ofSanta
Rosa
onIndefatigable
Island(Santa
Cruz),
September
1982.Photo
byA
ndersG
.J.R
hodin.A
ND
ER
SG
.J.R
HO
DIN
Tortoises
inthe
Mist
Tortoises
inthe
mist
timeless
creatures,m
oisture-kissedneath
shroudedtrees
andm
ossylace
sinceeons
past,still
exist.
On
craterrim
,a
magic
placetortoises
move,
attheir
pacefrom
calderafloor
belowponderous
steps,gentle
grace.
Ringed
byfire,
lavaflow
sspared
thew
halers’deadly
blows
surviveserene,
Alcedo
home
enchantedisles,
Galapagos.
Anders
G.J.
Rhodin
lOA
pril1996
Tortoises
inthe
mist
onthe
craterrim
abovethe
calderaof
Volcan
Alcedo,
Albem
arleIsland
(Isabela),July
1986.D
uringdry
periodstortoises
migrate
fromthe
drycaldera
floorup
tothe
craterrim
where
them
ist(garua)
ofthe
cloud-catchment
dripzone
providesthem
them
oisturetheyneed.S
eethepreceding
editorialprologueand
theaccounton
pp. 71—73
forfurtherdetails.Photo
byP
eterC.H
. Pritchard.
The
Darw
inC
onnection
“1have
notas
yetn
oticed
byfar
them
ostrem
arkab
lefeatu
rein
then
atural
histo
ryof
thisarch
ipelag
o;
itis,
that
thedifferen
tislan
ds
toa
con
siderab
leex
tent
arein
hab
itedby
adifferen
tset
ofb
eing
s.M
yatten
tion
was
firstcalled
tothis
factby
theV
ice-Go
vern
or,
Mr.
Law
son
,d
eclaring
thatthe
torto
isesdiffered
fromthe
differen
tislan
ds,
andth
athe
could
with
certainty
tellfrom
which
island
anyone
was
bro
ug
ht...
The
inhab
itants,
asI
havesaid,
stateth
atthey
candistin
guish
theto
rtoises
fromthe
differen
tislan
ds;
andthat
theyd
iffernot
onlyin
size,but
ino
ther
characters.
Cap
tainP
orter
hasd
escribed
tho
sefrom
Ch
arlesand
fromthe
nearest
island
toit,
namely,
Hood
Island,
ashaving
their
shells
infro
nt
thick
andtu
rned
uplik
ea
Sp
anish
saddle,w
hilst
theto
rtoises
fromJam
esIslan
dare
rounder,
black
er,and
havea
better
tastew
henco
ok
ed.”
—
Charles
I)arwin
(1845,Jo
urn
alof
Research
es)
..
v_
-\‘,
/%
\
-.-
-?‘&(,
“Iknew
Darw
in.
Nice
guy.”S
auers
(ca.1980.
The
New
Yorker)
AB
ST
RA
CT
PR
ITC
HA
RD
,P
ET
ER
C.H
.1996.
The
Galap
agos
Torto
ises:N
om
enclatu
ralan
dS
urv
ival
Statu
s.C
helonianR
esearchM
onograp
hs
1:1—85.
The
historyof
human
attentionto
Galapagos
tortoisesis
brokendow
ninto
fourphases.
These
are:the
Discovery
Phase
(1535—1840),
duringw
hichthe
islandsw
erediscovered
andearly
visitorsrecorded
theirobervations,
oftenw
ithconsiderable
attentionto
thegiant
tortoises;the
Taxonom
icP
hase( 1824—
1917),duringw
hichspecim
ensof the
variousisland
tortoisepopulations
(many
lackingcollecting
data)reached
major
museum
sand
were
describedand
named;
theR
escueP
hase(1925—
1955),during
which
zoologicalvisitors
concludedthat
thetortoises
were
doomed
intheir
nativehabitat
andcould
onlybe
savedby
captivebreeding
inzoological
institutions;and
theIn
Situ
Phase
(1957—present),philosophically
areversal
ofth
eR
escueP
hase,duringw
hichincreasing
effortsw
erem
adeto
conserve,protect,
andprocreate
thetortoises
inthe
Galapagos
Islandsthem
selves.T
echniquesused
duringthe
InS
ituP
haseincluded
environmentaleducation
programs
forG
alapagosresidents
andsettlers,
controlof
feralm
amm
alsdepredating
orcom
petingw
iththe
tortoises,protection
ofw
ildtortoise
nestsw
ithrock
cairnsor
transferof
neststo
artificialincubation,
head-starting
ofyoung
tortoisesthrough
thefirst
yearsof
life,and
closed-cyclecaptive
breeding.F
urthermore,
theestablishm
entof
Galapagos
National
Park
gaveat least
nominal
legalprotection
totortoise
habitatthroughout
thearchipelago.
The
scientificnom
enclatureof
Galapagos
tortoisesis
examined
andfound
tohave
many
problems.
The
recentlycom
monly
usednam
e,G
eocheloneelephantopus,
isnot
theoldest
availablenam
efor
thespecies
andshould
nolonger
beused.
The
oldestnam
efor
Galapagos
tortoisesis
Testudo
californ
iana,
andunder
currentT
CZ
Nregulations
thisgenerally
unfamiliar
name
would
needto
beutilized
unlesspetition
ism
adefor
itssuppression.
Inthe
interim,
theoldest
availablefam
iliarnam
efor
allG
alapagostortoises
isT
estudonig
raQ
uoyand
Gaim
ard,1824,
andits
useis
herebyadvocated
asthe
combination
Geochelone
(Chelonoidis)
nigra.T
hevarious
forms
fromthe
differentislands
areallconsidered
assubspecies
ofG
.nigra.
Itisalso
necessaryto
petitionthe
ICZ
Nto
suppressthe
name
Testudo
ephippium,
asenior
synonymof
Testudo
abingdonifrom
Abingdon
Island,but
thathas
beenutilized
extensivelyfor
theD
uncanIsland
tortoise.O
nA
lbemarle
Island,the
names
ofthe
variouspopulations
anddescribed
subspeciesof
allbut
thenorthernm
ostvolcano
aresynonym
izedunder
Geochelone
nig
ravicina,
onthe
groundsthat
thedescribed
differencesare
eitherattributable
toenvironm
entaldifferences
(especiallyof
rainfall,food
availability,and
humidity),
ordo
notshow
geographiccorrelation
butare
artifactsof
ageand
sex.S
everalpotential
butunnam
edsubspecies
arelisted,
includingthose
ofnortheastern
Chatham
Island,northw
esternIndefatigable
Island,and
theextinct
forms
thatm
ayonce
haveinhabited
Barrington
andJervis
islands.P
articularattention
ispaid
toan
analysisof
theargum
entsand
conclusionsof
Garm
an(1
917),
thisbeing
thelast
comprehensive
analysisof
thetaxonom
yof
Galapagos
tortoisesand
onethat
hasnever
beenform
allychallenged
despitem
anydeficiencies,
asw
ellas
certainrem
arkableexam
plesof
insightor
prescience.T
hesurvival
statushistory
ofeach
ofthe
tortoisepopulations
fromthe
time
offirst
discoveryto
thepresent
(oruntil
thetim
eof
extinction)is
reconstructedto
theextent
thatthe
written
recordperm
its.It
isobserved
thatconsiderable
conservationprogress
hasbeen
made
inthe
decadessince
theIn
Situ
Phase
startedin
1957,but
inthe
lasttw
oor
threeyears
enormous
andpossibly
insurmountable
challengeshave
arisenin
theform
ofa
generalbreakdow
nof
respectfor
con
serva
tionlaw
inthe
archipelago.
KE
YW
OR
DS
.—
Reptilia;
Testudines;
Testudinidae;
Geochelone
nig
ra;tortoise;
turtle;co
nserv
ation;
managem
ent;repatriation;
head-starting;introduced
species;extinction;
survivalstatus;
nomenclature;
taxonomy;
distribution;
natu
ral
histo
ry;
Ecu
ado
r;G
ala
pago
s
AC
KN
OW
LE
DG
ME
NT
S
My
heartfeltthanks
areoffered
tothe
mem
oriesof
fourtruly
greatm
enw
ho,a
generationago,
tooktim
eto
encouragea
youngO
xfordchem
istryundergraduate
totravel
tothe
Galapagos
Islandsand
tom
akeobservations
ontheir
giant tortoises.T
heyw
ereP
rofessorV
ictorvan
Straelen. D
r. David
Lack,
Sir
Peter
Scott,
andS
irA
listerH
ardy.F
orseveral
yearsm
yfield
work
inthe
Galapagos
Islandsw
asfinanced
bythe
Worldw
ideF
undfor
Nature
(thenthe
World
Wildlife
Fund,
U.S
.N
ationalA
ppeal).A
lthoughthis
work
was
primarily
onsea
turtles,I
utilizedthe
opportunityto
visitevery
islandin
thearchipelago
fromw
hichtortoises
hadever
beenrecorded
—and
arrivedon
Abingdon
just
intim
eto
witness
therem
ovalof
thelast
livingtortoise
fromthat
island.D
uringthese
visitsI
receivedm
anycourtesies
froma
successionof
Charles
Darw
inR
esearchS
tationD
irectors,especially
Roger
Perry,
Peter
Kram
er,and
Craig
MacF
arland.I
would
alsolike
tothank
several veteranG
alapagosscientists,
includingD
aniel Weber,
Tjitte
deV
ries,M
ichaelH
arris,and
Ole
Ham
ann,for
sharingtheir
insightsinto
Galapagos
biology—
andlogistics.
Asuccession
ofb
oat captains
also, especiallyJohnny
Angerm
eyer, Fritz
Angerm
eyer,Julian
Fitter,
Herm
ogenesM
oncayo,José
Villacis,
andthe
late“C
ocaC
ola,”provided
reliablecharter
service—
acrucial
factorduring
fieldw
orkon
waterless
anduninhabited
islands.A
rnaldoT
upiza,José
Villa,
and“P
escadito”guided
me
toextrem
elyrem
otegalap
aguera
areasof
southernA
lbemarle,
Chatham
island,and
Hood
Island.respectively.
Ithank
Miguel
Cifuentes
forserving
asfield
assistanton
my
expeditionsof
I971
and1972,
andfeel
pridethat
heso
quicklythereafter
became
anoutstanding
Director
ofthe
Galapagos
National
Park.L
indaC
ayotvery
generouslyshared
bothher
time
andunrivaled
knowledge
ofthe
currentsurvival
statusof
theG
alapagostortoises,
critiquinga
draftof
them
anuscriptand
steeringm
etow
ardsliterature
thatI
hadoverlooked.
Hobart
Sm
ithand
Roger
Bour
alsoprovided
masterful
critiquesof
them
anuscriptand
gavem
ethe
benefitof
theirlegendary
insightsinto
theC
odeof
Zoological
Nom
enclature.In
all cases, theguidance
theyoffered
constitutedpeer
reviewatits
finest,and
anyquality
thatm
aybe
foundin
thefinished
work
owes
much
tothese
threecolleagues.
Ith
ankJean
Lescure
ofth
eM
useumN
ationald’H
istoireN
aturelle(P
aris)and
AS
.C
larkeof
theR
oyalS
cottishM
useum(E
dinburgh)for
supplyingphotographs
ofcrucial
typespecim
ens,and
forperm
issionto
publishthem
.A
llother
photographsare
bym
yselfexcept
fora
fewby
Rollo
Beck,
Roger
Bour,
Anders
Rhodin,
andD
avidS
nowthat
arenoted
inthe
figurelegends.
My
wife,
Sibille,
traveledw
ithm
eon
most
ofmy
Galapagos
expeditions,and
herpresence
made
me
am
uchhappier
camper
inm
anyw
ays.F
inally,I
thankm
yfriend
Anders
Rhodin,
Director
ofC
helonianR
esearchF
oundation,for
editingand
publishingthis
work.
Itis
arare
pleasureto
haveone’s
compositions
editedby
onew
honot
onlyknow
sthe
subjectm
atterintim
atelybut
who
alsohas
asuperlative
graspof
theprinciples
ofE
nglishgram
mar
andsyntax.
INT
RO
DU
CT
ION
There
ism
uchthat
isironic,
surprising,or
evenp
aradoxical about the
Galapagos
Islandsandtheir fam
ousgig
an
tictortoises.
The
associationbetw
eenthe
islandsand
thegiant
tortoisesseem
sso
absolute,and
indeedis
foreverform
alizedin
thevery
name
ofthe
archipelago.Y
etthe
Spanish
word
galdpagodoes
notrefer
exclusivelyto
largechelonians,
oreven
justto
terrestrialones.
The
Co
nq
uista
dors,possessed
ofor
acquiringfrom
thenative
Am
en-cans
—a
modest inventory
ofvemacular
names
fortortoisesand
turtle
s(to
rtuga,caguw
na,jic
ote
a,g
ald
pag
o,rn
orro
coyo)
parcelledthem
outto
thenew
chelonianspecies
asthey
fcundthem
intheir
New
World
territories.T
hegiant
tor
toises,the
most
remarkable
inhabitantsof
thenew
Pacific
archipelagodiscovered
bythe
off-courseB
ishopof P
anama
in1535, and
ina
real sensea
first-wave
ofarmored
Conquis
tadorsthem
selves,becam
egaldpago,
butso
dida
comm
onside-necked
pondturtle
inV
enezuela,for
example,
thatrarely
exceedsa
lengthof
30cm
.A
ndthere
isan
evensm
aller,sem
i-aquaticturtle
inE
nglish-speakingT
rinidadthat,
tothis
day,is
calledgalap.
Moreover,
thevery
existenceof
thelargest tortoises
inthe
world
onm
inusculeislands
inthe
vastP
acificO
ceanstrikes
many
asodd
—creatures
that represent theepitom
eofterrestriality, that cannot dive
andbarely
swim
, inoceanic
islandecosystem
sw
hereno
freshwater
turtlespecies
hasever penetrated. A
lmost all big
landanim
alsneed
bigspaces
elephantsin
thevast plains
ofeastAfrica,
or bisonon
theA
merican
prairies,com
eto
mind.
What
isthe
linkbetw
eengigantic
tortoisesand
small islands?
How
didthey
get there,crossing
vastocean
barriers,w
henaquatic
turtles(apart
fromthe
trulym
arinespecies)
failed?H
owcom
ethe
Galapagos,
with
theirspectacular
herpetofaunaand
gen
erally
absent terrestrial mam
malian
fauna,can
soevocatively
mim
icM
esozoictim
es—
theglorious
Age
ofR
eptiles—
while
being(the
geologiststell
us)so
new?
How
isit
thatsuch
aunique
andslow
-growing, slow
-breedinglife
formas
agiant
tortoisecould
haveevolved,
andreached
giganticsize,
sorapidly?
And,
while
thevulnerability
ofsuch
huge,slow
creaturesto
mankind
isunderstandable,
why
shouldsuch
impregnable
behemoths
beso
easilyharm
ed,or
evenextirpated,
bysm
allerm
amm
als—
evenblack
rats,of
allthings, w
henlesser
tortoiseshave
coexistedw
ithsm
all mam
mals
incontinental
ecosystems
form
anym
illionsof years?
These
questionsare
answerable.
Firstly,
thelink
be-tw
eengiant tortoises
andsm
all oceanicislands
isillusory, or
atleast
anartifact
ofstrictly
modern,
human-dom
inatedconditions.
Huge
terrestrialchelonians
usedto
bealm
ost
everywhere. T
heyhave
nospecial
affinityfor
small, rem
oteislands,but they
were
ableto
reachthem
andcan
surviveon
them,and
theseare
theonly
placesthat w
ehave
left forthem.
Up
toand
duringthe
Pleistocene, great tortoises,
two, three,
four,even
fivefeet
inlength,
havebeen
reportedas
fossilsfrom
allcontinents
exceptA
ustraliaand
Antarctica,
andfrom
many
small to
largeislands
asw
ell (Auffenberg,
1974;H
ooijer,1971;
Jimenez
Fuentes
andC
arbajosaT
amargo,
1982).But
thespread
ofm
ankindover
theface
ofthe
globe,m
orethan
anyother
factor,has
plungedthe
giantsinto
ignominious
retreat. The
opportunistic, ingeniousnew
pred
aton, literally
asom
nivorousas
apig
andm
uchm
oredanger-
ous, provedtoo
much
for tortoisesbeyond
acertain
size. The
greattortoises
tooktoo
longto
reachm
aturity,found
ittoo
difficult tohide, w
eretoo
helplessand
slow. T
heirarm
or—
acrucial
survivalfactor
inpre-hum
antim
es—
was
easilysundered
byprim
itiveim
plements.
One
hugespecim
en,found
asa
sub-fossilin
Florida,
stillenclosed
thespear
thathad
killedit
andretained
thecharring
ofthe
carapacethat
showed
thatit had
beencooked
upside-down
ona
openfine
(Holm
an,1981).
After
theunequal
strugglehad
wiped
thegiants
offthe
faceof
thecontinents,
theylingered
onin
exilein
theirow
nnem
otelittle
Edens, the
archipelagoesofthe
GahIpagos,
theS
eychelles,and
theM
ascarenes,for
thousandsof
years,insular ecosystem
suntouched
bythe
withering
handofm
an,indeed
unknown
tom
an.B
ut finallythese
islandsurvivors
toow
erefound,
andtheir
veryabundance
andtoughness
underabusive
conditions—
“tenaciousof
life”is
howthey
were
always
described—proved
tobe
theirundoing. Whole
islandsw
erestrippedoftheirlasttortoise
sothatthosew
hoselife
styleskept
themat
seafor
months
ata
time
—w
halers,buccaneers,
explorers—
might
bevictualled
with
some-
thingbetter
thansalt pork,
something
that couldcontinue
tolive
form
anym
onths(if
youcall
thatliving),
unfedand
unwatered,
deepin
thew
oodenhull
ofa
sailingship.
The
tortoisesof
theM
ascarenesare
gone—
thelast
onedied
around1804,
avictim
ofthe
same
frenzyof
exploration,colonization,
anddestruction
thatw
ipedout
theinnocent
solitaire,asw
ellas
them
orefam
ousdodo
(North-C
oombes,
1976). The
tortoisesofthe
Seychelles
disappearedalso, w
iththe
exceptionofthe
singlepopulation
oftherem
oteisland
ofA
ldabra.O
nlyin
theG
alapagosdid
some
diversityof
gianttortoise
lifesurvive
untilm
ankindfinally
developeda
con-science, perhaps
inthenickoftime.O
rperhapsitw
astoo
late—
recentnew
sfrom
theG
alapagosis
notgood.
18CH
ELON
IAN
RESEARCH
MO
NO
GRA
PHS,
Num
berI
—I 996
The
Mesozoic
“look”ofthe
Galapagos
faunatoday
isan
illusion.T
hespectacular reptiles
oftheislands
allbelongto
modern
groups.L
argeground
iguanasare
alsofound
inthe
Baham
asand
theG
reaterA
ntilles,and
eventhe
remarkable
spectacleof
blackiguanas
sittingon
oceanbeaches
isa
familiar
one,n
otju
stto
Galapagos
visitors, buttothose
who
havew
alkedthe
blackbeaches
oftheP
acificcoast ofC
entralA
merica.
And
thesnakes
andlava
lizardsof
theG
alapagosw
ouldnot be
considerednotew
orthyin
am
ainlandtropical
setting. The
giant tortoisesare
remarkable
fortheir
size,but
arefam
ousonly
becausew
e—
humanity
—extirpated
themon
thecontinents,
sow
enow
haveto
goto
Galapagos
tosee
them.So
theislands
areno
Lost
World
inthe
Conan
Doyle
sense,but
bycontrast
aresurprisingly
new.
The
oceaniccrust upon
which
thearchipelago
sitsm
aybe
asm
uchas
ten
million
yearsof
age,but
theislands
themselves
probablyem
ergedno
more
than3—
5m
illionyears
ago(S
imkin,
1984).O
fcourse, establishingdefinitive
datesor ages
isim
possiblew
hennew
lavaflow
sm
aycom
pletelyconceal
olderrocks,
but thephysiognom
yofthe
islandssuggests
that thew
esternislands
of
Narborough
andA
lbemarle,
with
theiractive
vulcanism,
lofty,w
ell-definedcalderas,
andextensive
fresh
lavaflow
s, arethe
newest, w
hileH
oodIsland, in
theextrem
e
southeast—low
-lying, probablyuplifted, craterless, eroded
—m
aybe
theoldest. T
hepicture
onegets
isthat o
fashifting
archipelagoplodding
out tosea
onheavy-footed, slow
steps,w
ithnew
erislands
emerging
inthe
west
asthe
easternones
finallydisappear beneath
thew
aves, asthe
Nazca
Plate
uponw
hichall
theislands
sitcreeps
inthe
oppositedirection,
towards
theS
outhA
merican
continent.B
uttheG
alapagosw
erenever partof the
continent,and
thetortoises
must have
reachedthem
bydrifting,
despitethe
assumption
of
Gadow
(1901)that
“accidentaltransport
or
migration
areout ofthe
question,”orthe
musings
ofGarm
an
(1917)
thattheym
ayhave
beenintroduced
bym
an. Perhaps
theycam
efrom
South
Am
erica,and
indeedthe
Hum
boldt
Current
sweeps
convenientlyup
thecoast
ofC
hileand
Peru
beforediverting
westw
ardsat
Equatorial
latitudes.B
utthe
cis-Andean
seaboard,w
ashedby
thecold
Hum
boldt current,
hasprobably
longbeen
anarea
of
remarkable
sterility,and
anotherpossibility
isthat
thetortoises
arrivedfrom
theA
ntillesduring
aperiod
when
theC
entral Am
ericanIsthm
us
was
stilla
seriesof
separateislands.
Certainly,
fossilgiant
tortoiseshave
beenfound
inseveral
ofthe
Antilles,
inclu
d
ingC
uba,M
ona,and
Navassa
(William
s,1950,
1952;A
uffenberg,1974).
But
wherever
theycam
efrom
,they
were
almost
cer
tainlyalready
verylarge.
Sm
alltortoises
frequentlyreach
islandsclose
tocontinental
shorelines,and
therethey
may
reachhigh
densities,although
re-colonizationseem
sto
occurfrequently
enoughfor
suchpopulations
notto
differ-
Figure
1.An
oldm
aleD
uncanIsland
tortoise,O
nan,surveys
hisF
igure
2. Three
adultDuncan
Islandtortoises
ina
galapagueraon
aridterrain.
August
1986.a
clifftopnear
theocean.
March
1970.
PR
ITC
HA
RD
The
Galapagos
Tortoises
19
entiatefrom
thoseofthe
mainland.B
ut islandsbeyond
afew
hundredkilom
etersseem
tobe
reachableonly
bygiants;
andif
“stepping-stone”islands
areabsent,
eventhey
canap
par
entlynot
survivethe
tripto
trulyrem
ote,m
id-oceanislands
orarchipelagoes.
Lines
ofreasoning
tobolster
theargum
entthat
gig
antism
was
apre-adapted
condition,useful
forsuccessful
colonizationof
oceanicislands,
includethe
following:
1) The
newness
oftheislands
inquestion
arguesagainst
therehaving
beentim
efor
anin
situevolution
of gigantism.
2)The
islandofA
ldabra, inthelndianO
cean—
theonly
otherisland
with
surviving, indigenousgianttortoises
—is
thoughtto
havebeen
completely
submerged
severaltim
esduring
thelast few
tensofthousands
ofyears(B
raithwaite
etal.,
1973),yet
eachtim
e,it
was
recolonizedby
gianttortoises.
They
must
havearrived
bydrifting
fromother
IndianO
ceanislands;
but notethatit
cannothappen
again,because
tortoisesare
nowextinct
onall
thepotential
“do-nor”
islands.3) In
theocean
environment,a
giant tortoisew
ouldhave
many
survivaladvantages
overa
small
one.It
would
havem
oreextensive
fatreserves;
itcould
holdits
headhigher
abovethe
waves,
thusavoiding
accidentalingestion
ofsea
water; it w
ouldhave
am
orefavorable
surface/volume
ratio,thus
reducingosm
oticw
aterloss;
and, once(D
eovolente)
alive
landfallwere
made,a
giantwould
bebetterequipped
forlocom
otionin
brokenor
boulder-strewn
littoralterrain,
forsurvival
underuncertain
orextrem
etherm
alconditions,
ortoleration
ofprotracted
dessicatingconditions.
4)T
hevariation
observedin
thesurviving
populationsof
Galapagos
tortoises(Figs.
1—5)
suggeststhat
thepost-
arrival specializationsthat have
occurred—
specifically, thedevelopm
ent ofsaddlebacked, flared
shell forms,
generallyon
smaller, m
orearid
islands—
havebeen
characterizedby
areduction
inadult
size(partly
genetic,partly
aresult
oferratic
availabilityof
foodand
water).
Far
from“islands
creatinggiants,”
thereal
trendm
ayhave
beentow
ardssecondary
dwarfism
,although
itis
possiblethat
thetrend
was
complicated
orblurred
bysexual
selectionpushing
in
theopposite
direction, thesuccessful m
alesgenerally
beingthe
biggestanim
als.W
hyare
thegiant tortoises
sovulnerable,
sosubjectto
utterdisappearance
within
decades,or
at most
acentury
ortw
o,of
discovery?T
heydon’t
seemdelicate,
andeven
inrelatively
mediocre
zoosw
ithout evena
closefacsim
ileofan
appropriateclim
ate,w
heretortoises
ofother
speciesrarely
survivefor
long,G
alapagostortoises
may
live,w
ithoutproblem
s,for
many
decades—
althoughbreeding
themis
anotherstory.
The
answeris
complex. F
irstly, massiveness
andph
ysi
calim
pregnabilityevolve
underconditions
ofprotracted
stability, andare
not thekeys
tosurvival ordom
inancein
thefast-changing
modern
world.
Today
thefellow
travelerson
Planet
Earth
thatthreaten
todom
inateus
areviruses,
notV
elociraptor;tsetse
flies,not
Triceratops.
Moreover,
while
lifem
aybe
hardon
islandsthat
arefrequently
without
water,
andsom
etimes
scorchedby
vul
canismas
well
asthe
tropicalsun,
theabsence
oflarge
predatorshas
made
thetortoises
“soft”in
certainw
ays,to
usean
unlikelym
etaphor.O
nthe
continents,tortoises
havebeen
forcedto
surviveby
concealment
—hiding
inrocky
crevices,burrow
s,or
densevegetation;
bycrypsis,
with
camouflaging
shell markings
ordesigns;byelaborate
mod
ifications
ofthearm
or, with
anterior extensionsto
protecttheextrem
ities,orby
hingingof
varioussections
oftheshell,to
allowcrucial
lifefunctions
likem
atingand
ovipositionw
ithoutleaving
theanim
alopen
toattack
throughgaping
shellopenings;
byingrained,
appropriate,cautious
beh
avioral patterns
thatreducethe
exposureto
discovery;and
byequilibrium
populationdensities
thatareoften
quitem
odest.In
many
continental areasw
heretortoises
arequite
success-ful,
theynevertheless
areso
inconspicuousthat
theyare
hardlyever
seen.O
nthe
islands,these
constraintsdisappear.
The
tor
toises,instead
of beingelusive
masters
ofsurvival,
become
dominant.
They
achievehigh
densities,can
fallasleep
orfeed
wherever
theyw
ishw
ithoutany
obligationto
keepout
ofsight,
andcan
modify
theirrigid
carapacialenvelope
Fig
ure
3. The
snow-w
hiteanteriorhead
colorationofan
adult male
Hood
Islandtortoise.
September
1982.F
igu
re4.
Spuriouslyferocious
appearanceof
atortoise
fromIndefatigable
Island.Septem
ber1986.
20C
HELO
NIA
NR
ESEAR
CH
MO
NO
GR
APH
S,N
umber
1—
1996
(towards
saddlebackedform
s, or reducedthickness)
inw
ays
thatm
ayfacilitate
oviposition,therm
oregulation,or
locom
otionin
brokenterrain,
butthat
certainlydo
notenhance
securityfrom
predators.T
helatter,
afterall,
were
absentuntil
thedogs
andpigs
arrived.So
what
worked
well
undervirgin
conditionsbecom
esa
liability
underviolated
ones.T
hevulnerability
ofhatchling
Galapagos
tortoisesto
ratsis
curious.A
fterall,rats
areubiquitous
underco
ntin
en
talregim
esw
heretortoises
havealw
aysbeen
exposedto
them. M
oreover, hatchlingand
smalijuvenile
tortoisesofall
species
arevirtu
allydefen
seless,w
ithscant
bonein
their“shells,”
andoffering
anutritious,
lipid-richpackage
thatcan
neith
ero
utru
na
pred
ator
norev
entu
rnan
dnip
it onthe
snout.The
explanationprobably
liesin
thefact
thatthe
eco
logical
scenario
that
allow
sso
me
hatch
ling
torto
isesto
surviveis
always
adelicate
one,and
thepredators
them-
selvesm
ust
beunder
vario
us
ecolo
gical
constrain
tsif
theyare
notto
consume
allof
thehatchlings.
Thus,
variousrelativ
elysu
btle
land-u
sepattern
sin
theC
aliforn
iadesert
areas(esp
eciallythe
Mojav
e)h
ave
caused
un
natu
ralp
rolif
erationof
ravens,a
nativespecies
thatoften
preysupon
hatch
ling
andju
ven
ileto
rtoises.
Tod
ay,
raven
sco
nsu
me
so
man
yy
ou
ng
desert
torto
isesin
som
eim
pacted
areasth
atthe
capacity
of
theto
rtoise
po
pu
lation
tosu
rviv
eis
inqu
estion.
Sim
ilarlyin
theG
alapag
os,
intro
du
cedblack
ratsare
presenton
many
ofthe
islandsw
herethe
tortoiseslive,
butin
most
casesthe
ratsthem
selveshave
predators,including
feralcatsand
dogs,asw
ellasnative
owls
oftwo
species, andG
alapagoshaw
ks.O
nsuch
islands,som
enatural
recruit-m
entof
tortoisesoccurs.
On
theother
hand,on
Duncan
Island,the
onlyintroduced
mam
mal
isthe
blackrat,
whose
populationsprobably
cyclem
assivelyunder
constraintsof
climate
andfood
availability, but thatcan
reachhigh
po
pu
lationsquickly
when
conditionsare
good(H
oeck,1984).
The
rats’only
predatoris
therelatively
scarceG
alapagoshaw
k.Under these
conditions, it appearsthat naturalrecruit-
men
tof
theD
uncan
torto
isecan
almost n
ever
occu
r,an
dthe
population, despiterem
arkablelongevityofthe
adults, would
hav
eb
eend
oom
edw
itho
ut
thein
tensiv
eeg
g-rescu
ean
d
head
-starting
pro
gram
un
dertak
enby
theG
alapag
os
Na-
tionalP
arkand
theC
harlesD
arwin
Research
Station.
This
effort h
assp
elledn
ewh
ope
for
ap
op
ulatio
nofto
rtoises
that
seemed
tohave
fl() future; perhapsitalso
signalsthatthere
isev
enh
op
efor
man
kin
d.
Figure
5.C
opulatingtortoises
inthe
lushcaldera
ofV
olcanA
lcedo,A
lbemarle
Island.M
arch1975.
HIS
TO
RIC
AL
OV
ER
VIE
W
The
voluminous
literatureon
Galapagos
tortoises,re
centlyinventoried
byB
eaman
(1985),can
bedivided
intofour
chronologicallydefined
phases.T
hesephases
are:D
iscovery(1535—
1840),T
axonomy
(1824—1917),
Rescue
( 1925—1955),and
InSitu
(1957—present).T
hephases
reflectthe
historicaldevelopm
entof
human
attentionto
thegiant
tortoisesof
theG
alapagosIslands.
Tortoises
areknow
nor
suspectedto
haveoccurred
onI1
ofthe13
major
islandsin
thearchipelago
(seeM
ap1
onnext
page).
Discovery
Phase
The
firstphase,
andthe
longest,m
aybe
termed
the“D
iscovery”phase
(1535to
1840).F
romthe
discoveryof
thearchipelago
byB
ishopT
omás
deB
erlangain
1535to
theepochalvisitofC
harlesD
arwin
exactlythree
centurieslater,
thepresence
ofgigantic
tortoisesin
theislands
was
do
cuinented
bym
any,includingD
ampier( 1697),C
olnett(1798),M
itchill(1815),
Delano
(1817),F
itzR
oy(1839),
andD
arw
in(1
839).B
erlanga’sow
nobservations
onthe
islands,including
thetortoises,
were
transmitted
ina
letterto
theH
olyR
oman
Em
peror,C
arlosV
,in
aletter
dated26
April
I535, andpublished
inE
nglishtranslation
byS
levin(1959).
Many
ofthese
authorsduring
the“D
iscovery”phase
gaveinsight
intothe
naturalhistory,
abundance,distrib
ution,
andeconom
icor
subsistencevalue
ofthe
giantto
rtoises,and
gaveatleastanecdotalinform
ationon
theirinter-
islanddifferences.
How
ever,itwas
thegeneralpractice
at thetim
eto
refertoallisland
gianttortoises,worldw
ide,asa
singlespecies,called
Tortue
desindes
bythe
pre-Linnaean
authorityPerrault
(1676),and
Testudo
indicaby
Schneider
(1783).
Taxonom
icP
hase
Overlapping
onlyslightly
with
the“D
iscovery”phase
WL
Sa
secondperiod
(1824to
1917)thatm
aybe
termed
the“T
axonomic”
phase.T
hisperiod
startedw
iththe
firstpro-
posalsofscientific
names
forGalapagos
tortoises—
Testudo
calijbrnianaand
7’.nig
ra,by
Quoy
andG
aimard
(1824a,I824b),
andT.
elephantopusby
Harlan
(1827).D
uringthis
phase,E
uropeanand
Am
ericanzoologists,
usuallyasso
ciated
with
major
museum
s,described
andgave
names
tothe
specimens
thatarrivedin
theirinstitutions.T
hesew
ereoften
fromunknow
norvague
sourcesthe
provenanceofnone
ofthe
typespecim
ensofthe
above-named
taxacan
betraced
toa
particularisland,
norcan
itbededuced
fromthe
morp
hol
ogyof
thespecim
ens,since
allw
erejuveniles.
Indeed,the
juvenilethatservedas holotype
forbothT
estudocalijbrniana
and1’.n
igra
(MN
HN
P9550, carapace
length26.8
cm,Figs.
6-8)w
asreported
tohave
come
fromC
alifornia,and
tohave
beendonated,
asa
livinganim
al,to
M.
deF
reycinetby
M.
Meek,captain
oftheA
merican
shipB
ostonE
agle(l’A
iglede
Boston).T
hedonation
was
made
inH
awaii
(IlesS
andwich).
The
allegedC
aliforniaorigin
ofthespecim
enm
ayhave
beensupposition
basedupon
thesuperficialsim
iliarityo
fayoung
Galapagos
tortoiseto
anadult
ofthe
California
deserttortoise,G
opherus(X
erobates)agassizi, although
itmustbe
admitted
thatthere
isno
evidencethat
Quoy
andG
aimard
were
familiar
with
thisspecies.
Nevertheless,w
herethe
typem
aterialstillexists,q
ues
tionsas
tothe
preciseorigin
ofthese
specimens
may
notrem
ainforever
unanswered
asbiochem
icaland
genetictechniques
andD
NA
andprotein
analysesbecom
eever
more
sophisticated.T
henum
erousw
ritingsof
Gunther
(1875,1877,
1896,1898,
1902)and
Rothschild
(1896,1901,
1902a,1902b,
l902c,1902d,
1903,1915a,
l9lS
b,
1925,1928)
contributedm
uchduring
thisphase,
althoughneither
couldresist
theurge
tonam
enew
speciesbased
oninadequate
material
—
oftensingle
specimens
ofunknow
norigin.
GU
nther’s1877
monograph
onthe
gianttortoises,beautifullyillustrated
andoffering
detaileddescriptions
oftheosteology
ofmany
speci
mens,
stillsfalls
shortofprovidingreal
insightintothe
distri
butionand
relationshipsofthe
differentG
alapagosform
s.T
histask
fellto
JohnV
anD
enburgh(l914)
who
pub-lished
byfar
them
ostcom
prehensiveaccount
ofthe
Galapagos
tortoisesever
written.
Copiously
illustratedw
ithphotographs
ofspecim
ensfrom
standardizedangles
andbased
uponnew
ly-collectedm
aterialtotalling
256sp
ecim
ensof
preciselyknow
norigin
(collectedon
theA
cademy
expeditionof
1905—06,
largelyby
Rollo
Beck
andJoseph
Slevin),
itstands
asa
work
ofenorm
ousvalue
eventoday.
Moreover,
sincethe
currentprohibition
oncollecting
livespecim
ensof
Galapagos
tortoisesis
likelyto
continue,and
sincesom
eof
theisland
populationsare
extremely
rareor
extinct,thereis
nopossibility
ofduplicatingV
anD
enburgh’sstudy
series,housed
todayin
theC
aliforniaA
cademy
ofS
ciencesin
SanF
rancisco.Y
etboth
Van
Denburgh
andG
unther—
andalso
thetalented
Georg
Baur
—fell
intothe
trapof
attempting
todeterm
inethe
provenanceof certain
specimens
ofunknown
originby
aprocess
oflogical
argument.
Specim
ensin
hand(often
singleskulls
orshells)
were
compared
with
isolatedspecim
ensof
known
origin,accounts
byancient
mariners
were
consulted,and
rumors
asto
when
certainisland
forms
hadbecom
eextinct
were
discussed—
theintellectual
pot
was
stirredand
fermented
and,w
ithan
almostvisible
flour-
ish,m
agisterdixit,
andthe
originof
thespecim
enw
as
declared.S
uchguesses
were
presentedas
gospel,or
at least
assufficientbasis
formaking
nomenclaturaldecisions
often
involvingthe
naming
ofnewtaxa,even
thoughit is
clearfrom
Van
Denburgh’5
numerous
photographs—
andalso
fromthe
inabilityofm
odernzoo
curatorstoidentifythetortoises
intheir
charge—
thatthevariation
isenough
tom
akesuch
guesswork
exceedinglyunw
ise.Yetw
henhe
hadadequate
material,V
an
Denburgh
was
capableof greatand
cautionaryw
isdom,
asin
hiscom
ments
aboutG
alapagostortoise
skulls:“In
aseries
of 24skulls
fromV
ilamil,A
lbemarle,Ifind
allof
thevariation
which
GU
ntherm
entions;and
uponcareful
comparison
ofthis
seriesw
ithone
skullfrom
Hood
Island(N
o.8
125),one
fromIndefatigable
(No.
8381),
onefrom
James
(No.
8105),three
fromD
uncan(N
os.8378,
8379,and
8380),four
fromC
hatham(N
os.8127,
8128,8130,
and8131),
andone
fromIguana
Cove
(No.
8179),I
canfind
noconstant
differencesin
theskulls
ofthe
various
races.In
theskulls
fromV
ilamil,
thefrontal
regionm
aybe
flat orsomew
hatconvex.The
occipitalspinem
aybe
shortor
long,not
reachingthe
posteriorborders
ofthe
mastoid
processesor projecting
farbehindthem
,andm
ayorm
aynot
risem
uchabove
thelevel
ofthe
skull.T
hereis
much
variationin
theshape
ofthe
tympanic
caseand
cavity.T
he
fossain
frontof
theoccipital
condylem
aybe
deepor
very
shallow.
The
tuberosityfor
thetem
poralm
usclem
aybe
quitesm
allor
verylargely
developed.T
henasal
opening
may
beas
highas
broad,or
broaderthan
high.T
hepalatal
regionvaries
much
inshape.It m
aybe
narrowor broad,
andthe
pterygoidedges
may
besharp
orblunt.
The
alveolar
ridgesalso
varyin
positionand
degreeof
development.W
e
may
safelysay
thatnoconstantdifferences
existam
ongthe
skullsof
thevarious
racesof
Galapagoan
tortoises.”
Although
Rothschild
publishedshort
papersin
1925
and1928,S
amuelG
arman’s
1917revision
oftheG
alapagos
tortoises,based
uponthe
collectionsof
L.
Agassiz
andG
.B
aurat
theM
useumof
Com
parativeZ
oologyat
Harvard
22C
HELO
NIA
NR
ESEAR
CH
MO
NO
GR
APH
S,N
umber
1—
1996•1
IND
LO
E)
N
;i::rov
EsA
$‘;:
ISL
AS
AN
SA
LV
AD
OR
-
BA
LT
RA
(IND
EF
AG
AL
E)
BA
RR
ING
TO
N
ISA
NT
AM
AR
IA4;’S
AE
SP
AO
LA
jb1L
vlL
AS
AN
CR
ST
oA
L
Map
1.Galapagos
Islands.Tortoises
areknow
nor suspected
tohave
occurredon
11separate
islands:Narborough
(Fernandina)(probablyextinct),
Albem
arle(Isabela)
(fiveseparate
populations,distributed
onthe
separatevolcanoes),
Abingdon
(Pinta)
(probablyextinctin
the
wild,
oneindividual
male
alivein
captivity),Jam
es(San
Salvador),Jervis
(Rábida)
(extinct,possibly
introduced),D
uncan(Pinzón),
Indefatigable(SantaC
ruz),Barrington
(SantaF
é)(extinct,possibly
introduced),Charles
(SantaM
arIaorF
loreana)(extinct),Chatham
(San
Cristóbal)
(two
separatepopulations,
oneextinct),
andH
ood(E
spañola).B
lackdots
representextant
populations,lightly
shadeddots
representextinct
orextirpated
populations.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises23
University,
essentiallyrepresents
thefinal
flourishof
the“T
axonomic”
phaseof
Galapagos
tortoisedocum
entation.T
othis
day,this
idiosyncraticw
orkm
ustbe
consideredthe
lastcom
prehensiveattem
ptto
establisha
nomenclature
forG
alapagostortoises.T
hatsuchan
uneveneffortshould
havew
ithstoodform
alchallengeforthree
quartersofa
centuryis
littleshort
ofincredible.
Yet,
inthe
fieldof
gianttortoise
biology,the
cyclesof
assertation,argum
ent,and
counter-argum
entm
aybe
asslow
asthe
tortoisesthem
selves.W
it-ness,
forexam
ple,the
fieldaccount
ofthe
IndianO
ceanR
odriguestortoises
(Cylindraspis)offeredby
Leguat(1708),
following
hissojourn
onR
odriguesduring
theyears
1691—
94.It took
overtw
ocenturies
forA
tkinson(1922)
toq
ues
tionL
eguat’sw
orkby
thedevastating
techniqueof
arguingthat L
eguathad
neverexisted;
thebook,
Atkinson
claimed,
was
afictitious
accountby
anarm
chairtravelogue
writer.
More
thanhalf
acentury
more
was
topass
beforeN
orth-C
oombes
(1979)defended
bothL
eguat’sexistence
andhis
observationsin
am
eticulouslyresearched
andextraordinar
ilythorough
thesis.
Rescu
eP
hase
Beginning
approximately
with
Beebe’s celebrated
1925book
Galdpagos
—W
orld’sE
nd.and
continuingw
iththe
Figure
6. Originaiplate
oftheholotype
ofTestudo
nigra(M
NH
NP
9550)from
Quoy
andG
aimard,
1824h.Photo
courtesyof
Bihliothèque
duM
useumde
Paris.
writings
ofCharles
Tow
nsend(1924a,1924b,
1925a,1925b,
1928a,1928b,
1931a,193lb,
1932.1942),
Ralph
De
Sola
(1929,1930),
andothers,
athird
or“R
escue”phase
ofG
alapagostortoise
writings
ensued.W
iththe
assumption
thatthetortoises
were
sorare
andso
stressedin
theirnative
habitat(by
directtake
bysettlers,
andby
competition
orpredation
fromferal
mam
mals)
thattheir
survivalw
asun
likely,itwas
consideredim
perativeto
transferthemto
zoos,w
herethey
would
beencouraged
toreproduce.
The
effortsstarted
shakily,w
ithB
eebe’scollection
ofa
singleD
uncanIsland
tortoisethatdied
aftera
week,butT
ownsend
in1928
was
ableto
collecta
largeseries
of180
mostly
juvenilespecim
ens,largely
fromsouthern
Albem
arleIsland.
Som
eof
theselater
reproducedin
zoos,and
afew
surviveto
thisday
(Fig.9).
The
captivegroups
were
alsothe
subjectofbehavioralstudies
byE
vans(1949),E
vansand
Quaranta
(1949,
1951),and
others.D
uringthis
phase,only
De
Sola
(1930)gave
anew
name
(Testudo
vandenburghi)to
thelive
tortoiseshe
collected,andsince
thisvery
briefpaperdidnotidentify
typespecim
ensnor
featuresby
which
thenew
speciescould
bedistinguished
fromothers,
confusionrather
thanclarifica
tionw
asthe
result.T
ypicalof
thew
ritingsof
thisperiod
was
theo
bserv
ation
ofB
anning(1933),
who,
injustifying
thetaking
ofa
Figures
7and
8. Ventraland
lateralviewofholotype
ofTestudo
izigm(M
NH
NP
9550)as
itexists
today.Photos
courtesyJean
Lescure,
Museum
National
d’Histoire
Naturelle.
24CHELONTAN
RESEARCH
MO
NO
GRA
PHS,
Num
berI
—1996
seriesof
18G
alapagostortoises
(12actually
obtainedfrom
Guayaquil
onthe
mainland
ratherthan
fromthe
islands
them
selves)
toS
anD
iego
bythe
Hancock
expeditionof
1933,com
mented:
“Ifthe
‘galapago’the
prodigious
gianttortoise
fromw
hichthe
islandstook
theirnam
e—
isto
surv
ive,
itis
scarcelyprobable
thathis
nativesoil
will
providethe
environment.”
The
“Rescue”
phasefinally
achievedfulfillm
entw
ith
theannouncem
entofsuccessulcaptivebreeding
ofGalapagos
tortoisesin
thezoos
ofS
anD
iego(S
taedeli,1958;
Shaw
,
1959,1961a,
1961b)and
Honolulu
(Throp,
1969,1975).
The
realizationfollow
ed,how
ever,that
captivebreeding
would
bestbe
undertaken—
where
necessary—
inthe
islandsthem
selves,utilizing
breedingstock
orknow
nand
homogeneous
islandm
aterial.The
“Rescue”
phasereached
fullcycle
when
two
representativesof
therarest
island
forms,
aD
uncantortoise
inthe
Bronx
Zoo
(Fig.10)
anda
Hood
Islandtortoise
inthe
SanD
iegoZ
oo(Fig.
I 1),w
ere
returnedto
theislands
(Bacon,
1978;P
ritchard,1979).
Significantly,
inboth
casesthese
individualsw
ereof
wild-
caughtstock
ratherthan
captive-bred.N
evertheless,som
e
individualsofthe
original1928
collectionstillsurvive
inU
S
institutions,and
inrecent
yearsthere
hasbeen
excellent
captivereproduction,
e.g.,at
theG
ladysP
orterZ
ooin
Brow
nsville,T
exas,and
atthe
Life
Fellow
shipF
acilityin
Seffner,
Florida
(where
85hatchlings
were
producedin
1988—89, w
ithsom
efem
alesnesting
fivetim
esin
aseason).
As
of1994,
Life
Fellow
shiphad
hatchedand
keptalive
a
totalof
411
Galapagos
tortoises.
InSitu
Phase
During
World
War
IIand
fora
decadethereafter,
opportunitiesto
visitthe
Galapagos
were
few(except
for
thoseon
Baltra
Islandon
US
military
service),and
nofield
studiesw
ereundertaken.
How
ever,starting
around1957
with
theU
NE
SC
O/L
ifeM
agazineG
alapagosE
xpedition
underthe
scientificleadership
ofI.
Eibl-E
ibesfeldtand
Robert
Bow
man,
anew
phasew
asinitiated,
continuingto
thepresent,
thatm
aybe
calledthe
“inSitu”
phase.E
vents
nowunfolded
rapidly,leadingnotonly
tothe
constructionof
theC
harlesD
arwin
Research
Station
andthe
establishment
oftheG
alapagosN
ationalP
arkof the
Republic
ofEcuador,
butalso
toa
vastand
increasingvolum
eof
literaturebased
uponfield
studiesof
theG
alapagostortoises.
Som
econ-
spicuouscontributions
duringthis
phasew
erethose
of E
ibl
Eibesfelt
(1959),S
now(1964),
MacF
arlandet
al.(1974a,
1974b),M
acFarland
andR
eeder(1974),
Schafer
(1982,
1983),F
owler
(1983),and
Cayot
(1987),although
these
titlesrepresentonly
asm
allfractiono
ftho
segenerated
from
I958to
date.D
etailsofthe
“inSitu”
phaseconservation
activitiesare
givenin
theS
urvivalS
tatussection
laterin
thisw
ork.T
he
essenceof
thestrategy
devisedby
theexperts
at theC
harles
Figure
9.Scenein
Houston
Zoo,T
exas,1970. Z
oogoersadm
irea
largeadultm
aletortoise
fromA
lbemarle
Island(Isabela), “rescued”
asa
juvenilefrom
theIguana
Cove
(Cerro
Azul)
areain
1928by
Charles
Tow
nsend.
PR
ITC
HA
RD
Th
eG
alapag
os
Torto
ises25
.
.)
Fig
ure
10.Adult fem
aleD
uncanIsland
(Pinzón) tortoise, captured
byC
harlesT
ownsend
for theB
ronxZ
ooin
1928, andreturned
toD
uncanin
1972.T
hew
hitehead
istypical
bothof
oldG
alapagostortoises
andof
representativesof
thesaddlebacked
races.
Fig
ure
11.A
dultm
aleH
oodIsland
(Española)
tortoise,resident
inthe
SanD
iegoZ
oofor
over40
yearsuntil
itsreturn
tothe
Galapagos
in1977,
where
itis
nowa
contributorto
thesuccessful
captivebreeding
programfor
thisendangered
population.M
ayI970.
26CH
ELON
IAN
RESEARCH
MO
NO
GRA
PHS,
Num
ber1
—1996
Darw
inR
esearchS
tationand
theG
alapagosN
ationalP
ark
was
arecognition
thatthe
tortoisesw
erem
enacedby
a
mosaic
ofstress
factors,the
precisecom
binationof
which
was
uniqueto
eachpopulation. T
hus, aninventory
ofdiffer-ent
techniquesw
ouldbe
required,w
iththe
leastm
anipula
tive(and
the‘east
expensive)being
appliedw
heresuch
techn
iqu
esw
ou
ldbe
adeq
uate,
orw
here
po
pu
lation
sizes
were
sufficient toabsorb
some
losses, but with
progressively
more
ambitious
techniquesapplied
where
necessary.T
hestress
factorsidentified
includedcapture
ofto
r
toisesby
smugglers
forsale
overseas,slaughter
of
tortoises
bysettlers, food
competition
byferal
herbivores(especially
goats,but
alsocattle
andburros),
predationon
primarily
subadulttortoises
bydogs,
predationon
juvenilesand
eggsby
pigs, andpredation
onhatchlings
byintroduced
rats, orby
nativespecies
includingthe
Galapagos
hawk.
Furtherm
ore,som
epopulations
were
sodepleted
thateven
sporadic,
accidentalm
ortalityof
adultscould
notbe
tolerated,or
reproductionhad
effectivelyceased
becauseencounters
between
males
andfem
alesno
longeroccurred.
For
some
populations,the
solutionw
asa
combination
oflaw
enforcement,
education,and
controlor,
where
po
s
sible, elimination
offeral
mam
mals. H
owever, in
thecase
of
thetortoises
of
Duncan
Island,w
hereferal
ratsconsum
ed
everyhatchling, it becam
enecessary
tocollect eggs
fromthe
wild, incubate
themartificially, and
head-start theoffspring
for several yearsbefore
releaseon
their ancestral island(Fig.
12).T
ransferof
eggsto
artificalincubation
andhead-
startingof
theresulting
youngw
asalso
practicedfor
the
tortoisesof
James
andC
hath
amislan
ds,
where
theeggs
andyoung
were
threatenedby
feralpigs
anddogs,
resp
ectively
.O
nly
inthe
caseo
fthe
extrem
elyd
epleted
torto
isesof
Ho
od
Island
was
them
ostlab
or-in
tensiv
e
techniqueof
all—
thecapture
andconveyance
ofthe
entire
wild
populatio
nto
acap
tive-b
reedin
gfacility
—
foundto
benecessary.
The
ultimate
challengew
aspresented
bythe
discoveryof
asingle
survivingm
aletortoise
onA
bingdonIsland
in
I972.T
heanim
alhas
residedin
captivityat
theC
harlesD
arwin
Research
Station
forthe
ensuing24
years,but
attempts
tom
atehim
with
females
fromV
olcanW
olf
(Albem
arleIsland)
havefailed, and
ifanyofhis
genesare
tosurvive,
technologicalrem
ediesw
illhave
tobe
soughtand
applied.
Fig
ure
12. Juvenilecaptive-hatched
Duncan
Islandtortoises
beinghead-started
at theC
harlesD
arwin
Research
Station.A
ugust1986.
NO
ME
NC
LA
TU
RA
LS
TA
TU
S
Inthe
sectionsthat follow
variousaspects
ofthenom
enclature
ofG
alapagostortoises
arediscussed.
Many
pro
blem
sexist,
fromthe
actualscientific
names
usedto
thevalidity
anddistinctness
ofthevarious
populationsorform
s.T
hecurrently
comm
onlyused
speciesnam
eforallG
alapagostortoises,
Geochelone
elephantopus,w
hoseusage
(with
separatesubspecies
forthe
variousisland
forms)
datesonly
backto
Mertens
andW
ermuth
(1955),is
notthe
oldestavailable
name
forthe
speciesand
thereforeneeds
tobe
corrected.Many
populationsbearinaccurate
orerroneouslyallocated
names,
severalnam
edpopulations
needto
besynonym
ized,and
some
distinctform
sneed
tobe
named.
The
firstsection
belowreview
sand
critiquesthe
lastmajor
taxonomic
work
regardingG
alapagostortoises
(Garm
an,19
17).T
hecurrent
statusof
thenom
enclatureis
thenan
alyzed
inthe
secondsection,
followed
bya
summ
aryof
recomm
endations.
Garm
an’sT
heG
alapagosT
ortoises
Sam
uelGarm
an[1843—
1927]w
as73
yearsold
when
hepublished
The
Galapagos
Tortoises
in1917
—his
onlypublication
ontortoises.
Afam
ousand
distinguishedsenior
herpetologistandheir
tothe
Agassiz
mantle
at theM
useumof
Com
parativeZ
oology(M
CZ
)at
Harvard
University,
Garm
anhad
made
notablecontributions
tothe
studyof
seaturtles,
havingdescribed
two
newspecies
(Lepidochelys
kempi
andC
heloniadepressa)
in1880.
These
remain
theonly
validspecies
ofsea
turtleto
havebeen
describedsince
Bocourt’s
descriptionof
Chelonia
agassiziin
1868.Y
etG
arman
was
ahighly
eccentricand
lessthan
meticulous
worker.H
isodd
andsecretive
working
habitsare
describedby
Adler(l989).G
arman’stype
seriesofC
heloniadepressa
includeda
perfectlytypicaljuvenile
greenturtle,
Chelonia
mydas
(Barbour,
1914),w
hilein
hisdescription
ofL
epidochelyskenipihe
repeatedpurely
hearsayinform
ationto
theeffect thatthis
speciesnested
inthe
Florida
Keys
fromD
ecember
toJanuary
(infact,
itis
virtuallyconfined
asa
nestingspecies
toT
amaulipas,
Mexico,
anditonly
nestsin
thespring
months).
Garm
analso,in
describingthe
newand
validm
udturtle
speciesK
inosternonbaurii
in1891,
reported
thisspecies
notonly
fromFlorida,
where
itindeed
occurs,although
nolonger
at thetype
localityofK
eyW
est(L
azell,I989),
butalso
fromC
uba(w
hereno
mud
turtlesoccur).
Garm
an’S
1917
work
onthe
Galapagos
tortoises,pub-
lishedabout
aquarter
centuryafter
theclose
oftheauthor’s
primary
herpetologicaloutput,
isalso
somew
hatpuzzling,
inconsistent,and
superficial.Although
chronologicallythe
lastm
ajorpaper
inthe
“Taxonom
y”phase
ofG
alapagostortoise
documentation,
itis
certainlynot
thelast
word.
Perhapsit
was
more
comm
onin
1917than
todayfor
thecontributions
ofdistinguishedelder
statesmen
ofscienceto
receiveonly
superficialcritical
comm
entby
presumably
youngerand
certainlydeferential
technicalreview
ers,especially
when
theyw
eresubm
ittedto
journalsover
which
theauthors
hadadm
inistrativeauthority.
Cer
tainlythis
seems
tohave
beenw
hathappened
with
Garm
an’sw
ork.G
arman’
5statedstim
ulusfor w
ritinghis
revieww
asthe
largecollection
ofG
alapagostortoise
material
—m
ainlyshells
thathadaccum
ulatedatthe
Museum
ofCom
parative
Zoology.
But
italso
hearkenedback
toa
much
earlierphase
ofGarm
an’slife—
to1872,w
henhe
was
inW
yoming
collectingfossils
forE
dward
Drinker
Cope.
Cope
firedG
arman
form
akingunreasonable
salarydem
ands,and
Garm
andrifted
westw
ard, arrivingin
SanF
ranciscoin
time
tosee
Louis
Agassiz
andhis
partyarrive
fromthe
Galapagos
onthe
shipH
assler.Agassiz
recognizedG
arman’s potential
onthe
spot,andoffered
hima
placeatH
arvardC
ollegeashis
pupil.T
hus,the
tortoisespecim
ensand
Garm
anhim
selfarrived
atHarvard
ataround
thesam
etim
e.Perhaps
Garm
an‘s
work
was
alsom
otivatedby
alittle
transatlanticrivalry
Rothschild
(1915a,l915b)
hadjust
publisheda
catalogueand
accountofthelarge
collectionof
gianttortoisesinhispersonalm
useumat T
ring.YetG
arman’s
materialin
many
caseshad
dubiousorabsentcollecting
data,and
evenw
henthe
islandof
originw
asrecorded
—as
with
thejuvenilesfrom
Charles
Island(=Santa
Maria
orFloreana)obtained
byA
gassizon
theH
asslerexpedition
in187
1—72
—
thetortoises
were
purchasedfrom
localsw
hohad
pre
sumably
broughtthem
tothis
islandfrom
elsewhere,
thelocal
tortoiseshaving
become
extinctvery
shortlyafter
Darw
in’svisit
in1835
(Seemann,
1846,in
Van
Denburgh,
1914).H
owor
why
Garm
anconcluded
(p.266)
thatthe
[-fasslerspecimens,too
smallto
showany
distinctiveisland
characteristics,w
erereferable
tofour
differentspecies,
iscom
pletelyunclear.
Had
Garm
an’spaperappeared
beforeV
anD
enburgh’s1914
monograph,
thepurpose
—ofdoing
thebestpossible
with
availablem
aterial—
would
havebeen
evident.A
sa
subsequentpaper,however,itseem
sto
takescientific
under-standing
ofthe
Galapagos
tortoisesa
stepbackw
ard.G
arman’s
material
mighthavejustified
thepublication
ofa
28CHELONIAN
RESEARCHM
ONOGRAPHS,A
/umber
1—
1996
catalogue,but
them
aterialin
theM
CZ
collectiondid
notw
arrantusingitas
virtuallythe
solebasis
fora
reviewofthe
group.V
anD
enburgh(1914)
hadrecognized
fourteensp
ecies
ofG
alapagostortoise,
severalof
them(T
estudohoodensis, T.chatham
ensis,T.darw
ini,and7’.phantastica)
describedby
himself
afew
yearsearlier
(Van
Denburgh,
1907).G
arman
assigned1’. darw
ini,T.
wallacei
(which
heincorrectly
attributedto
Van
Denburgh
ratherthan
toR
othschild),and
theunnam
edform
thatV
anD
enburghreported
fromC
owley
Mountain
(=V
olcanA
lcedo)to
thesynonym
yofT
estudonigrita
Dum
érilandB
ibron,1835
. He
synonymized
T.hoodensis,
T.becki,
andT.phantastica,
asw
ellas
theearlier
species1’.
ephippiumand
T.abingdoni,
with
Testudo
elephantopusH
arlan,1827.
The
latternam
ew
asused
fora
mixed
assemblage
ofsaddlebacked
forms,
eventhough
itsholotype
was
ajuvenile
ofunknown
islandorigin,andirretrievablylost.(A
.E.B
rown,in
Van
Denburgh,
1914,had
locatedthe
cleanedleg
bonesof
oneside
ofa
tortoise,and
apart
ofthe
legsof
theother
sidestillbearing
some
driedskin,
inthe
collectionofthe
Philadelphia
Acad
emy
ofS
ciences;these
were
probablyall
thatrem
ainedof
thetype
ofT.
elephantopus).F
urthermore,
Garm
an(p.
269)did
nothelpthose
seek-ing
hisactual
views
bylisting
T.abingdoni,T.becki,
andT.
duncanensis,all
expressedin
binomial
form,
as“varieties”
ofT.elephantopus,whilelater(p.294)
helistsT.ephippium
,T.
abingdoni,T.
becki,T.
hoodensis,and
T.phantasticus
[sicjas
“so-calledspecies.”
The
name
T.duncanensis
was
quotedonly
once,inpassirn,
andis
anom
ennudum
.We
cansuspect
fromG
arman’s
quotationofB
aur’s(1889)
insight-fulargum
ent(pp.295—296)thatthe
holotypeofT
.ephippiumw
asnot
infact
fromD
uncanIsland
butw
asinstead
anA
bingdontortoise
(generallyknow
nsince
1877as
T.abingdoni),
andthat
heintended
T.duncanensis
asa
newnam
efor
theD
uncanIsland
tortoisenow
thatT.
ephippiumw
asunavailable;
butthis
isassum
ptiononly.
While
disposingofV
anD
enburghs
newnam
esin
thissum
mary
but ambiguous
fashion,Garm
anpresented
severalnew
names
ofhis
own.
Although
aware
(inhis
veryfirst
paragraph)thatG
alapagostortoise
populationsshow
edsuch
individualvariation
asto
requireanalysis
ofsubstantial
seriesto
characterizethem
adequately,he
basedhis
newspecies
Testudo
clivosaon
asingle, unrem
arkablebony
shellw
itha
fewscutes
stillattached.
The
islandof
originw
asunknow
n.S
imilarly,
nolocality
was
givenfor
eitherthe
holotypeor
theonly
otherreported
specimen
ofthe
newspecies
Testudo
typica.T
hetw
ospecim
ens(P
lates22
and34)
seemto
differconsiderably
inrelative
width
andround-
ness,w
hilethe
similarity
ofthe
lateralprofiles
leadsone
tosuspecterror
inthe
measurem
ents(p. 286)
thatindicatethat
thetw
oshells
differby
fiveinches
instraight-line
carapacelength
butby
onlyone
inchin
curvedlength.
The
lengthof
thelarger
(MC
Z5260)
isincidentally
givenas
29inches
on
p.286
butas
28.5inches
inthe
captionto
Plate
34—
acareless
inconsistency,but anunderstandable
errornon
ethe
less;repeated
measurem
entsof
largeturtle
sheHs
rarelyyield
identicalresults.
Garm
analso
proposedthe
name
Testudo
macrophyes
asa
substitutenam
efor
theT
agusC
ovetortoise,
describedearlier
asT.
niicrophyesby
Gunther
(1875,1877)
Ofl
thebasis
ofseveral
specimens
ofknow
norigin
collectedby
J.C
ookson,but
with
aholotype
(BM
NH
75.12.29.1)of
un
known
originthat
Garm
anconcluded
was
abnormal,
aged,and
dwarfed.
Perhaps
Garm
anm
ayhave
beenjustified
indism
issingthis
tortoiseas
asuitable
holotypefor
theT
agusC
ovetortoise,
butin
viewof
hiscondem
nationof
thenorm
alityofthe
specimen
itissurprising
thatheretained
thenam
eT.
rnicrophyesas
avalid
species,apparently
fromsouthern
Albem
arle,ofwhich
T.guntheri,T.galapagoensis,and
evenT.
chatharnensis—
fromthe
farside
ofthe
archipelago—
were
listedas
synonyms.
What
was
notparticularly
surprisingw
asthat
Garm
an’snew
names
(i.e.,T.
inacrophyes,T.
clivosa,T.
typica,and
T.duncanensis)
were
neverused
againby
anyone.A
fewcom
ments
onpoints
raisedby
Garinan
inthe
introductorysection
ofhisw
ork(pp. 261—
270)are
appro
pri
ate.P
articularlyenigm
aticis
theargum
entpresented
atthe
topof
p.262.
inw
hichG
arman
arguesthat
theG
alapagosorganism
sreached
theislands
eitherby
“originin
place,”or
by“accidental
importations;”
hisdiscounting
ofthe
latterpossibility
suggeststhathe
subscribedto
theform
ertheory.
Fig
ure
13.D
orsalview
ofcotype
ofT
estudorotunda,
Latreille,
1802,ahatchling
Rodrigues
tortoise.Photo
byR
.Bour.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises29
Had
thephrase
been“evolution
inplace”
rather than“origin
inplace,”
theargum
ent mighthave
beenm
oreacceptable; as
itstands
itappears
toargue
forthe
independentcreation
offinished
organisms
inthe
Galapagos
Islands—
aview
thatreflected, perhaps,the
influenceofL
ouisA
gassiz(or, through
Agassiz,
evenof
Georges
Cuvier).
On
pp.262—
263,G
arman
reflectedupon
theS
panishand
English
names
byw
hichthe
differentGalapagos
islandsw
ereknow
n,suggesting
thatthe
Spanish
names
were
older,that
Ecuador
stillclung
tothem
,and
thatthey
would
ulti
mately
prevail.Y
etin
factthe
English
names
hadbeen
establishedby
seventeenthcentury
English
piratessuch
asD
ampier
andC
owley,
whereas
theS
panishnam
es,alm
ostallofw
hichreflectsom
easpectofC
olumbus’
voyageto
theA
mericas
(e.g.,the
shipsP
intaand
Santa
Maria;
Queen
IsabelaofSpain;
SanS
alvador,theisland
onw
hichC
olu
mbus
firstlanded;
Genovesa,
Colum
bus’birthplace,
etc.)w
eregiven
onthe
occasionof
thefourth
centenaryof
Colum
bus’firstvoyage
tothe
Am
ericas(i.e.,
in1892)
only25
yearsbefore
Garm
an’sw
orkw
aspublished.
How
ever,som
eS
panishnam
esare
older,Jam
esIsland
havingbeen
renamed
Olm
edaand
Charles
IslandF
loreanaby
Col.
IgnacioH
ernándezin
1832.In
thisw
ork,I
electto
usethe
English
names
(butalso
giveS
panishequivalents
inthe
Sum
mary
andS
urvivalStatus
sections),notjust forreasons
ofchronological
priority(an
important
considerationto
taxonomists), but
alsobecause
thispaper
isnotintended
asa
fieldguide,
hutrather
asan
academic
contributionand
comm
entaryupon
theearlier
biologicalliterature
onthe
Galapagos
tortoises,w
hichin
general(exclusively,
inearlier
works)
utilizesthese
names.
For
exhaustivesy
nonym
iesof
Galapagos
Islandnam
es,see
Black
(1973)and
Woram
(1989).M
ysteriousindeed
isG
arman’s
theorythat
theG
alapagostortoises
hadbeen
introducedby
man.
Acentury
earlier,it
hadbeen
thoughtthat
allisland
gianttortoises,
inboth
theIndian
andP
acificO
ceans,w
ererefer-
ableto
asingle
species,T
estudoindica,
which
would
natu
rallylead
oneto
seekevidence
ofhuman
introductionto
thefar-flung
islandson
which
thespecies
was
supposedto
occur.D
arwin
himself
subscribedto
thisview
inthe
firstedition
ofhisJournalofR
esearches(1839), as
didF
itzR
oy(1839),
buthe
correctedhim
selfin
afootnote
tohis
firstedition, and
inthe
textofhissecond
edition(D
arwin,
I845),in
which
hereferred
tothe
Galapagos
tortoisesas
“Testudo
nigra,form
erlycalled
Indica,”but
alsocom
mented
thatthere
were
“two
orthree
speciesor
racesin
theG
alapagosIslands.”
The
keyinfluence
inthis
revisionof
opinionhad
beenG
abrielB
ibron,w
hohad
addressedthe
Zoological
Society
ofL
ondonon
28F
ebruary1838,
andw
hohad
insistedthathe
hadseen
at leasttwo
speciesoftortoises
fromthe
Galapagos
Islandsalone
(Sullow
ay,1982).
Neverthe
less,as
lateas
1870,John
Edw
ardG
rayw
rote“T
heblack
tortoisesof A
siathathave
beencalled
Testudo
indica, which
arefound
spreadover
allparts
of theA
siaticregion,
alsoon
theislands
of theeast
coastof
Africa,
andin
California
and
Figure
14.Adult m
aleJam
esIsland
(SanSalvador)
tortoise, inthe
western
highlandsof Jam
es.Note
remnantScalesia
pedunculatavar.
pedunculatatrees
ina
habitatlargelyconverted
tograssland
orpampas. February
1971.
30C
HELO
NIA
NR
ESEAR
CH
MO
NO
GR
APH
S,N
umber
I—
1996
theG
alapagos..
. Modern
writers
onthe
subjecthaveunited
theseinto
asingle
speciesunder
thenam
eT
estudoindica”
(Gray,
1870).A
gassiz(1
857) failedto
noteD
arwin’s
I845co
rrection,
referringto
thescientific
name
ofthe
“Gallapago
turtle”as
Cylindrapis
[sic]indica,
i.e.,still
utilizingthe
“universal”
specificepithet for
insulargianttortoises.
Cylindraspis
hadbeen
proposedas
anew
genusby
Fitzinger
(1835),w
ith
subsequentdesignation
ofthe
Rodrigues
saddlebackto
r
toise(T
estudovosm
aeri)as
thetype
species(Fitzinger,
1843).Garm
anclaim
edthat
severaldistinctspeciesw
erep
ecu
liartovarious
oftheG
alapagosIslands,butthis
leavesentirely
unanswered
thequestion
ofthe
originalhabitat
ofw
hathe
consideredm
ayhave
been“introduced”
populations.A
ctually,thesuperficialsim
ilaritiesbetw
eenthe
Indian
Ocean
andthe
Galapagos
tortoises—
largesize,unpatterned
darkshells,enlarged
shellopenings,long
necks,absenceof
osteo
derm
sin
thelim
bscales,
shortenedplastra
—are
striking,and
would
surelylead
am
oderncladistic
phylogeneticistunschooledin
zoogeographicconsiderations
orother
specialknowledge
ofthe
groupto
considerthem
atrue
(monophyletic)
dade,evenif nota
singlespecies
.Even
thevariation
within
thegroups
—forexam
ple,thedevelop-
ment
ofsaddlebacked
shellsam
ongsom
eof the
Galapagos
forms
andalso
inthe
IndianO
ceanspecies
Cylindraspis
vosmaeri
—is
remark
ably
parallel;
sadd
leback
edsh
ells
app
earto
bea
shared
deriv
edch
aracteror
synap
om
orp
hy
ifever
therew
asone.
As
arem
arkablelatter-day
comm
entaryupon
theparal
lelismbetw
eenthe
Galapagos
andthe
Mascarene
tortoises,
RogerB
ouradvises(in
litt.,19
Decem
ber1989)
thatthetw
ohatchling
tortoises,preserved
inspirit
inthe
MN
HN
Pco
llections
(Fig.13),
onw
hichL
atreille(1801)
basedhis
descrip
tion
of
Testudo
rotunda,are
infact
Mascarene
tor
toises,
pro
bab
lyfro
mR
od
rigu
es,w
hose
torto
isesw
erestill
marginally
extantwhen
Latreille
wrote
hisaccount.
Infact,
thesam
especim
enshad
earlierbeen
figuredby
Lacépède
(1788)—proofpositive
that theyw
erecollectedin
or before1788.
They
were
notG
alapagostortoises,
asB
ourhim
self
( 1980)had
earlieridentifiedthem
,andas
theyhad
alsobeen
consideredby
thosefew
twentieth
centurysystem
atistsw
hohad
takenany
noteof
theexistence
ofthis
name.
Bour’s
comparisons
hadbeen
hinderedby
thefactthatno
collection
hasu
neq
uiv
ocal
Mascaren
eto
rtoise
hatch
ling
s,an
dthe
Paris
collectionlacked
hatchlingsfrom
theG
alapagosalso.
Moreover,the
name
1’. rotundaoffers
littleclue
(rotundityis
acharacteristic
of alltortoisehatchlings);and
theactions
ofthe
review
ersin
theyears
follo
win
g1801
were
decid
edly
confusing.T
henam
eis
anim
portantone
—if
it were
basedupon
aG
alapag
os
torto
ise,it
isth
eo
ldest
availab
lenam
efo
ra
Fig
ure
15.Adultm
aleJam
esIsland
tortoise,from
thew
esternhighlands
ofJam
es.T
heanterior
elevationof
thecarapace
isgreater
than
istypical
ofthis
population.F
ebruary1971.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises31
mem
berof
thatgroup,has
extanttypem
aterial,and
isnot
anonien
obliturn.Schw
eigger(1812)notedthatL
acépèdehad
consideredthis
speciesto
bea
synonymof
theE
uropeanpond
turtle,T
estudoorbicularis
orE
rnyseuropaea
(bothof
which
names
herendered
inthe
genitiverather
thanthe
nominative
case),but
hedisagreed
vigorouslyw
iththis
conclusion,observing
thatthe
speciesw
asclearly
entirelyterrestrial,
andhe
utilizedthe
name
Testudo
rotundain
unalteredform
,as
avalid
species.N
evertheless,M
errem(1
820)continued
toconsider
Testudo
rotundato
bea
syn
onymof
Em
ysorbicularis,
anerror
thathas
survivedto
recenttim
es(W
ermuth
andM
ertens,1977).
Schw
eigger’sdescription
of theholotype
ofT.
rotundaincluded
severalusefuldiagnostic
characters.For
example,
thestatem
ent“bracteae
marginales
vigintitres,
anticaeaequales”
(marginals
23,the
frontonesequal)
indicatethat
thenuchalw
asabsent,
asin
bothG
alapagosand
Mascarene
butnotSeychelles
tortoises.On
theotherhand,the
statement
“sternum...in
duodecimareas
divisuin”(12
plastralscutes)
would
favorthe
Galapagos
ratherthan
theM
ascareneto
rtoises;
butthisis
notanecessary
conclusion.Although
most
adultM
ascarenetortoises
havea
singlegular
(orfused
gulars),givingeleven
plastralscutes,certainsubfossilplastra
fromM
auritiusattributedto
Cylindraspis
triserratahave
thegulars
pairedand
indeedanteriorly
extendedand
divergent(B
our,I985).
The
gularis
alsooccasionally
dividedin
the
Reunion
Cylindraspis
(R.
Bour,
pers.com
m.),
andis
do
rsally
dividedin
theyoung
ofAstrochelysyniphora
andin
theholotype
ofC
ylindraspisgraii.
Furtherm
ore,turtle
speciesare
known
inw
hicha
median
epidermal
shellelem
entis
dividedin
theyoung,
butby
aprocess
ofgradual
fusionbecom
esontogenetically
undivided.T
hisoccurs
especiallyin
areasw
herethe
scutesgrow
bydistalratherthan
proximal
accretionofnew
substance,asis
thecase
with
theanalscutes
ofcertain
Cuora
species,or
thesupracaudal
scuteof
Peltocephalus.
Som
eof
theknow
nadult
shellsof
theR
odriguesto
rtoise
showactual
separationof
theanteriorm
ostpair
ofm
arginalscutes,with
thefirstvertebralscute
forming
part ofthe
anteriorm
arginof
thecarapace.
How
ever,this
ispro
bably
againan
ontogeneticcharacter,
associatedw
iththe
developmentof the
raised,saddledanteriorpartofthe
shell,and
was
apparentlyexacerbated
alsoin
adultsby
damage
tothis
areapossibly
resultingfrom
intraspecificbiting
orfighting,
with
resultantloss
ofshell
material
andform
ationof
scartissue
(pers.obs.).
Important
inS
chweigger’s
de
scriptionis
thereference
tothe
spuron
theend
ofthe
tail(“C
audacrassa
exserta,apice
corneo”),a
massive
tailspur
beinga
featureof
adultm
aleM
ascarenetortoises.
Butdespite
thisconfusion,the
skullsalone
aresufficiently
differenttojustifyseparation
ofthethree
gianttortoisestocks
atthe
genericlevel
—G
eochelonefor
theG
alapagosform
s,
Figure
16.Old
adultmale
tortoiseundera
barbedw
irefence
incattle
pasturenearS
antaR
osa,IndefatigableIsland
(Santa
Cruz).T
hepeeling
textureofthe
carapacescutes
isprobablyaresultofinfestation
with
fungalandotherm
icroorganisms
inthis
veryhum
idenvironm
ent.July1986.
32CHELONTAN
RESEARCH
M0N
ouRA
PIIS,N
umber
I—
1996
Fig
ure
17. Holotype
(CA
S8127)
ofTestudo
chathamensis,
Van
Denburgh, 1907, from
southwestern
Chatham
Island(San
Cristóbal).
Photofrom
Van
Denburgh
(1914),plate64, figure
2.
Aldabrachelys
forthe
Aldabra-S
eychelles-Madagascar
tor
toises,and
Cylindraspis
forthe
Mascarene
forms.
onpp. 264—
266G
arman
discussedhis
beliefthatpart ofthe
problemin
c’assifyingG
alapagostortoises
layin
thelikelihood
thattortoises
were
extensivelytransported
fromone
islandto
another,resulting
inhybrid,
morphologically
confusingpopulations.
Such
inter-islandtransport
ofto
r
toisesundoubtedly
occurredfrom
time
totim
e,but
onnothing
likethe
scaleoftortoise
transport between
islandsof
theIndian
Ocean. T
hem
orphologyofthe
survivingp
op
ula
tionsdoes
not showevidence
ofgeneticcontam
inationof the
different islandstocks,
althoughin
afew
cases(such
asthat
ofthe
Volcan
Wolf
tortoise)the
morphological
variationseen
todayis
somew
hatdifficult
toexplain.
The
main
element
ofuncertainty
presentedby
theinter-island
trans
portationpossibility
relatesprim
arilyto
theextinct form
s—
toknow
ingw
hetheror
not driedbones
orsubfossil
remains
foundon
suchislands
asC
harles,B
arrington,and
Jervisare
trulyreferable
toendem
ic,extinct
subspeciesrather
thanto
tortoisesbrought
fromother
islands.O
npp.
267—269, G
arnunentered
adetailed
discussionof the
topicof
sloughing,w
ithexam
plestaken
fromvarious
unrelatedreptiles,
includingrattlesnakes
andthe
flatbacksea
turtle.S
ome
freshwater
turtles,especially
einydidsand
chelids,m
ayslough
theouter
layero
fa
growing
scutein
a
singlepiece,
hutthis
doesnot
occurw
ithG
alapagosto
r
toises. The
questionofw
hy
thescutes
ofG
alápag
os
tortoises
differin
textureand
surfacesculpturing
isa
complex
one.
discussedto
some
degreeby
Fritts
(1984),but
Garm
an’s
conclusionsthat
thecarapacial
textureof
Galapagos
tor
toisesis
aproduct
of
thesloughing
of
entirescutes
doesnot
appeartobe
basedon
actual observation, nor issuch
slough-ing
known
tooccur
inthese
tortoises.T
hecorrect
interp
re
tationis
probablythat
smooth-shelled
individualsbecam
em
echanicallyabraded
toa
smooth
conditionafter
theyreached
maturity,
althoughperhaps
smaller,
chronically
undernourishedtortoises
may
alsoshow
smooth
shellsat
times.
The
former
hypothesism
ayexplain
inpartw
hyadult
female
Galapagos
(andA
ldabra)tortoisesareusually
smooth-
shelled.H
owever,
itisunlikely
tobe
thecom
pleteex
plan
a
tion,since
abrasionby
contactw
iththe
plastronand
theforelim
hsofthe
male
duringcopulation
would
not affect allparts
ofth
ecarapace
equally.T
hequestion
isnot peculiar
to
Galdpagos
tortoises;m
anylong-lived
chelonianspecies,
especiallyterrestrial ones
(includingsuch
generaas
Gop
herusand
Terrapene),
may
includeboth
smooth-shelled
adultsand
othersw
ithw
ell-sculpturedannuli.
Fig
ure
18.Y
oungadultm
aletortoise
surveysthe
floodingof the
lowlands
of northeasternC
hathamIsland
(SanC
ristobal)during
theE
lN
iñoepisode
of1972.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises33
Garm
an’sdiscussion
onp.
269ofpossible
differencesbetw
eenG
alapagostortoises
collectedin
earlyand
inrecent
yearsis
exceedinglyinteresting.
Fritts
(1983)w
rotethat
“preliminary
comparison
ofm
useumspecim
enscollected
duringtheperiod
1890-4910w
ithspecim
ensfoundliving
inG
alapagosin
1976and
1977indicate
thatpopulationdiffer-
encesare
stableand
donotvary
with
short-termclim
aticor
ecologicalfluctuations.”Y
etbothG
arman’s
andsom
em
orem
odernobservations
suggestotherw
ise,and
factorsm
orecom
plexthan
theprogression
ofthe
Duncan
Islandtortoise
populationfrom
avigorous
oneat the
turnofthe
centuryto
asenile
one(w
ithrepatriated
younganim
alspresent,
butnone
intheir
middle
years)m
ayhave
beenat
work.
For
example,
Garm
anquotes
Porter’s
(1815)account
ofthe
round,plum
p,black,
andthin-shelled
tortoisesof
James
Island(F
ig.14),
whereas
acentury
laterV
anD
enburghreported
thetortoises
onthis
islandto
belarge,
heavy,thick-shelled,
andsom
ewhat
saddlebackedrather
thanrounded.
Furtherm
ore,m
yow
nobservations
inrecent
yearssuggest
thatsom
echanges
havetaken
placesince
Van
Denburgh’
5specim
ensw
erecollected
atthe
beginningof
thiscentury.
Insom
ecases
thechanges
may
relateto
alterationsin
thepopulation
structure.F
orexam
ple,a
century
ago,the
tortoisesof
James
andIndefatigable
Islandsw
ereregularly
preyedupon
bysettlers,
andreally
large,oldanim
alsm
ayhave
beenvery
scarce.T
oday,the
threatis
primarily
uponthe
hatchlingandjuvenile
stages(e.g.,by
rat,pig,ordog
predation),andlarge
animals
areessentially
safe.T
headult
specimens
foundtoday
onJam
es(Fig.
15)and
onIndefatigable
(Fig.16)
includeindividuals
largerthan
any
reportedby
Van
Denburgh,
andm
anym
aybe
seenw
ithsm
ooth,unstriatedshells,w
hereaseven
theadults
collectedby
Van
Denburgh
showed
theshell
sculpturingtypical
ofyounger
adults.T
hetortoises
reintroducedto
Duncan
andH
oodare
growing
largerthan
theirparents
(Rhodin
etal.,
1983;P
ritchard,1985).
The
tortoisesof
Chatham
Island,reportedto
beflat-shelled
byV
anD
enburgh(Fig.
17),are
nowsignificantly
saddlebacked(Fig.
18)(indeed,
asdescribed
below,they
areofa
differentpopulationaltogether,although
onthe
same
island,and
curiouslysharing
thefrequent
anomaly
ofpectoralscutes
thatfailto
meetin
them
idline,a
detailrecordedby
Van
Denburgh,and
which
Ifoundin
9of
18specim
ensexam
inedin
thefield).
And
thetortoises
ofV
olcanW
olf(Fig.
19),described
assaddlebacked
byV
anD
enburgh(although
with
considerablevariation
inthe
de
greeofanterior
elevationofthe
carapace)are
nowknow
nto
includeboth
domed
andflattened
individualsas
well
asa
wide
varietyof
saddlebackedspecim
ens.T
hecauses
ofthese
differencesrem
ainconjectural.
Som
ehypotheses
arepresented
elsewhere
inthis
paper.The
verylarge
sizeof
theadult
male
tortoiseson
James
andIndefatigable
todaym
aystem
fromthe
changein
populationstructure
discussedabove,
butalso
may
beassociated
with
theprogressive
conversionoflarge
areasofthe
highlandsof
theseislands
intolush
savannahsor
pasturelandslocally
known
aspampas
(Fig.14).T
hesehave
developedas
aresult
ofhum
anland-clearing
forpasture
onindefatigable
Island.and
offeral
goats,present
intens
ofthousands,
destroyinglarge
amounts
ofnative
uplandvegetation
onJam
es.T
heecological
changesw
roughthave
beenserious
andm
any
Figure
19.A
dultfemale
Volcan
Wolf
tortoisenesting
ina
small
patchof
soilsurrounded
bypahoehoe
lavainland
fromB
ank’sB
ay,A
lbemarle
Island(Isabela).February
1971.
34Cf1EIoN
1AN
RESEARCHM
oNouR
APTs,
A/uniber
1—
1996
plantspecies
may
disappear.B
ut,as
acurious
secondaryresult,
thegrasslands,
thathave
developedat
cool,high
altitudesatw
hichtortoises
arerarely
indanger
fromex
cessive
insolationeven
when
noshade
isavailable,
provideexcellent
tortoisepasture,
andthe
individualsthat
grazethere
aregrow
inglarge
andfataccordingly.
There
seems
tohe
noevidence
forthe
speculationofF
ritts( 1984)
that cool,m
oist(i.e.,
upland)habitats
may
resultin
aslow
ingof
growth
ofall
adults(m
aleand
female),
althoughthere
may
beevidence
thatsom
eindividuals
donot
move
tosuch
altitudesuntil
alreadyw
ell-grown.
Peter
andR
osemary
Grant
andtheir
studentsand
co
workers
studiedthe
finches(G
eospizinae)of
theisland
ofD
aphneM
ajorfor
more
thantw
odecades
(Weiner,
1994).T
heirextraordinarily
detailedobservations
indicatedthat,
among
thesesm
allbirds, reproductioncould
occuratan
ageof
lessthan
threem
onthsin
especiallyrainy
years,and
thatthe
populationson
Daphne
showed
fluctuationsor
pro
gres
sions,especially
inbeak
dimensions
andoverall
bodysize,
ona
year-to-year basis. Detectable
microevolution
atsuch
atem
pow
ouldnot
beexpected
among
Galapagos
tortoises,that take
atleast two
decadesto
reachfirstm
aturityand
thatshow
greatindividual resistanceto
stressful conditionssuch
asprolonged
drought(conditions
thatquickly
decimate
finchpopulations, leavingpresum
ablyonly
thebest-adapted
individualsas
survivors). Nevertheless, the
same
phenomenon
couldconceivably
beoccurring
inthe
variouspopulations
ofG
alapagostortoises
ona
veryprotracted
scale—
measurable
onlyover
decadesrather
thanover
months
oryears.
Garm
anw
entto
some
lengthto
redescribethe
speciesT
estudotab
ulata,
basinghis
descriptionlargely
uponan
allegedlyP
uertoR
icanspecim
enof
which
heprovided
photographs.A
lthoughcurrent
opinionsuggests
thatthis
modem
species, while
relatedsubgenerically
tothe
Galapagos
tortoises, iscertainly
not ancestral tothem
,Garm
an(p. 272)
thoughtotherw
ise.T
henam
eT
estudotab
ulata
was
pro-posed
byW
albaum(1782),
butthis
work
was
notco
nsis
tentlybinom
ial inits
nomenclature, and
theL
atinizednam
esincluded
areonly
citedin
passim
,astranslations
ofvern
acular
names
(Vanzolini,
1977),and
arethus
notconsidered
availablefornom
enclaturalpurposes. The
specieslong
known
asT
estudotabulata
isin
facta
composite
oftw
ospecies,
Geochelone
denticulataand
G.
carbonaria,
whose
distinct-ness
was
overlookedby
most
twentieth
centuryauthorities
untilthe
matter
was
finallyclarified
byW
illiams
(1960).M
oreover,the
specimen
selectedby
Garm
anfor
de
taileddescription
was
aconfusing
one.Its
statedorigin
of“P
uertoR
ico”is
clearlym
istaken.W
hilethe
specimen
may
conceivablyhave
beenpurchased
there,no
tortoisesare
nativeto
Puerto
Rico,
nordo
Schw
artzand
Thom
as(1975)
norP
ritchardand
Trebbau
(1984)report
evenintroduced
tortoisesO
flthat island. The
specimen
itselfislarge
CL
57cm
andon
thosegrounds
would
appearto
bereferable
tothe
largerof
thetw
ospecies,
Geochelone
den
ticulata,
with
G.
carbonaria
notbeingknow
nto
exceed5
1 .2cm
CL
andusually
much
srnaller(Pritchard
andTrebbau,
1984). On
theother
hand,the
uniformly
blackcarapace
with
asingle
yellowspot
onthe
areolaof
thesecond
vertebral,is
more
typical ofG.
carbo
naria,
althoughin
thisspecies
theyellow
spotsare
usuallypresent on
eachofthe
vertebralsand
costals(but
uniformly
blackspecim
ensm
ayoccur
inB
olivia).O
neincidental com
ment ofG
arman’ sprovides
interest-ing
insightintothe
slowness
ofcomm
unicationbetw
eenthe
Galapagos
andthe
United
States
atthe
beginningof
thecentury.
Garm
an(p.
265)com
mented
onthe
Cobos
settle-m
ent onC
hathamIsland, and
observedthatit“m
ayyetbe
inexistence.”
Infact,
thiscolony
hadcom
eto
anend
thirteenyears
earlier,in
1904,w
henC
obos’ow
npeons,
tiredof
histyrannical
andviolent
rule,had
murdered
himw
iththeir
machetes.
But
despitethe
defectsof
Garm
an’sw
ork,it
alsohad
positiveaspects.
Som
eof
Garm
an’staxonornic
insight,although
oftendiscounted
inrecent publications,
appearsto
bevalid.
Garm
anw
asin
some
casesable
todiscern
thevalidity
ofspecies
names
(suchas
C’helonia
(=N
atatoñ
depressaand
Testudo
(=G
eochelone)nig
ra)that were
ignoredby
hiscontem
porariesand
onlyrestored
tovalidity
decadesafter his
death. His
discussionofthe
apparent changesin
theisland
populationsw
ithtim
ehas
similarly
beenrecently
vindicated.F
romthe
pointof
viewof
exploringthe
taxo
nomic
historyof
theG
alapagostortoises,
Garm
an’sex
tensive
seriesof
platesare
most
valuable.A
ndhis
conclusionthat
thetype
specimen
ofT
estudoephippiurn
came
fromA
hingdonIsland. in
agreement
with
Baur
(1889), notCharles
orIndefatigable
ashad
beenassum
edby
Gunther
U877).
norD
uncanIsland
asassum
edby
Van
Denhurgh
(1914)
andvirtually
allm
odemauthors,
seems
tohave
beencorrect.
Curren
tN
omenclature
Van
Denhurgh
( 1914)recognized
14species
ofG
alapagostortoises
( 13of
themnam
ed),all
ofw
hichhe
includedin
thegenus
Testudo.
Van
Denburgh
hadavailable
tohim
theseries
of256
specimens,
ofknow
nisland
origin,recently
collectedby
theA
cademy
expedition,and
hew
asthus
ableto
describeand
illustratethe
variousgeographi
callyisolated
forms
andto
preparea
keyfor
theiridentifica
tion. Van
Denhurgh
recognizedfive
speciesfrom
Albem
arleIsland
(T.vicina,
T.gilniheri,
T.inicrophyes,
T.becki,
andthe
unnamed
formfrom
Volcan
Alcedo), and
oneeach
fromN
arborough( T. p
han
tastica),Jam
es(T
darwini), Jervis
(T.
wailacei), Indefatigable
(T. porteri), D
uncan(T.ephippium
),A
bingdon(T.
ahingdoni),C
hatham(T.
chathumensis),
andH
ood(T
.h
ood
ensis).
The
Charles
Islandspecies
(T.
gala
pag
oen
sis)and
theform
fromB
arringtonIsland
were
foundto
beextinct.
Testu
do
ph
anta
stica
was
basedupon
asingle
specimen
(Fig.20),
fortunatelyadult;
theim
mature
specimen
fromV
olcanA
lcedolacked
distinctivecharacter-
istics(although
unusuallyhigh-dom
ed),and
was
wisely
leftunnam
ed.V
anD
enburghfound
hissingle
specimen
fromJervis
Islandto
hesom
ewhat sim
ilarto
Rothschild’s
typeof
T.w
allaeei,and
hethus
associatedthis
name
with
theJervis
thrmeven
thoughthe
two
specimens
inquestion
lackedany
remarkable
orobvious
identifyingfeatures.
andthere
was
II
PRITCHARDT
heG
alapagosT
ortoises35
considerablesuspicion
thattheone
specimen
known
tohave
actuallybeen
foundon
Jervishad
beenintroduced
(byG
.B
aur).Apart
fromthe
curiouschanges
wrought
byG
arman
(that foundlittle
acceptance), subsequentlypublished
mo
di
ficationsoradditions
toV
anD
enburgh’s classificationhave
beenas
follows:
i)D
eS
ola(1930),
havingcollected
aseries
oflive
tortoisesfrom
thevicinity
ofC
artagoB
aynear
Perry
Isthm
us,A
lbemarle
Island(in
thecoastal
lowlands
between
Volcan
Alcedo
andV
olcanS
ierraN
egra), declaredthese
tobe
referableto
anew
species,Testudo
vandenburghi.U
nfo
rtunately,
De
Sola
providedneither
adiagnosis
ofthis
formnor
adesignation
ofthe
typespecim
en.ii)
Mertens
andW
ermuth
(1955)reduced
thevarious
islandform
sto
subspeciesw
ithinthe
speciesT
estudoelephantopus,
andsubstituted
theolder
name
T.nigrita
forT.
porteri.M
ertensand
Werm
uthconsidered
Testudo
californianaand
1’.nigra,bothofQ
uoyand
Gaim
ard,1824
(butpublishedin
two
papers,theone
afew
months
beforethe
other),to
bequestionable,
andgave
preferenceto
thelater
name
T.elephantopus
Harlan,
1827.T
heseauthors
alsoconsidered
thefive
describedform
sfrom
Albem
arle(or
six,if
we
includeT.
macrophyes
Garm
an,1917)
asidentical,
listingall
underT.
e.elephantopus.
iii)L
overidgeand
William
s(1957)
establish
edG
eocheloneas
theappropriate
genusforthe
Galapagos
(andm
anyother)
tortoises,the
Galapagos
forms
beingincluded
(asG
.elephantopus)
with
theS
outhA
merican
mainland
speciesin
thesubgenus
Chelonoidis
(renderedincorrectly
asC
helonoidesby
Auffenberg,
1967and
1971).iv)
Werm
uthand
Mertens
(1961),in
arevision
oftheir
inventoryof
thew
orld’sliving
turtlespecies,
continuedto
useT
estudofor
theG
alapagostortoises.
v)In
afurther
revisionpublished
afterthe
deathof
Mertens,
Werm
uthand
Mertens
(1977)still
usedT
estudo,but
includedthe
Galapagos
tortoisesin
thesubgenus
Chelonoidis.
Moreover,
althoughin
previouseditions
theyhad
listedthe
varioussubspecifically
indeterminate
names
forG
alapagostortoises
assynonym
sof
T.elephantopus,
inthe
I977edition
thesew
ereallincluded
inthe
synonymy
ofT.e.elephantopus.T
hecopious
literaturebetw
een1961
and1977
listingthe
Galapagos
tortoisesas
forms
within
thegenus
Geochelone
was
ignoredeven
inthe
synonymies,and
certainsubspecies
known
tobe
living(those
ofA
bingdonand
Duncan
islands),as
well
asthe
long-extinctC
harlesIsland
form,
were
alllabelled
aspresum
ablyextinct
(“vermutlich
ausgestorben”).N
osuch
labelwas
attachedto
theone
subspecies(phantastica)
thatmay
indeedbe
extinct,but
thathas
notyet
beenproven
tobe
so.vi)
Bour
(1980),in
ashort
paperbased
more
ona
philosophicalstriving
fortaxonom
icbalance
within
thefam
ilyT
estudinidaerather
thannew
biologicaldata,
pro-m
otedthe
subgenusC
helonoidisto
genericstatus,
andw
ithit
elevatedthe
varioussubspecies
ofG
alapagostortoises
tofull
speciesstatus
within
thatgenus.
Fig
ure
20.A
nteriorand
posteriorview
sof
holotypeof
Testudophantastica
(CA
S8101).
36CH
ELON
IAN
RESEARCH
MO
NO
GRA
PHS,
Num
ber1
—1996
Figures21
and22.
Dorsal
andventral
views
ofthe
holotypeof
Testudoephippium
inthe
Royal
Scottish
Museum
,E
dinburgh.P
hotocourtesy
AS
.C
larke,R
oyalS
cottishM
useum.
vii)C
rumly
(1982)considered
theG
alapagostortoises
tobe
subspeciesof
Geochelone
elephantopus,but
subse
quentlyreversed
himself
(Crum
ly,1984),
arguingfor
fullspecies
statuso
feach
population,ironically
ina
paperdem
onstratingthe
virtualimpossibility
ofdistinguishingthe
islandpopulations
byskull
analysis.C
rumly’
s justificatio
nw
asthat
Pritts
(1983,1984)
consideredV
anD
enburgh’sanalysis,
thatused
binomials,
tobe
theauthority
ofchoice
untilsuperceded.
viii)F
ritts(1983),
ina
studyof
thecom
parativem
orphologyof
Galapagos
tortoises,largely
(andpro
bably
wisely
)refrain
edfrom
providingscientific
names
forthe
differentpopulations,
with
thenotable
exceptionof
thecaption
toa
figurein
which
thenew
combinations
Geo
chelo
ne
abin
gd
on
i,G
.becki,
G.
darw
ini,
G.
epphip
ium
[sic].and
G.
porteri
were
perpetratedw
ithoutcom
ment.
Further
informal,
perhapseven
unrealized,new
combinations
(Geochelone
chathamensis.
G.
guntheri,G
.vicina,G
. hoodensis,G. m
icrophyes,G. nigrita,G
. wallacei,
G.
phantastica,and
G.
vandenburghi)w
erepublished
byC
rumly
(1984).ix)
Pritchard
(1984),in
afootnote
toa
paperon
theA
hingdonisland
tortoise, observedthatthetype
specimen
ofT.
ephippium(F
igs.21
,22),as
hadpreviously
beenargued
byB
aur(1
889),alm
ostcertainly
came
fromA
bingdonIsland,
notD
uncan;and
thatthe
name
T.nigra
Quoy
andG
aimard,
1824b,w
aspublished
threeyears
earlierthan
T.elephantopus
Harlan,
1827,w
asnot
anom
enoblitum
,and
thereforew
asthe
validspecific
epithetfor
theG
alapagos
tortoises(and
hadthe
incidentaladvantage
ofan
extantholotype).
x)B
our( 1985)
partiallyreversed
hisearlier
positionthat
theG
alapagostortoises
were
allfull
species,consider-
ingthem
acom
plexof
speciesand
subspecies,and
agreedw
ithP
ritchard(1
984)thatnigra
was
theappropriate
specificepithet
forthis
complex.
Inview
ofthisconfusion,
itisclear
thatanew
analysis,from
firstprinciples,
with
referenceto
thelatest
editionof
theC
odeof
Zoological
Nom
enclature(IC
ZN
,1985),
andusing
modern
conceptsof
speciesand
subspecies,is
neces
saryto
establish
anap
pro
priate
nom
enclatu
refor
theG
alapagostortoises.
This
shouldfollow
thesesteps:
1)D
etermination
ofthe
appropriategeneric
orsubgeneric
name
forthe
group.2)
Determ
inationof
thebiological
relationshipbetw
eenthe
variouspopulations.3) S
trictappli
cationof
theIC
ZN
Rules
ofN
omenclature
tothe
hio
logi
callydistinguishable
populations. Elaboration
ofth
esethree
stepsfollow
s:1)
Itm
ustbe
observedthat
genusis
asom
ewhat
more
subjectiveconcept
thanspecies,
andthat
notruly
objectiveor
biologicaldeterm
inationm
aybe
possible.M
ayr(1963)
alludesto
anum
bero
fisland
“monotypic
genera”thatturned
outtobe
little(ornothing)
more
thansubspecies
ofcon
tinen
taTform
s.and
observed,onthe
otherhand, thatintraspecific
‘ariationin
some
vertebratespecies
may
includesuch
fun-dam
entalcharacters
asnum
berof
cervicalvertebrae
ornum
berof
digits.B
our’s(1980)
proposalthat
thesubgenus
Chelonoidis
heraised
togeneric
rankw
asfollow
edby
..
•.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises37
Pritchard(1984)
primarily
onthe
groundsthat
aproposed
taxonomic
change, iffoundunacceptable
byanother w
orker,shouldbe
refutedratherthan
ignored, aprincipleclarified
byL
azelland
Michener
(1976).H
owever,
theG
alapagosto
rtoises
andtheir
SouthA
merican
relativeslack
them
ajorshared
derivedfeatures
thatdistinguish
theSeychelles!
Aldabra/M
adagascartortoises
(notablythe
extreme
skullm
odifications),the
Mascarene
tortoises(single
gular,by-
pertrophiedodontoid
tubercies,labial
borderof
mandible
stronglydentate,
extensiveshell
andskull
ankylosis—
atleast
inthe
veryold
adultsthat
constitutem
ostof
theav
ailable
material),
orthe
gen
eraM
anouria
andIndotestudo,
andrecent
authorssuch
asF
ritts(1983,
1984), Crum
ly(1985),
andPritchard
andT
rebbau(1984)
havenot accepted
Bour’s proposal, although
Obst (1985)
did.The
decisionof
Loveridge
andW
illiams
( 1 957)to
listthe
Galapagos
tortoisesas
mem
bersof
thesubgenus
Chelonoidis
within
thegenus
Geochelone
was
nota
un
ani
mous
oneby
thesetw
oauthors.
William
sprevailed,
while
Loveridge
disclaimed
ina
lengthyfootnote.
Nevertheless,
sinceboth
therevived
genusG
eocheloneand
thesubgenus
Chelonoidis
were
adequatelycharacterized
morphologi
callyin
thispaper, the
proposal shouldbe
considereda
validone
eventhough
thesubgenus
Chelonoidis
was
onlyfound
todiffer from
thesubgenus
Geochelone
(fromA
fricaand
theIndian
subcontinent) intw
ocharacters :the
concealedpostotic
fenestraofthe
former and
thenarrow
edpectoral scutes
of thelatter.
Insom
eG
alapagostortoises, the
pectoralsm
aybe
sonarrow
edas
tofailto
make
midline
contact;but tortoisesof
thesubgenus
Geochelone
havepectorals
that arenot m
erelynarrow
edbutthatshow
acharacteristic, parallel-sided
form,
althoughthere
ism
uchvariation
andthe
character isa
weak
oneatbest.
There
isconsiderable
ontogeneticinfluence
inthe
development of narrow
edpectorals, and
thevariation
ofpectoral narrow
ingw
ithinthe
extinct genusC
heirogaster isillustrated
anddiscussed
byB
ourgatand
Bour
(1983).It
appearsto
beassociated
with
aphylogenetic
shorteningof
thebridge
andexpansion
ofthe
humeral
scutes.U
seofthe
subgenusC
helonoidishas
beenaccepted
bythe
majority
ofsubsequent authors. Subgenericnam
esbeing
optional, thepublication
ofthebinom
ial without the
subge
nusdoes
notconstitutedisagreem
ent with
theoriginal
pro-posal.
The
conspicuousexceptions
tothis
usagehave
beenW
ermuth
andM
ertens(1961,
1977)w
hopersisted
with
Testudo
forthe
Galapagos
tortoises,w
ithoutpublishing
ajustification
fortheir practice,andindeed, by
failingto
evenlist
combinations
involvingG
eocheloneor
Chelonoidis
intheir
synonymies,
seemingly
declinedto
acceptthat
thesenew
allocationshad
evenbeen
proposed.In
my
opin
ion,
thearg
um
ents
establish
ing
Geochelone
Fitzinger,
1835,as
theappropriate
genusfor
theG
alapagostortoises, and
Chelonoidis
Fitzinger,
1856,as
thesubgenus,
areadequately
documented
andaccept-
able.T
hegeneric
names
Chersine
Merrem
,1820,
andT
estudinitesW
eiss,1830, both
havechronological prior-
ityover
Geochelone.
How
ever,C
hersineis
onlyin
part
asenior
synonymof
Geochelone,
andL
indholm(1929)
hasdesignated
Testudo
graecaas
itstype
species. There-
foreC
hersine(partim
)as
asynonym
ofG
eochelonedoes
notaffect
thestability
ofthe
latter.T
estudinitesW
eiss,1830, is
definitelybased
uponS
outhA
merican
material,
butitssuppression
asa
longunused
name
uponappeal to
theInternational
Com
mission
onZ
oologicalN
om
encla
ture(IC
ZN
)is
assured.If
theG
alapagostortoises
shouldbe
consideredsubgenerically
(orgenerically)
distinctfrom
theSouth
Am
ericanm
ainlandform
s,a
newnam
ew
ouldhave
tobe
proposed.T
henam
eE
lephantopus, usedby
Gray
(1873)as
agenus,has
beenproposed(Bour, 1988),but,as
Bourpoints
outSthis
name
isajuniorhom
onymofthe
coelenteratenam
eE
lephantopusA
gassiz,1846,
andis
thereforenot
availablefor
theG
alapagostortoises.
2)R
egardingthe
biologicalrelationships
between
thevarious
populationsof
Galapagos
tortoises,the
following
pointsare
relevant:a) In
noknow
ncase
cantortoises
ofdifferent Galapagos
populationsbe
morphologically
distinguishedas hatchlings,
Fig
ure
23.Probably
thelargesttortoise
inthe
world
today,a
male
indefatigableIsland
tortoisew
eighingover
350kg
atthe
Life
Fellow
shipR
anchnear
Seffner,F
lorida. February
1993.W
eight inO
ctober1995
was
400kg.
38C
HE
LO
NIA
NR
ES
EA
RC
HM
ON
OG
RA
PH
S,
Num
berI
—1996
Figure
24.Juveniletortoises
(ca.30cm
carapacelength), from
Hood
Island(left)
andfrom
Cerro
Azul, A
lbemarle
Island(right),captive
raisedunderidentical conditions
attheC
harlesDarw
inR
esearchStation. N
otethat the
elevationofthe
anterior endofthe
carapaceisalready
evidentinthe
Hood
Islandtortoise. indicating
geneticcontrol of the
saddlebackedtrait.
September
1982. Photoby
A.G
.J. Rhodin.
andin
many
cases(including
thegreat
majority
ofzoo
specimens), the
adultstoo
cannotbeattributed
toa
particularisland
oforigin
with
anyconfidence.
While
Van
Denburgh
( 1914)
didpublish
akey
byw
hichthe
differentisland
populationscould
beidentified,
thisw
asin
many
casesbased
uponvery
small
sample
sizes,and
inthree
cases(phantastica,
wallacei,
andthe
unnamed
formfrom
Volcan
Alcedo)
onlysingle
specimens
were
available.b)
Genetic
studiesby
Marlow
andP
atton(1
981)have
revealedextrem
elyclose
relationshipsbetw
eenthe
variousG
alapagostortoise
forms.
C)
Although
Van
Denburgh
(1914)used
binomials
forthe
differentpopulations,he
was
writing
atatim
ew
henthe
subspeciesconcept w
aslittle
used, andin
histext he
repeat-edly
referredto
thesepopulations
as“races”
ratherthan
as“species.”
Yet
thoserecent
authors,such
asF
ritts(1983,
1984)w
hohave
utilizedbinom
ialscite
Van
Denburgh
asthe
authorityfor
thisusage.
d)W
hileit has
beenm
aintainedby
some
that thefailure
ofGalapagos
tortoisesto
reproduceregularly
orabundantly
inzoos
isa
resultof
am
ixedstock
consistingof
different“species”
oftortoise
(seee.g.,
Bacon,
1980),this
argument
ignoresthe
fact thatsome
ofthezoo
populations(especially
atSanD
iego)w
ererelatively
largeand
composed
ofat most
two
orthreeisland
populations, sothat even
ifcross-matings
were
unproductive,there
would
beam
pleopportunity
form
atingsbetw
eenm
embers
ofa
singlepopulation.
e)R
yder(1978)
was
unableto
distinguishthe
popula
tionsofG
alápagostortoises
byelectrophoretic
separationof
serumproteins.
f)Itis
universallyadm
ittedthat the
Galapagos
tortoisesare
more
similar
toeach
otherthan
theyare
toany
otherspecies
within
(oroutside)
thesubgenus
Chelonoidis.
Insum
,theabove
pointsare
supportiveofthe
hypothesisthatthe
Galapagos
tortoisesare
allreferableto
asingle
species.R
ecognitionofthe
variousisland
orotherw
iseisolated
populationsas
subspeciesis
alsoa
pointw
orthyof
discus-sion.
The
populationsare
generallydistinguished
byadult
sizeand
theshape
ofthe
carapace(especially
theanterior
profile,the
width
atm
arginals2—
3,the
scuteor
pointat
which
thehighest point is
reached,and
theflaring
abovethe
hindlim
bs). How
ever, some
othercharacters
havealso
beenused, including
theam
ountofw
hiteor
yellowcoloration
onthe
faceand
chin;w
hetherthe
shellis
blackor
brown,
striatedorsm
ooth; orscutedetails
suchas
thetendency
inthe
Abingdon
formfor
theupper
edgeof
marginal
8to
benarrow
edoreven
excludedfrom
contact with
costal3, orfor
thepectorals
tofail to
meet on
them
idlineofthe
plastron,asin
some
specimens
fromC
hatham(V
anD
enburgh,19
14and
pers.obs.),
fromD
uncan(van
Lidth
deJeude,
1898;V
anD
enburgh,19
14), orin
thesingle
survivingindividual
fromA
bingdon(pers.
obs.).S
ome
ofthese
charactersare
betterthan
others.C
olorand
textureof
theshell
areclosely
involvedw
ithsuch
featuresas
age,sex, growth
rate, andthe
overallhumidity
ofthe
environment.
The
scutevariations
aretoo
unstableto
havem
aj ortaxonomic
value, andthey
appeartobe
primarily
asuperficial m
anifestationofchanges
inthe
overallshape
ofthe
shell(carapace
andplastron).
The
extentof
theunpigm
entedarea
ofthe
faceand
throat,w
hilegenerally
more
developedin
thesaddlebacked
forms, definitely
variesw
ithage.
Moreover,
adultsize
islikely
tobe
stronglyinfluenced
byfeeding
conditions.T
helargest
tortoisere
portedin
recent times
isa
captiveraised
specimen
collectedas
ahatchling
onIndefatigable
Islandas
recentlyas
1957(Fig.
23).Ithas
beenm
aintainedon
anabundant
dietat
the
..
•
PRITCHA
RD—
The
Galapagos
Tortoises
39
Life
Fellow
shipfacility
incentral
Florida,
andnow
,al
thoughshow
ingno
signsofobesity, w
eighsabout400
kg(G
.M
oss,pers
.C
Oflirn
.). Another
feature,the
degreeof bossing
of thecarapace
scutes,also
appearsto
berelated
tospeed
ofgrow
thor
toprotein
orcalcium
contentof
thediet,
andis
sometim
esvery
marked
incaptive-grow
nanim
als—
butreaches
itsgreatest
development
inw
ildadult
females
ofsuch
dwarfed
tortoisespecies
asthe
South
African
tenttortoise,
Psainnw
batestentorius
tentorius.N
evertheless, thedevelopm
ent oftheopen
shell frontinthe
saddlebackedform
sseem
sto
beundergenetic
control, asthe
youngtortoises
ofDuncan
andH
oodIslands, raised
fromhatching
atthe
Charles
Darw
inR
esearchS
tationunder
conditionsidentical
tothose
underw
hichyoung
tortoisesof
dome-shelled
racesare
raised,startto
showthe
characteristic
elevationof
thenuchal
areaw
ithina
fewyears
andat
acarapace
lengthof
onlyabout
30cm
(Fig.24).
On
theother
hand,those
fewindividuals
ofthe
characteristicallydom
e-shelled
populationsthatdisplay
some
degreeof
saddlingof
thecarapace
areinvariably,as
Fritts
(1984)pointed
out, “thelargest
andpossibly
oldesttortoises.”
Moreover,
Marlow
andP
atton( I98
1)presented
persuasivegenetic
evidencethatthe
varioussaddlebacked
populationshad
evolvedin
de
pendentlyfrom
dome-shelled
forms,
insharp
contrastto
Garm
an’5 unsupported
conclusionthatthe
forms
T.becki,T.ephippium
,and
T.hoodensis
were
basedupon
youngto
medium
-agespecim
ensof
T.elephantopus,
andthat
T.p
han
tasticaand
T.abingdoni
were
basedupon
oldand
very
oldsp
ecimen
sof
thesam
esp
ecies.T
hese
saddlebackedtortoises
sharea
number
ofunusual,
de
rivedm
orphologicalfeatures,
andin
theabsence
oftrenchant differences,
am
oderncladistm
ightalso
regardthem
asm
onophyleticallyderived
froma
dome-shelled
ancestor.D
iscussionof
the“subspecies
question”—
them
iddleground
between
consideringthe
variouspopulations
asfull
species,or
combining
allinto
asingle
monotypic
taxon—
requiresthatconsideration
betaken
ofthechanging
conceptofth
esubspecies
among
modern
evolutionarybiologists.
Inaddition,
comparisons
andcontrasts
needto
bem
adew
iththe
criteriathat
havebeen
usedby
biologistsevaluating
variabilityin
other cheloniansuperspecies,hybridizing
spe
cies,in
tergrad
ing
subsp
ecies,arch
ipelag
ictax
a,R
assenkreisen,and
thelike.
The
subspeciesconcept m
ayhave
beenfirstform
alizedin
aneditorial in
Novitates
Zoologicae
(Anonym
ous,1894),
inw
hichitw
asproposed
thattheterm
“variety”be
discardedin
favorof
thetw
oterm
s“aberration”
(forindividual
varia
tions)and
“subspecies”for
geographicalform
sw
hichcan-
notrank
asfull
species.Jordan
(1896)elaborated:
“Asub-
speciesis
alocalized
groupof
individualso
fa
species,the
mean
ofthe
charactersof
which
isdifferent
fromthe
mean
Fig
ure
25.Dead
youngadult
male
repatriatedtortoise
onD
uncanIsland,
July1986. T
hisindividual perished
afterthe
severepost-N
iflodrought
of1984—
85.
40CHELONIAN
RESEARCHM
ONOGRAPHS,N
umber
I—
1996
ofthe
charactersof
allother
localizedgroups,
andw
hichw
ill,under
favorablecircum
stances,fuse
togetherw
ithother
groups.”W
orkingdefinitions
of thistype
were
utilizedfor
much
ofthetw
entiethcentury, although
thew
ording“underfavor-
ablecircum
stances”clearly
neededrefinem
ent.T
hedev
eloping
consensusw
asthat
subspeciesgenerally
intergradedw
heretheirareas
ofdistributionjoined, butitwas
alsonoted
thatcaptive
conditionsm
ightprom
otehybridization
be-tw
eentaxa
thatshould
stillbe
consideredas
distinctspecies.
Indeed,observations
summ
arizedby
Karl
etal.
(1995)indicate
thatm
anyif
notm
ostof
thepossible
hybridsbetw
eendifferent
gen
eraof
cheloniidm
arineturtles
may
beproduced,
evenunder
wild
conditions,although
allare
veryrare.
Wilson
andB
rown
(1953)challenged
thesubspecies
concept, raisinga
stormofprotest from
the“establishm
ent”that rivalled
thecontrary
outburst when
subspeciesw
erefirst
proposedsixty
yearsearlier.
Wilson
andB
rown’s
argument
centeredupon
theobservation
thatso-called
subspecificcharacters
were
frequentlydiscordant
—that
is,that
selection
ofdifferent
characters(for
example,
absolutesize,
orcoloration,
orrelative
headlength)
canresult
incom
pletelydifferent concepts
ofsubspecific
variationw
ithina
species.T
heyconcluded
thatthe
subspeciesconcept
was
arbitrary,and
thushad
noplace
inscience.
This
proposal gatheredm
omentum
inthe
ensuingyears,
andthe
tempo
ofdescriptionsofnew
trinomials
faltered. But
Wilson
himself
subsequentlyrecognized
that heand
Brow
nhad
overstatedtheir
case(W
ilson,1994),
andthat,
inactual
fact, many
speciesindeed
showed
geographicvariation
with
genetictraits
thatchanged
ina
concordantrather
thandiscordant m
anner. Inrecent years,the
general approachhas
beento
utilizethe
criterionofintergradation
fortaxa
whose
naturalranges
come
intocontact.
From
suchcases
may
bedeveloped
aconcept ofthe
overalldegree
of morphological
divergencethatcharacterizes
subspeciespairs
asopposed
tospecies
pairs(thatm
aytypically
exhibit widespread
sym
pa
tryw
ithoutintergradation),
andto
applysuch
aconcept
todeterm
inationofthe
statusofallopatric
populations.N
ever-theless, this
policyis
notaunanim
ousone,and
indeedE
rnstand
Barbour
(1989)regularly
utilizeallopatry,
ratherthan
sympatry
without
intergradation,as
acriterion
fordistinct-
nessofchelonian
taxaatthe
specieslevel.B
ut thereis
atleastgeneral agreem
ent thatasubspecies, in
ordertobe
valid,hasto
begeographically
definable,and
cannotbe
merely
anassem
blageof
individualsfound
randomly
throughoutthe
rangeof
aspecies.
The
saddleb
acked
torto
isesof
Du
ncan
,H
ood,A
bingdon,and
Chatham
aredistinguishable
morp
ho
logically.V
anD
enburgh(1914)
translatedthese
differ-ences
intoquantitative
criteriaand
percentages,but in
gen
eral thetortoises
ofllo
od
andD
uncanare
smalland
havethe
highestpoint
ofthe
shellreached
atthe
extreme
front.T
he“saddle
horn”of
theH
oodtortoise
isnarrow
edand
well-
defined(Fig.
1 1);in
theD
uncanform
itis
broadand
ill-defined
(Fig.25). The
Abingdon
tortoise—
atleast theadult
males
—is
largerand
hasa
much
thickerneck
andthe
shellreaches
itshighest
pointnear
thecenter
ofvertebral
2.T
heextant
populationof
Chatham
tortoiseshas
notbeen
de
scribedin
anydetail.
butit
isrem
iniscentof
theH
oodtortoise
while
lessextrem
e—
notas
high,nor
asnarrow
ed,in
front. The
extinct formfrom
Charles
Island(see
Figs.44—45)
was
characterizedm
orphologicallyby
Broom
(1929).I
believeit
isjustified
toregard
thesefive
forms
asvalid
subspecies.F
ritts(1984)
alsom
entionsthe
Barrington
Is-landrace, know
nonly
frombony
fragments, as
asaddlebacked
race,but
thisis
theonly
mention
inthe
literatureof
them
orphologyof
thispopulation,
andno
detailsw
eregiven.
The
subspeciessituation
isless
clearon
thecentral
islandsof Indefatigable, Jam
es,and
Albem
arle.A
mong
theother reptiles
onthese
islands, thepopulations
oflavalizards
(Tropidurus
albem
arlensis),
geckoes(P
hyllodactylusgalapagoensis), andland
iguanas(C
onolophussubcristatus)
arenot
considereddistinct
evenat
thesubspecies
leveldespite
theexistence
ofwell-differentiated
full speciesw
ithinthese
generaelsew
herein
thearchipelago.
The
habitof
distinguishingthe
tortoisesof
thecentral
islandsas
eitherspecies
orsubspecies
may
belargely
am
atterof
habit,or
arelict from
thetim
ew
henthese
populationsw
eredescribed
asfull species
onthe
basisof individual type
specimens
with
littleor
nosupporting
hypodigmor
otherinform
ationon
populationvariability.
The
concept ofadifferent subspecies
oftortoiseon
eachofthe
fivevolcanoes
of Albem
arleIsland
iscom
plicatedby
thefact
thatonly
onV
olcanA
lcedoare
thetortoises
most
comm
onlyfound
nearthe
summ
itor
inthe
caldera.E
lse-w
herethey
arem
oretypical
ofthe
lowlands
andinterm
ediate
altitudes(although
theydo
reachthe
5540foot sum
mit of
Cerro
Azul),
andindeed
populationshave
beenreported
inthe
lowlands
equidistantfrom
them
ountainsof
Cerro
Azul
andS
ierraN
egra(near
Cabo
Rosa),
between
Sierra
Negra
andV
olcanA
lcedo( w
hereD
eS
olacollected
hisspecim
ensof
T.vandenhurghiatC
artagoB
ay,in
anarea
subsequentlycovered
bynew
lavaflow
s),and
between
Volcan
Alcedo
andV
olcanD
arwin
(atU
rvinaB
ay).T
heshell
colorand
textureof
thesetortoises,
while
subjectto
variation,m
aybe
environmentally
determined
(Fritts,
1983).as
may
bethe
adultsize.
The
Volcan
Darw
intortoise,
ina
ratherxeric
environment,
issm
allerthan
thoseofthe
threevolcanoes
tothe
south;A
.Rhodin
andIen
cou
ntered
acopulating
male
ofthis
populationw
itha
carapacelength
ofonly
77cm
.T
herem
aythus
heno
adequatebasis
for distinguishingthe
Albem
arlepopulations, exceptfor that
ofVolcan
Wolf, even
atthesubspecies
level. The
character-istic
ofthe
unusuallyeroded
shellsof
adulttortoises
onS
ierraN
egra,attributed
byV
anD
enburgh(1914)
tovolca
nicactivity,m
aym
orereasonably
beexplained
asa
formof
shell rot orfungusresulting
fromthe
unusuallydam
pco
ndi
tionsunder
which
thehighland
enclaveof
thispopulation
lived. Beck(1903)
notedthat, as
oneascended
Sierra
Negra,
thetortoises
increasinglyhad
erodedor
scarredshells,
consideredbyFritts
(1984)tobecharacteristic
oftheguentherim
orphotype.S
imilar
shellerosion
isfound
inw
ildor
intro-
PRITCHA
RD—
The
Galapagos
Tortoises
41
ducedpopulations
ofthe
mainland
speciesG
eochelonecarhonaria
invery
damp
regionssuch
asin
theD
arienof
Panam
aor
onO
ldP
rovidenceIsland
(pers.ohs.).
Itism
oreof
atestim
onyto
thetaxonom
icm
icroscopethat
hasbeen
directedat the
Galapagos
tortoisesfor
more
thana
centurythan
toany
realisticsubspecific
charactersthatsuch
environ-m
entallyderived
featuresw
ouldeven
beconsidered
tohave
nomenclatural
value.N
evertheless,recent
writings
bythose
most
familiar
with
southernA
lbemarle
tortoisesin
thefield
(e.g.,C
ayotand
Louis,
1994)do
followF
ritts(1984)
inrecognizing
aguentheri m
orphotypein
southernA
lbemarle, although
theyconsider
ittobe
farm
orerestricted
thandid
Van
Denburgh
(1914).V
anD
enhurghconsidered
theguentheri m
orphotype(or
race)to
hedistinguished
bythe
relativelyflat
carapace,especially
characteristicoflarge
adultmales, and
tobe
distributedgenerallyoverthe
southernslopesofS
ierraNegra(”V
ilaniilM
ountain”),although
itdid
notreach
asfar
eastas
Cabo
Rosa
(where
aform
indistinguishablefrom
microphyes
occurred),and
evenw
ithinthe
statedrange
ofguentheri
therew
asa
generousadm
ixtureof
thevicina
morphotype.
Picking
throughthe
ruinsof
thesedevastated
po
pu
lations
today, CayotandL
ouis(1994)reported
thattheguentherim
orphotypestill
existed,but
onlyin
thevicinity
ofC
incoC
erros(i.e.,
southeasternslopes
ofC
erroA
zul,not
SierraN
egra—
thesupposedly
exclusiverange
ofvicina).
Itis
difficultto
make
senseof
allof
this.O
fcourse,
tortoisesdo
move
veryextensively
when
theterrain
permits
—and,over
time
(atleastupto
thetim
eof the
1926lava
flow),
tortoisesm
ayhave
wandered
throughoutthe
vegetatedareas
ofsouthern
Albem
arle.In
thatthe
guentherim
orphotypecannot
beassociated
with
anyd
efinitiv
egeo
grap
hic
range,w
asrep
orted
byV
anD
enburghto
bem
icrosympatric
with
specimens
ofthe
adjacentrace
vicina,and
hascharacters
thatare
reallyonly
expressedin
adultm
ales,it
appearsbest
toregard
itas
asy
non
ym
of
vicina,pending
thepublication
ofD
NA
studiesby
Edw
ardL
ouis.T
henorthernm
ost populationoftortoises
onA
lbemarle
isa
differentm
atter.A
tleast
presentlyisolated
fromthe
Volcan
Darw
inpopulation
bylava
flows,
thespecim
ensof
thistortoise
seenin
thedry
lowlands
atB
anksB
ayand
atP
untaAlbem
arleinclude
asignificantproportion
ofstronglysaddlebacked
individuals(pers.ohs.). V
anD
enburgh(1914)did
notreport
dome-shelled
tortoiseson
thism
ountain,but
hisphotographs
dodem
onstrategreatvariation
inthe
degreeof
saddlebackingand
anteriornarrow
ingof
theadultm
ales,and
thesingle
adultfem
alecollected
(CA
S8
120)w
as,surprisingly,
them
oststrongly
saddlebackedtortoise
foundon
Volcan
Wolfeven
thoughithad
acarapace
lengthofonly
21.75inches. R
othschild(1901),in
describingT.becki from
northernA
lbemarle
(onthe
basisof
asingle
largem
ale)
—41’
!•‘•‘
;:‘r
Figure26.V
eryold
female
James
Islandtortoise, naturally
imprisoned
form
anyyears
ina
steep-sidedtuffcrater
southofJam
esB
ay.January
1970.T
heanim
alhas
sincebeen
liberatedin
theinterior
ofJam
esIsland.
42CHELONIAN
RESEARCHM
ONOGRAPHS,N
umber
1—
1996
comm
entedthatthis
newform
“was
ofthe
greatestinterest,because
itdem
onstratesthe
presence,not
onlyof
athird
speciesofG
iantLand
Tortoise
onA
lbemarle
Island,but alsoa
tortoiseof
atotally
differentkind
fromthe
othertw
ospecies.”
Yet
nowadays,
asI
haveseen
atboth
Bank’s
Bay
andP
untaA
lbemarle,
theV
olcanW
olfpopulation
includesan
apparent majority
oftortoisesthatare
somew
hatflattenedor
dome-shelled,
andnot
saddlebacked.P
ossiblya
long-iso-lated
populationof
saddlebackedtortoises,
adaptedto
thearid
andprecipitous
conditionsofV
olcanW
olf,was
invadedby
dome-shelled
tortoisesfrom
thesouth
duringa
periodw
henthe
lavaseparating
thesem
ountainshad
become
sufficientlyeroded
andvegetated
asto
bepassable.
Ifthis
postulateis
correct,thenthe
productionofm
orphologicallyinterm
ediatetortoises
onV
olcanW
olftoday
isfurther
evidencethat
thepopulations
aresubspecifically
relatedonly,and
donotshow
eithermutualinfertility
norbehavioralreproductive
isolatingm
echanisms.
The
persistenceof
ex
treme
forms
asw
ellasinterm
ediateones
suggestseitherthat
thepopulations
metcom
parativelyrecently
andhave
notyetbecom
efully
intergraded,or
thatrelatively
fewgenes
con-trol
thesaddling
ofthe
shell.O
necould
thenadd
theV
olcanW
olftortoiseto
thelist
ofvalid
subspeciesof
Galapagos
tortoiseson
thebasis
ofcertain
phenotypesthat
aredefinable
anddifferent
fromother
saddlebackedtortoises
inbasically
thew
aysdescribed
byR
othschildand
Van
Denburgh,
althoughthe
populationis
sovariable
todaythat
arigorous
diagnosisof
thesu
bsp
ecies
would
probablybe
impossible.M
oreover, itwould
stillbe
appropriateto
synonymize
theother
foursubspecies
reportedfrom
Albem
arle;they
simply
cannotbe
distin
guishedw
ithany
certainty,andthe
mostobvious
characters(size,
texture)are
environmental
responses.T
hepopulations
ofJames
andIndefatigable
arenotvery
differentfromthose
ofthesouthern
volcanoesof A
lbemarle,
butlargem
ales,atleast,tendto
bedistinctive
inm
inorways,
andcertainly,being
onseparate
islands,theym
aybe
consid
eredgenetically
isolated.T
heJam
esIsland
tortoiseis
alarge,
semi-saddlebacked
orinterm
ediateform
(Figs.15,
26),diagnosed
byV
anD
enburghas
havingthe
carapacew
idthat
marginals
2—3
equalto
48—58%
ofthe
straightcarapace
length,and
theheightatthe
nuchalnotch
42—45%
ofthe
straightlength.T
heyounger
adultmales
onIn
defati
gablehave
anextrem
elyrounded,
highlydom
edcarapace,
with
theheightatthe
nuchalnotch
30to
40%ofthe
straightlength,and
thew
idthat m
arginals2—
3from
53to
71%
ofthestraight
length.V
erylarge,
oldm
alesfrom
Indefatigable(none
ofwhich
were
collectedby
Van
Denburgh)
may
tendtow
ardselongation
ofthe
shell,or
posteriorincrease
inw
idth,producing
aless
prominently
domed
form(Figs.
I6and
27,also
seephotos
onp.
6in
theeditorialprologue
andon
theback
cover).Butthe
James
andIndefatigable
tortoisesm
aybe
consideredas
validsubspecies
onthe
groundsof
geographicisolation
andat
leastm
arginalm
orphologicaldifference.
3)U
nderthe
ICZ
NR
ulesof
Nom
enclature,the
oldest
availablenam
ethat
isnot
anom
ennudum
,w
hoseholotype
isclearly
arepresentative
ofthespecies
inquestion,and
thatw
aspublished
ina
work
exclusivelyem
ployingthe
binomial
system,is
theone
thatshouldapply.F
ormerly,long-forgot-
tensenior
synonyms
couldbe
dismissed
asnom
inaoblita,
butthe
latestedition
ofthe
Rules
(ICZ
N,
1985)requires
formal
petitionbe
made
beforesuch
names
canbe
consid
eredunavailable.
Testudo
rotundaL
atreille,1801
,althoughthoughtto
applyto
aG
alapagostortoise
byB
our(1980),has
subsequentlybeen
shown
bythe
same
authorto
applyto
aM
ascarenespecies
(R.
Bour,
inlitt.),
sothe
firstnam
eto
beapplied
toa
Galapagos
tortoisein
thescientific
literatureis
Testudo
californianaQ
uoyand
Gaim
ard,1824a.
This
description,consisting
ofa
briefL
atindiagnosis
(“Testudo
totocorpore
nigro;testagibba;scutellisdorsalibusprioriposteriorque
elevatis; loricaecunctis
margine
striatis;lateribus
subcarinatis”)w
asfollow
edby
aseries
ofdim
ensions,using
theancient
systemofpieds
orpedes,poucesor
polles, andlignes
orlines. This
isa
duodecimalsystem
,with
1pouce
equivalentto
27m
m.
The
paperannouncing
thisnew
speciesw
asread
atthem
eetingofthe
Société
d’Histoire
Naturelle
deP
arison
7N
ovember
1823(1.
Lescure,
pers.com
m.),
byM
.G
aimard.
The
proceedingsof
thism
eetingw
erepublished
inthe
secondsection
oftheB
ulletinU
niverselledes
Sciences
etdel’Industrie,
partof
theB
ulletindes
Sciences
naturelleset
Géologie.T
heprecise
dateofthis
publicationis
unclear,butIam
advisedbyL.Jescure
(inlitt.,
12M
arch1977)thatitw
asvery
probablybefore
24A
pril1824,and
certainlybefore
I8S
eptember
1824.T
hedate
isim
portant,in
that,later
inI824,
anew
descriptionofthe
same
tortoisew
aspublished,also
byQ
uoyand
Gaim
ard(1824b),
within
thew
orkV
oyageA
utourde
Monde
executesur
l’Uranie
etIa
Physicienne
pendantles
années181
7—1820
byM
L.
deF
reycinet.T
hisw
aspub-
lishedin
fascicles;F
ascicle4,
containingthe
descriptionof
thenew
tortoise,thistim
ecalled
Testudo
nigra,appearedon
18S
eptember
1824,w
iththe
plate(P1.
40)published
later
( 18D
ecember).T
henew
description(differing
byonly
afew
words
fromthe
earlierdescription
ofT.
californiana)w
assupplem
entedby
apage
anda
halfofFrench
textdescribingthe
specimen
infairdetail,and
givingthe
supposedorigin
asC
alifornia.It
iscurious
that,after
allof
them
ariners‘
andbucca
neers’accounts
ofgiant
tortoisesin
theseventeenth
andeighteenth
centuries,it
seems
tohave
beenstill
un
recog
nizedin
thescientific
world
thatgianttortoiseseven
existed—
inthe
paperim
mediately
following
thedescription
ofT.
californiana,Fitzinger(1
824)gives
anaccountofhis
visittothe
Menagerie
inV
ienna,w
herehe
identifieda
Brazilian
tortoisethat
hecalled
Testudo
schweigeri
(presumably
theform
nowknow
nas
Geochelone
denticulata,a
largebutnot
giantspecies)
asthe
largestknow
nland
tortoise(“la
plusgrande
tortuede
terreque
l’onconnaisse”).
Schw
eigger’sdescription
ofthis
speciesas
T.gigantea
isdiscussed
byP
ritchard(1986).
But
thenam
eT.
schweigeri
(“corrected”by
Boulenger
[1889]to
Testudo
schweiggeri),
isnorm
ally
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises43
attributed(e.g.,
byB
oulenger[1889]
andW
ermuth
andM
ertens[1961
1) toa
laterw
riterthan
Fitzinger
(1824),i.e.,
Gray
(1831),and
basedupon
arather
smallshell
sopuzzling
inappearance
thatnoone
hasbeen
ableto
identifyit.(Itm
aybebasedupon
aspecim
enofG
opheruspolyphemus
showing
numerous
abnormalities
ofboth
formand
coloration).B
utsom
etimes
thetask
ofm
akingprecise
interpretationsfrom
decidelycasual
writings
bynineteenth
centurynaturalists,
who
alltoo
frequentlytook
nonote
ofeach
others’w
ork.leads
oneto
thetype
ofconclusion
expressedby
Vanzolini
( 1977),w
hocom
mented
that“G
ray’scarelessness
inciting
scientificandpersonalnam
es,hisdisregard
forotherpeople’sopinions
andprevioususage,and
wretchedproofreading,all
conspiretogether
tom
akethis
pretentiouslittle
work
[i.e.,G
ray,1825]
veryunpleasant.”
Testudo
califrirnianaQ
uoyand
Gaim
ard,I824a,
was
notutilizedby
subsequentreviewers.
Under
earliereditions
oftheR
ulesofN
omenclature,this
name
couldbe
dismissed,
eitherbecause
itw
asa
nomen
obliturnor
becausethe
descriptionw
asan
unrigorousw
rittensum
mary
ofa
verbalpresentation.
Under
the1985
Rules,how
ever,theseco
nsid
erationsno
longerapply,
andthe
name
will
haveto
beused
unlesssuccesful
petitionis
made
tothe
ICZ
Nto
haveit
repressed.O
nthe
otherhand,
evenunder
theold
Rules,
Testudo
nigracould
notbedism
issedas
anoinen
oblitum.
Anam
eused
byC
harlesD
arwin
(1845)
inhis
Journalof
Researches
ishardly
aforgotten
one,and
Garm
an( I 917)
usedthis
name
forthe
tortoiseofC
harlesIsland.
Ttwas
alsocited
byC
uvier(1829),
Dum
ériland
Bibron
(1835),W
iegmann
(1835),S
trauch(1862,
1865,1890),
andB
oulenger(1889),
andw
asdiscussed
byV
anD
enburgh(1914)andby
Rothschild
andG
Unther(inR
othschild,1902c),
althoughthese
lastauthors,
convincedthatthe
islandp
opu
lationsshould
allbereferred
todifferent
species,dism
issedthe
name
asunusable
becausethe
holotypew
astoo
smallfor
theisland
oforigin
tobe
determined.
Nevertheless,
thisproblem
isonly
operativeas
longas
eachpopulation
isconsidered
adistinct
species.W
henM
ertensand
Werm
uth(1955)
subsumed
alltheG
alapagostortoises
within
asingle
species,theyshould
haveselected
nigraas
thevalid
epithetw
ithpriority,
anddealt
separatelyw
iththe
issueof
which
islandpopulation
shouldbe
consideredthe
nominate
sub-species.
The
speciesnam
eelephantopus
ofHarlan
(1827)has,of
course,been
usedin
avast
number
ofpapers
duringthe
lastthirty
years,and
acase
couldbe
made
fora
petitionto
theIC
ZN
toretain
it. How
ever,Idonotrecom
mend
thisstep, in
partbecause
useof
elephantopusfor
allthe
Galapagos
tortoisesis
relativelynew
(since1955),
andthus
lacksthe
statusof
anom
enveneraturn
asdefined
byS
mith
andS
mith
(1980),and
inpart
becausethe
islandof
originof
theholotype
isconjectural.
Van
Denburgh’s
assumptions
lead-ing
tohis
guessthat
thism
ightbe
theC
harlesIsland
formw
ereclearly
flawed
(Broom
,I929).M
oreover,theholotype,
Figure
27.A
dultm
aletortoise
inlush
vegetationnear
theC
aseta,southeastern
highlandsof
IndefatigableIsland.
July1986.
44C
HE
LO
NIA
NR
ES
EA
RC
HM
ON
OG
RA
PH
S,
Num
ber1
—1996
beinglost
(theappalling
confusionresulting
fromefforts
tofind
oridentify
thisspecim
enbeing
discussedby
Garm
an,1917),
isnot
availablefor
future,m
oresophisticated
mor
phological,genetic,
orbiochem
icalanalysis
thatm
ighthave
clinchedthe
identification.I
thereforeconclude
that thecorrect
specificnam
efor
theG
alapagostortoises
isG
eochelone(C
helonoidis)n
igra
(Quoy
andG
aimard,
I824b).A
tthe
subspecieslevel,
furthernom
enclaturalpro
blem
sexist.
Ifnigra
isthe
applicableepithet for
thespecies,
which
raceis
thenom
inotypicalsubspecies,
Geochelone
nigranigra?
Until
suchtim
eas
Captain
Meek’s
logbooks
arefound
andstudied
andthe
actualorigin
ofthe
typespecim
endeterm
ined(an
improbable
development),
oneoption
would
beto
make
anarbitrary
determination,
vali
datedin
ways
dictatedby
acarefulconsiderationofthe
Rules
ofNom
enclature. How
ever,perhapsthe
least painfuloptionw
ouldbe
toturn
adeliberate
blindeye
tothe
weaknesses
inG
arman’
S arguments
thatthisform
was
fromC
harlesIsland,
andto
acceptthatdesignation
onthe
groundsthatit
at leastcannot readily
bedisproved.
The
procedurew
ouldhave
theadvantage
ofnot
affectingthe
nomenclature
ofany
extantsubspecies.
Furtherm
ore, onlythree
ofthesubspecies
ofGalapagos
tortoises(hoodensis, phantastica,and
darwini)
seemto
haveunclouded
taxonomic
status,w
ithdem
onstrablem
orp
ho
logicaldistinctness,
known
islandsof
originof
thetype
material,
andno
known
seniorsynonym
s(V
anD
enburgh,1907),
althoughT.
phantasticaV
anD
enburgh,1914
doesrepresent
am
inorcorrection
ofthe
originalnam
eT.
ph
antasticu
sV
anD
enburgh,1907.
The
holotypesof
thesethree
forms
are,respectively,
CA
S8121,
CA
S8101,
andC
AS
8108.P
roblems
with
theother
names
areas
follows:
i)W
hileT.
chathainensisw
asbased
byV
anD
enburgh(1907)
upona
holotype(C
AS
8127,Fig.
17)definitely
collectedon
Chatham
Island,theform
describedfrom
there(based
ona
singleentire
specimen
anda
fewshells
andbones
oftortoisesthat had
diednaturally
orbeenkilled
bysettlers)
was
aflat-shelled
typefound
inthe
relativelyhum
idareas
southwest
ofthe
freshlava
flows
ofcentral
Chatham
.T
hispopulatio
nw
asalm
ostextinct
in1906,
when
Van
Denburgh’s
specimen
was
collected,and
itnow
appearsto
becom
pletelyso.
The
photographof
Chatham
Islandtortoises
inS
ulloway
(1984),w
iththe
captionindicating
thatthey
were
locatedin
northwestern
Chatham
Island,is
misleading;
thecaption
incorporatesa
typographicalerror
andshould
readn
orth
easternC
hathamIsland
(F.S
ulloway,
pers.com
m.).
The
extantpopulation
oftortoises
onC
hatham,
men-
tionedbyD
orst(1965), Castro
(1970), andP
ritchard(1979),and
numbering
several hundredindividuals, is
ratherm
ark-edly
saddlebacked(at least in
theadult m
ales),and
occursin
thedry
lowlands
northeastof
thebarren
volcaniccenter
ofC
hatham.Itthus
seems
thattheliving
populationis
nottruechatham
ensis,but
ratheran
undescribedsubspecies.
ii)The
typespecim
enofT
.becki(No.2
intheR
othschild
collection,an
adultoriginally
describedas
40.75inches
=
103.0cm
inlength[R
othschild,1901];clarifiedbyR
othschild[1915a]
asbeing
curvedcurved
length,and
with
astraight
carapacelength
of3
1.25inches
=79.4
cm),
was
reportedlynot
fromV
olcanW
olf,to
which
populationthis
name
iscustom
arilyapplied,but from
Cape
Berkeley
(Volcan
Ecu
ador),
atthe
northwestern
tipof
Albem
arle,and
partof
aseparate
volcano(now
bisectedby
downfaulting).
Itis
unclearif
thisis
simply
anerror
oflabelling,
orif
thespecim
enreally
came
fromC
apeB
erkeley.Itcould
alsobe
acase
ofgeographic
confusion.T
heoriginal
descriptionreferred
to“C
apeB
erkeley,northern
pointof
Albem
arleisland”
(Rothschild,
1901),although
thenorthern
pointof
Albem
arleIslandis
infactP
untaAlbem
arle, aknown
habitatfor
saddlebacktortoises.
Cape
Berkeley
isone
ofthe
least-know
nareas
ofthearchipelago
today, andw
hileno
tortoiseshave
beenfound
therein
recentyears,
thepossibility
thatsom
eare
therecannot be
dismissed.C
onceivablythere
is(or
was)
apopulation
ofsaddlebacks
onC
apeB
erkeleythat
intermittently
hadaccess
toV
olcanW
olf,and
thetortoise
populationon
thelatter
mountain
may
representan
intergradientone
between
saddlebackedand
domed
tor
toises.B
utthis
isspeculation.
iii)T
heform
Tw
allaceiw
asdescribed
byR
othschild
Figure
28.Illustrationfrom
Snow(1964)
ofthem
alesaddlebacked
tortoisefound
bythe
Angerm
eyerbrothers
innorthw
esternIn
de
fatigableIslandin
1962. Photo
byD
.W.Snow
.The
original captionread:
“The
oldm
alesaddle-backed
tortoiseon
Indefatigablefeed-
ingon
acactus
padfrom
theopuntia
treebehind
him.
The
dev
astation
ofthe
undergrowth
ism
ainlythe
work
ofgoats.”
PRITCHARD—
The
Galapagos
Tortoises
45
Figure
29.Juveniletortoise
foundby
theauthor
nearSaddleback
Hill(C
erroM
ontura),Indefatigablelsland,inMay
1972. The
small
sizeof
theanim
al(ca.
35cm
CL)
issuggestive
ofsuccessful
reproductionofthis
colonysince
itsdiscovery
in1962.
(1902d)on
thebasis
ofa
singlesm
ooth-scutedadult
male
shell8
1.9cm
instraight
length,N
o.42
inthe
Rothschild
collection,w
hoseorigin
was
unknown.
Originally
labelledw
iththe
catchallgianttortoise
name
T.indica,itcam
efrom
theprivate
collectionof
oneM
r.B
ullock.It
was
thenacquired
bythe
Wallace
collection,housed
inD
istington,C
umberland,andpassed
intothe
Rothschild
collectionupon
thedispersalofW
allace’scollection.R
othschildspeculated
thatthe
specimen
was
fromC
hathamIsland,
onthe
thinargum
entthat,between
1800and
1835,mostofthe
tortoisescollected
inthe
Galapagos
came
fromJam
esand
Chatham
Islands,and
thisspecim
en,not
beingas
roundedas
theJam
estortoises
were
known
tobe,
presumably
came
fromC
hatham.
Van
Denburgh
(1914)reported
onthe
collectionof
asinglelive
male
tortoiseon
JervisIsland
byB
eckand
Slevin,
which
hefound
tobe
generallysim
ilarto
thetype
ofT.
wallacei.H
ethus
felttentativelyjustifiedin
usingthis
name
forthe
tortoisesof
JervisIsland,
eventhough
itw
aspro-
foundlydoubtful
thatJervis
hadever
supporteda
nativetortoise
population.V
anD
enburghdid
concedethat
therew
asa
rumorin
theislands
thatBaurhad
introducedtortoises
toJervis,and
Rothschild
(1915a)
consideredthe
differencesbetw
eenthe
specimens
tobe
ratherprofound.
Inview
ofthese
circumstances,the
name
T.wallacei
Rothschild
1902should
beconsidered
asynonym
ofT.
nigraQ
uoyand
Gaim
ard,1824b.
Nevertheless,
theconclusion
ofP
ritchard(1979)
andothers
thatJervisIsland
neverhad
anative
tortoisepopula
tionm
ustbere-exam
inedin
thelightofreported
discoveriesof
fossilor
caverem
ainsof
bothtortoises
andland
iguanason
thisisland,
thatled
Steadm
anand
Zousm
er(1988)
tostate
thatJervis
oncesupported
nativepopula
tionsof
tortoisesand
landiguanas
thatm
ighthave
diedout
asa
resultof
predationor
increasedcom
petitionfor
foodsources.
Their
speculationis
questionable,in
thatthe
ratsand
(formerly)
goatson
Jervisw
ereboth
ofrecent
introduction,and
couldhave
playedno
partin
the
Fig
ure
30.R
easonablyfresh
tortoisedroppings
foundnear
SaddlebackH
illby
A.G
.J.R
hodinand
theauthor
inSeptem
ber1982.
extermination
ofterrestrial
reptiles.P
erhapsthe
tortoisebones
foundderived
froman
unsuccessfulintroduced
population;G
.B
aur,for
one,is
known
tohave
liberatedtortoises
onJervis.
iv)T
henam
esp
orteri
andnigrita
havebeen
usedvirtually
interchangeablyfor
thetortoises
ofIndefatigable
Island.T
estudonigrita
Dum
ériland
Bibron,
1835,is
theolder
name,
butis
basedupon
avery
youngand
a55.9
cmsubadultspecim
en,theform
erinthe
MN
HN
Pcollection,the
latter(on
which
thedescription
was
primarily
based,and
thusto
beconsidered
thelectotype)
inthe
Hunterian
Mu-
seum(R
oyalC
ollegeof
Surgeons).
GU
nther(1877)
laterassociated
alarge
(104.1cm
),dom
edcarapace
inthe
Royal
College
ofS
urgeonsC
ollection(now
inthe
BM
NH
)w
iththis
name,
andG
arman
(1917)figured
thisspecim
enin
hisP
late9.
Van
Denburgh
(1914)w
rotethat
“we
seemju
stifled...
insaying
thatG
unther’sT
estudonigrita
agreesw
iththe
Indefatigabletortoise,”
butstill
preferredthe
useof
thenam
eT.porteri R
othschild,1903,w
hoseholotype,N
o.44in
theR
othschildM
useum,w
asa
92.7cm
adultmale
definitelycollected
onIndefatigable
Island(by
R.B
eck).Sinceporteri
hasachieved
ratherw
idespreaduse
(e.g.,byV
anD
enburgh,1914;
Pritchard,
1967;B
ailey,1970;
Groom
bridge,1982;
andF
ritts,1983),its
continueduse
isjustified,especially
inview
ofits
more
satisfactorytype
material.
While
thelarge
Royal
College
ofS
urgeonsspecim
en,identified
asnigrita
byG
Unther
(1877),is
almost
certainlyan
Indefatigabletortoise
ongrounds
ofmorphology,
itm
ustberem
embered
thatthis
individualw
asnot
partof
thetype
series,and
itw
ouldthus
beappropriate
tom
akepetition
tothe
ICZ
Nto
suppressthe
name
nigrita.T
hereis
anotherisolated
andvery
smallpopulation
oftortoises
onIndefatigable,described
byS
now(1964)
andby
Pritchard
(1979)(Figs.
28—30).A
dulttortoisesofthis
po
pu
lationare
stronglysaddlebacked;
theylive
inthe
aridnorth-
western
cornerof
theisland,
separatedby
aconsiderable
distancefrom
thepopulation
ofporteri
insouthw
esternIndefatigable.
This
populationhas
notbeen
named,
and
46C
HELO
NIA
NR
ESEAR
CH
MO
NO
GR
APH
S,N
umber
1—
1996
thereare
nospecim
ensin
museum
collections.v)T
henam
eT. abingdoni G
unther,1877, is
customarily
usedfor
thetortoises
ofA
bingdonIsland,
andindeed
this
name
isbasedupon
anadequate
seriesofspecim
ens(B
MN
H76.6.21.38—
40,76.6.21
.44)of
definiteA
bingdonIsland
orig
in(collected
byJ.
Cookson
in1875)
(Fig.3
1,
seealso
frontispiecepainting). H
owever, as
Ihavepreviously
argued(P
ritchard,1984),
theholotype
ofT.
ephippiumG
Unther,
1875,custom
arilyallocated
toD
uncanIsland
byV
anD
enburgh(1914)
andsubsequent
authors,agrees
inboth
mo
rph
olo
gy
and
circum
stantial
details
of
itsco
llection
with
thetortoises
ofA
bingdon(B
aur,1889),
andis
theolder
name.
The
holotypeofephippium
(Figs.21—
22), althougha
relativelyyoung
animal, has
acarapace
lengthof83
.8cm
about9
cmlonger
thanthe
largestof
86D
uncantortoises
collectedby
theAcadem
yexpedition(V
anD
enburgh,1914),
andabout7
cmlonger
thanthe
largest of26specim
ensin
theR
othschildcollection
(Rothschild,
I915a),
butabsolutely
typicalof
male
Abingdon
tortoises(R
othschild,19
15a).M
oreover, inlateral profile, the
specimen, stillin
goodshape
inE
dinburgh,show
sthe
Abingdon
tortoisecharacteristic
(con
trasting
with
the
Duncan
torto
isem
orp
ho
log
y)
of
the
highestpoint
ofthe
carapacereached
atvertebral
2rather
thanat
thevery
frontof
thecarapace.
The
circumstances
ofthecollection
ofthespecim
enare
alsopersu
asive.
Bau
r(1889),
referringto
thediary
ofC
ap-
tamB
asilH
all( I840),a
mariner
who
visitedthe
Galapagos
inI822,
determined
that Abingdon
was
theonly
islandthat
Hall had
actuallyvisited
inthe
archipelago,and
thathehad
takensom
etortoises
onboard,
althoughall
perishedin
thecold
weather offC
apeH
orn. One
ofthesew
aspreserved, and
“itm
aynow
beseen
inthe
Museum
ofthe
College
atE
dinburgh.”T
hiscollection(now
theE
dinburghM
useumof
Science
andA
rts)did
not givecatalog
numbers
tospecim
ensin
theearly
nineteenthcentury, but the
acquisitionlog
doesinclude
anentry
indicating“L
argeturtle
fromS
outhS
ea—
Captain
Basil
Hall,”
andit
onlyhas
asingle
Galapagos
tortoise—
thetype
ofT.
ephippium.
Thus,
T.abingdoniG
unther,1877, is
ajuniorsynonym
ofT.
ephippiurnG
Unther,
1875,and
theD
uncanIsland
tortoisew
illrequire
analternative
name;
thediscovery
thata
type
specim
enhas
been
misid
entified
isa
more
serious
problemthan
them
erediscovery
ofan
obscuresenior
syn
on
ym
,w
hich
canoften
bed
isposed
of
byp
etition
tothe
ICZ
N.
Itis
appropriateto
reviveG
arman’s
Testudo
duncanensisfor
thispurpose.
This
name
appearsonly
once,onp.
269, ofGarm
an(1917).
where
it isoffered
inbinom
ialfo
rm,
althoug
hd
escribed
asonly
a“v
ariety”
of
Testudo
elephantopus.L
aterin
thispaper
(pp.290—
296),G
arman
offers
adetailed
descrip
tion
of
hisco
mp
osite
taxon
T.elephantopus,
andon
pp.292—
293, thereappears
adetailed
descriptionof
a25
inchm
alespecim
en,M
CZ
11068,w
ith
:i
Fig
ure
31.Mounted
specimen
ofanA
bingdonIsland
tortoise,a
veryold
adult male
collectedby
J. Cookson
in1875; itdied
offCape
Horn
onthejourney
toE
ngland.S
pecimen
inthe
British
Museum
(Natural H
istory). Seealso
thefrontispiece
paintingw
hichappears
tobe
basedon
thisspecim
en.
PRITCHARD—
The
Galapagos
Tortoises
47
accuratecollection
data—
itwas
collectedon
Duncan
Islandby
G. B
aur.The
accountalsodescribes
aduitfemale
specimens
fromthe
same
island.Thus,M
CZ
11068should
beconsidered
thelectotype
oftheD
uncanIsland
tortoise,Geochelone
nigraduncanensis
(Garm
an,1917),
andis
sodesignated.
On
theother
hand,substitution
ofthe
name
ephippiumfor
abingdoniw
ouldcause
extensiveconfusion,
theform
erhaving
asubstantial
recentliterature
inall
ofw
hichit
isconsidered
torefer to
theD
uncantortoise, and
itisap
pro
pri
ateto
petitionthe
ICZ
Nto
establishthe
validityofabingdoni
byP
lenaryD
ecreedespite
theexistence
ofa
seniorsu
bjec
tivesynonym
.vi)
While
thispointbecom
esm
ootifthetortoises
ofthefour
southernvolcanoes
ofA
lbemarle
aresynonym
ized,G
arman
observedthat
thenam
eT.
microphyes
GU
nther,1875,custom
arilyused
tothis
dayforthe
tortoisesofT
agusC
oveand
Volcan
Darw
in, was
basedupon
asingle, probably
abnormal fem
ale(but ventrally
concave)specim
en, 57cm
inlength,
ofunknow
nprovenance.
Ithad
beenpurchased
bythe
British
Museum
fromthe
Museum
Com
mittee
ofthe
Royal
Institutionof
Liverpool.
GU
nther’ssubsequent
allocation
toT.
microphyes
ofa
seriesof
specimens
ofknow
nV
olcanD
arwin
origin(from
“asm
allelevated
plateaucoy-
eredw
ithstunted
bushand
high,very
coarsegrass”
aboutfourm
ilesinland
fromT
agusC
ove,where
tortoisesm
aystill
befound
duringm
oisttim
esof
theyear)
didnotexem
pt theholotype
fromchallenge,
andin
viewof
this,G
arman
(191
7)proposed
thealternative
name
T.m
acrophyesfor
theT
agusC
ovetortoises,
althoughfor
mysterious
rea
sonshe
con
tinu
edto
useT.
microphyes
forother
Albem
arlepopulations.
Another
detailconcerning
thenom
enclatureof
oneof
thesouthern
Albem
arletortoises
pertainsto
theform
de
scribedby
Baur
(1889)
asT. guntheri,w
rittenthus, i.e.,w
ithan
umlaut.
Many
authorsfrom
thento
nowhave
renderedthis
name
inidenticalfashion—
theseinclude
Van
Denburgh
(1914), Rothschild
(1915a), Garm
an(1917), B
eebe(1925),
Eibl-E
ibesfeldt(1960),
Werm
uthand
Mertens
(1961),H
endrickson(1964),P
ritchard(1967),IU
CN
(1968), Black
(1973),M
acFarland
etal.
(1974a,1974b),
andde
Vries
(1984).C
rumly
(1984)
bothutilized
andquoted
theepithet
without
theum
laut;how
ever,a
corrigenduminserted
intothis
paperindicated
thatguntheri
shouldbe
substitutedfor
guntherithroughout.
Despite
theabundance
ofauthorsw
housed
thegiintheri
form,
the1961,
1964,and
1985editions
ofthe
Rules
ofN
omenclature
(asw
ellas
thedraft
versionof
thefo
rthco
ming
edition)are
quiteexplicit;
inC
hapterV
II,Article
32(d)
(i)(2), they
specifythata
name
derivedfrom
theG
erman
andincluding
a“U
”shouldbe
emended
sothat“ue’is
substitutedfor
“u.”O
nthe
otherhand,
the1977
Rules
includedad
di
tionalw
ording,specifically:
“.
.exceptw
henthe
name
was
firstcorrected
bydeletion
ofthe
mark
concerned,in
which
caseit
cannotbe
correctedfurther.”
Authors
who
usedguentheri
includeH
onegger(1972,
1980),IU
CN
(1975,1979),
Co
rleyS
mith
(1977),G
roombridge
(1982),F
ritts(1983,
1984),O
bst(1985),
and
Werm
uthandM
ertens(1977).T
heselast authorsconsidered
thistaxon
invalid,listing
itin
thesynonym
yof
T.e.
elephantopus,m
isquotingthe
originalauthor
(Baur,
1889)by
usingthe
emended
form(item
sin
synonymy
shouldbe
citedexactly
asoriginally
written).H
owever,the
earliest useof this
emended
formappears
tohave
beenby
me
(Pritchard,
197la,
1971b) intw
opapers
inw
hich,tom
yem
barrassment,
Iused
firstthe
unaccented(guntheri)
formand
thenthe
emended
form(guentheri). Ifoundto
my
furtherchagrinthat
inm
y1967
book,the
accentedform
(guntheri)had
beenused
inthe
text,w
hilein
theappendix
tothe
same
volume
ithad
appearedin
unaccentedform
(guntheri). To
date,Bailey
(1970),
Iverson(1985)
andC
ayotand
Louis
(1995)are
among
thefew
who
havefollow
edm
ein
theusage
ofthe
unaccentedform
.F
urther problems
areencountered
with
thenam
eofthe
adjacentpopulation,
T.vandenburghi
De
Sola,1930.
This
name,
anom
ennudum
,can
onlybe
validatedif
atype
specimen
isdeclared
andif
thefeatures
byw
hichthe
formdiffers
fromother
Galapagos
tortoisesare
clarified.M
ore-over,
thenam
ecan
onlybe
legallyapplied
tothe
Volcan
Alcedo
tortoiseif
itcan
beshow
nnot
onlythat
thepopula
tionfrom
which
De
Sola’ sspecim
ensw
ereobtained
(alow
landpopulation
nearC
artagoB
ay)w
asidentical
tothe
Alcedo
population,but
alsothat
itdiffered
fromother
Albem
arlepopulations
with
oldernam
es.T
hisdem
onstration
will
bedifficult,
inthat
theC
artagoB
aypopulation
appearsto
havebeen
eliminated
fromthe
wild
byvolcanic
activity,butperhapsitw
ouldbe
possibleto
tracesom
eof the
specimens
De
Sola
senttozoologicalparks, ifthey
happenedto
endup
inm
useums
upontheir
demise.
How
ever,if
theysurvived
forany
lengthof
time
incaptivity,
theyare
likelyto
havedeveloped
some
minor
abnormalities
ofcaptive
growth
thatcould
mask
anysubtle
shellfeatures
uniqueto
thispopulation. Interestingly, eventhoughtheV
olcanAlcedo
tortoisepopulation
isthe
most
abundant inthe
archipelagotoday,
ithas
neverbeen
morphologically
characterizedin
theyears
sinceD
eS
ola’s1930
publication,and
Van
Denburgh’s
(1914)originaldesignation
oftheterm
Testudo
sp.forthispopulation
reliedentirely
upona
singlespecim
en(C
AS
8141)
thatw
asnot
onlytoo
small
(carapacelength
26.75inches)
todem
onstratepotential
subspecificch
aracters,
butw
asalso
atypicallyhigh
domed.
The
“seventyold
skeletons”observed
byR
olloB
eckon
Volcan
Alcedo
when
thelive
juvenilew
asfound
would
havebeen
useful,but
unfortunatelyw
erenot
collected.F
urtherquestions
arisein
connectionw
iththe
selectionof
theappropriate
trinomial
forthe
singlesubspecies
oftortoise
that,I
haveargued,
inhabitsthe
foursouthern
volcanoesofA
lbemarle.
Of
theavailable
names
(T.vicina,
T.elephantopus, T.microphyes, T.m
acrophves, T.guentheri,and
T.vandenburghi), itshouldbenotedthat the
originofthe
typespecim
ensof
thefirst
threeform
sand
ofT.
guentheriare
unknown;
T.m
acrophyeshas
neverbeen
usedsince
itsoriginal
proposal;and
T.vandenburghi
remains
anom
ennudum
.H
owever,
theholotype
ofT.
vicinaG
Unther,
I875(an
adultm
aleskeleton,
BM
NH
74.7.15.1,obtained
from
48CH
ELON
IAN
RESEARCH
MO
NO
GRA
PHS,
Num
ber1
—1996
Figure
32.V
eryold
adultm
aletortoise
onV
olcanA
lcedo,A
lbemarle
Island,w
ithatypically
elongate,saddlebacked
carapace.D
esquamating
carapacialtexture
isprobably
aresultof protracted
exposureto
extremely
moist conditions. M
arch1975.
Professor H
uxley), was
subjectedto
detailedm
orphologicalanalysis
byV
anD
enburgh(19
14)and
determined
byhim
tobe
allocatableto
theIguana
Cove
(southwestern
Albem
arle)population. T
estudom
icrophyesG
Unther,
1875, onthe
otherhand,
althoughhaving
pagepriority
overT.
vicina(it
was
describedin
thesam
epublication)
was
basedon
aspecim
entoo
small
tobe
allocatableto
aparticular
populationon
thegrounds
ofm
orphology.T
estudoguntheri
was
proposedby
Baur
(1889),for
aprobably
veryheterogeneous
seriesof
fivespecim
ensthat
Gunther
(1877)
hadallocated
toT.
elephantopus.B
aurconsidered
thatT.
elephantopusand
T.vicinaw
eresy
nony
mous,
and,in
thatG
unther’ sseries
was
not(in
Baur’ s
opinion)identical
toT.
vicina,it
neededa
newnam
e.T
heargum
entsboth
infavor
ofand
against thisallocation
were
farfrom
persuasive,especially
inview
of thefact
thatBaur
didnot form
allydeclare
aholotype
forhis
T. guntheri—
hew
asreally
onlyrenam
ingG
Unther’s
series,andG
untherw
as
notusing
anew
name,
butrather
allocatingnew
specimens
toH
arlan’s1827
name.
The
largestspecim
enin
Gunther’s
seriesw
asa
78cm
oldm
aleskeleton
inthe
Oxford
Un
iver
sityM
useum(O
UM
8656), oftheflat-shelled
typesim
ilartoV
anD
enburgh’ sm
alesfrom
Sierra
Negra.
The
remaining
specimens
(anim
mature
male
andfem
ale,and
two
juve
niles,fromthe
Free
Public
Museum
inL
iverpool, theB
ritishM
useum,
andthe
Royal
College
ofS
urgeonsM
useum)
serveto
confuserather
thanclarify,
inthat
nonehas
known
provenancenorobvious
distinguishingfeatures, yetallm
ustbe
consideredpartofthetype
series, with
theactualholotype
undeclared.T
heonly
specimen
largeenough
todisplay
thedepressed
carapaceconsidered
diagnostico
fthe
guentherisubspecies
ormorphotype
isthe
Oxford
University
Museum
specimen,
describedby
GU
nther(1877)
as:“A
nadult
male
example:
aperfect
skeletonw
ithcarapace,
butw
ithoutepiderm
oidplates.
The
carapaceis
31
incheslong.
History
ofth
especim
enunknow
n;purchased
ofa
dealerin
Paris
for
theO
xfordU
niversityM
useum,
andkindly
lentto
me
byP
rofessorR
olleston,F
.R.S
.”A
lthoughI
knowof
nolitera
turein
which
thisspecim
enis
identifiedas
theholotype
ofT.
guentheri,it
isso
identifiedin
theO
xfordU
niversityM
u-
seumcatalogue
(I.S
wingland,
inlitt.,
29O
ctober1974).
1
thereforehereby
formalize
what
hasalready
beenassum
ed,
anddeclare
thisspecim
en(O
UM
8656)the
lectotypeof
Testudo
guentheriB
aur,1889.
As
aparenthetical
footnote,R
othschild(i902b), after examining
thealleged
holotypeofT
elephantopus,decided
that therew
ereno
groundsfor
Baur
tohave
proposeda
substitutenam
efor
Gunther’s
series.It thus
seems
appropriatehere,as
first reviser, toutilize
theepithet
vicinaover
thesim
ultaneouslypublished
inicrophyesfor
thesubspecies
inhabitingboth
thehigh
andlow
terrainof
thefour
southernvolcanoes
ofA
lbemarle
Island.W
hilesom
em
inorm
orphologicaldivergence
may
[,“,
[.‘,.
..%
--
%.
-._
s•
.
...
PR
iTC
HA
RD
The
Galapagos
Tortoises
49
existam
ongthese
populations(e.g.,
guentherishow
sa
tendencyfor
largeadultm
alesto
besom
ewhatflat-shelled),
theproportion
ofspecim
ensin
which
distinctivecharacter-
isticsare
presentis
toolow
tojustify
nomenclatural
recog
nitionof
differentsubspecies.
Afew
individualsof
allof
thesepopulations
may
showa
tendencytow
ardsm
arkedlysaddlebacked
shells(e.g., alarge
male
Isawalive
onV
olcanA
lcedo[Fig.
32j;or
thesoutheastern
Albem
arlespecim
enfigured
byG
arman
[19171in
hisP
late16),but these
aretoo
rareto
affecttheoveralldesignation
of thesepopulations
asbasically
dome-shelled. T
heydo,how
ever,serveto
indicatethat
thegenes
forthe
saddlebackedshell
may
bepresent
inall
populationsin
acertain
frequency,and
thatw
hensuch
phenotypesare
favoredby
appropriateenvironm
entalcon-
ditionsthey
may
bem
anifestedthroughout
thepopulation
relativelyrapidly.
Butapart from
selectionfor
asaddleback
shellm
orphologyon
thedrier,m
orebarren
islands,mostof
theother
featuresthat
havebeen
utilizedto
differentiatesubspecies
orpopulations
—overall
size,shell
smoothness
orsculpturingand
degreeoferosion,degree
ofbossingofthe
carapacescutes,and
detailsofshellproportions
aresurely
responsesofthe
individualto
environmental
circumstances
ratherthan
geneticdifferences.
Moreover,underprim
ordialconditions,G
alapagostortoises
were
subjecttono
predationonce
theyhad
passedthe
veryearly
growth
stages,andas
theonly
largeherbivores
intheirenvironm
ent,thereis
noreason
tobelieve
thatfitnessw
ouldbe
lessin
tortoisesdem
onstrating
minordivergences
fromthe
typicalshellform,as
longas
thearchitectural,
supportiye,and
thermoregulatory
func
tionsof the
shelland
shellopenings
were
notcomprom
ised,and
mating
successw
asnot
reduced.T
helarge
sizeand
greatertendency
towards
shellsad-
dungofthe
adultmales
ofm
ostorall
Galapagos
racesm
aythus
bem
anifestationsof
sexualselection,
andthe
require-m
entsofenhanced
mating
success.Had
adultmales
notbeenavailable
totaxonom
ists,itis probablethat80%
ofthenam
esproposed
for“new
”G
alapagostortoise
speciesw
ouldnever
haveseen
thelight
ofday.
Sum
mary
ofP
roposedN
omenclature
The
Galapagos
tortoisesare
includedin
thegenus
Geochelone
andthe
subgenusC
helonoidis.T
echnically,the
specificepithet
shouldbe
californ
iana
Quoy
andG
aimard,
1824a,but
untilthis
name
canbe
petitionedto
besuppressed
bythe
ICZ
N,
inthe
interim,
asa
conve
nienceand
toavoid
givingcredence
toa
name
thatItrust
hasno
future,I
shallutilize
theepithet
nigra,a
name
orig
inally
pro
posed
asT
estud
onig
raQ
uoyand
Gaim
ard,
1824b,as
theco
mb
inatio
nG
eoch
elone
(Chelonoidis)
nigra.
Synonym
yof
Geochelone
(Chelonoidis)
nigra
1824aT
estudocaliforniana
Quoy
andG
aimard
Bull.
Sci.N
at.Paris
I:90.Type
locality:“C
alifornie.”
(An
unusednam
eto
besuppressed
bypetition
toIC
ZN
).I824b
Testudo
nigraQ
uoyand
Gaim
ardV
oy.U
ran.P
hys.Z
ool.,
174.T
ypelo
cality:
“Californie.”
1827T
estudoelephantopus
Harlan
J.A
cad.N
at.Sci.
Philadelphia
5(2):284.T
ypelocal-
ity:“G
alapagosIslands.”
Nom
endubium
(atsu
bsp
ecific
level).1835
Testudo
nigritaD
umdril
andB
ibronE
rpdtol.G
en.2:80.
Type
locality:unknow
n.N
omen
dubiu,n(at
subspecificlevel).
1854T
estudoplaniceps
Gray
Proc.
Zool.
Soc.L
ondon1853:12.
Type
locality:“G
alapagosIslands.”
Nom
endubium
(atsubspecific
level).1875
Testudo
rnicrophyesG
untherPhil.T
rans. Roy.
Soc.London
165:275.Type
locality:“H
ood’sIsland”
(bysupposition).
A/om
endubium
(atsubspecific
level).1902
Testudo
wallacei
Rothschild
Novit.
Zool.
London
9:619.T
ypelocality:
“Chatham
Island?”N
ornendubiuni
(atsubspecific
level).19
17T
estudoclivosa
Garm
anM
em.
Mus.
Com
p.Z
ool.30:283.
Type
locality:“M
ascarenes?”N
omen
dubium(at
subspecificlevel).
I917
Testudo
tvpicaG
arman
Mem
.M
us.C
omp.
Zool.
30:285.T
ypelocality:
un
known.
Nom
endubium
(atsubspecific
level).1952
Testudo
(Chelonoidis)
elephantopus,W
illiams
Bull.
Am
er.M
us.N
at.H
ist.99:555.
1967G
eochelone(C
helonoidis)elephantopus,
Pritchard
Liv.
Turt.
World.
156.1980
Chelonoidis
elephantopus,B
ourB
ull.M
us.N
at.H
ist.N
at.Paris
(4)2:546.
Irecognize
tensubspecies
ofG
eochelonenigra,
with
theirreferenced
maps,
photographs,type
localities,synonym
izednam
es,and
islandsor
areasofo
rigin
,listed
asfollow
s:
Geochelone
nigranigra
(Quoy
andG
aimard,
1824b)(M
aps1,
8;F
igs.6—
8,44—
45)T
ypelocality:
“Californie.”
Restricted
toC
harlesIs-
land(S
antaM
ariaor
Floreana)
(extinct).T
estudonigra
Quoy
andG
aimard,
1824b.T
estudogalapagoensis
Baur,
1889.T
ypelocality:
Charles
Island.T
estudoelephantopus
galap
ago
ensis,
Mertens
andW
ermuth,
1955.C
helonoidisgalapagoensis,
Bour,
1980.
Geochelone
izigraabingdoni
(Gunther,
1877)(M
aps1,
2;Figs.
21—22,
31,34—
35,frontispiece)T
ypelocality:
“Abingdon
Island”(P
inta)(extinct
inthe
wild;
onecaptive
survivor).Nam
erequires
valid
ation
bytheIC
ZN
bysuppression
oftheseniorsynonym
Testudo
ephippiuniG
unther,I875.
F50
CH
ETON
IAN
RESEA
RC
HM
oNouR
APfls,
Num
berI
—/996
Testudo
ephippiurnG
unther,1875.
Type
locality:C
harlesIsland
(inerror).
Nam
eprev
i
ously
used
erron
eou
slyfo
rthe
Duncan
Islandsub-
species.
Testudo
abingdoniG
unther,1877.
Type
locality:A
bingdonIsland.
Testudo
elephan
topus
ephip
piu
m,
Merten
sand
Werm
uth.1955.
Chelonoidis
abingdoni,B
our,1980.
Chelonoidis
ephippium,
Bour,
1980.G
eocheloneabingdoni,
Fritts,
I983.G
eocheloneepphippium
,F
ritts,1983.
Geochelone
nigrabecki
(Rothschild,
1901)(M
aps1,
10;F
ig.19)
Type
locality:“C
apeB
erkeley,northern
pointof
Albem
arleIsland.”
Northern
andw
esternslopes
of
Volcan
Wolf,
Albem
arleIsland
(Isahela).T
estudobecki
Rothschild,
1901.G
eocheloneelephantopus
becki,P
ritchard,1967.
Chelonoidis
becki,B
our,1980.
Geochelone
becki,F
ritts,1983.
Geochelone
nigrachatham
ensis(V
anD
enb
urg
h,
1907)(M
aps1,
5;Fig.
17)T
ypelocality:
“Chatham
Island.”S
outhwestern
andcentral
Chatham
Island(S
anC
ristóbal)(extinct).
Testudo
chathamensis
Van
Denburgh,
I907.T
estudoelep
han
top
us
chath
amen
sis,M
ertensand
Werm
uth,1955.
Geochelone
elephantopuschatham
ensis, Pritchard,
1967.C
helonoidis(hatharnensis,
Bour,
I980.G
eochelonechathaniensis,
Crum
ly,1984.
Geochelone
nigradarw
ini(V
anD
enburgh,1907)
(Maps
1,9;
Figs.
14.15,
26)T
ypelocality:
‘James
Island”(S
anS
alvadoror
Santiago).
Testudo
darwini
Van
Denburgh,
1907.T
estudoelephantopus darw
ini, Mertens
andW
ennuth
, I955.G
eocheloneelephantopus
darwini,
Pritchard,
1967.C
helonoidisdarw
ini.B
our,1980.
Geochelone
darwini,
Fritts,
1983.
Geochelone
nigraduncanensis
(Garm
an,
1917)(M
aps1,
3;Figs.
1,2,
/0,25,
36—4/, fro
nt
cover)T
ypelocality: “D
uncanIsland”
(Pinzón). T
hissu
bsp
e
ciesp
revio
usly
erron
eou
slydesig
nated
Testudo
ephippiurnG
unther,1875.
Testudo
duncanensisG
arman,
1917.N
ornennudum
.G
eochelonenigra
duncanensis,P
ritchard,I996.
Geochelone
nigrahoodensis
(Van
Denburgh,
1907)(M
aps1,
4;Figs.
3,1
1,24)
Type
locality:“H
oodIsland”
(Española).
Testudo
hoodensisV
anD
enburgh,1907.
Testudo
elephantopushoodensis, Mertens and
Wem
rnth, 1955.
Geochelone
elephantopushoodensis,
Pritchard,
1967.C
helonoidishoodensis,
Bour,
1980.G
eochelonehoodensis,
Crum
ly,
1984.
Geochelone
nigraphantastica
(Van
Den
burg
h,
1907)(M
aps1,
7;Fig.
20)T
ypelocality:‘N
arhoroughIsland”
(Fernandina)(prob
ablyextinct).
Testudo
phantasticusV
anD
enburgh,1907.
Testudo
phantastica,V
anD
enburgh,1914.
Testudo
elephantopus phantastica, Mertens
andW
ermuth,
1955.G
eocheloneelephantopus
phantastica,P
ritchard,1967.
Chelonoidis
phantastica,B
our,1980.
Geoche/one
phantastica,C
rumly,
1984.
Geochelone
nigraporteri
(Rothschild,
1903)(M
aps1, 6;Figs.
4,16, 23, 27, 42—
43, prologue,back
cover)T
ype
locality
:“In
defatig
able
Island
.”S
outh
ernm
dc-
fatigableIsland
(Santa
Cruz).
Nam
erequires
valid
ation
bythe
ICZ
Nby
suppressionofthe
possiblesenior
synon
ym
Testudo
nigritaD
umdril
andB
ibron,1835,
wh
ose
ho
loty
pe
isof
uncertain
pro
ven
ance.
Testudo
po
rteriR
othschild,I903.
Geochelone
elephantopusporteri,
Pritch
ard,
I967.
Geochelone
porteri,F
ritts,1983.
Geochelone
nigravicina
(Gun
ther,
1875)(M
aps1,
10,11;
Figs.5,
9,24,
32,48—
58, prologue)T
ypelocality:
unstated;given
as“Iguana
Cove, Süden
derinselAlbernarle”
byW
ermuth
andMertens
(1977).S
outhernand
middle
Albem
arleIsland
(Isabela), fromIguana
Cove,
Cerro
Azul,
toS
ierraN
egra,V
olcanA
lcedo,and
toT
agusC
ove,V
olcanD
arwin.
Testudo
vicinaG
unther,1875.
Testudo
gOntheri
Baur,
I889.T
ypelocality: unstated, givenas
“Villam
iel, SUdw
estender
Insel Albem
arle”by
Werm
uthand
Mertens,
1977.T
estudon’zacrophyes
Garm
an.
I917.
Typ
elo
cality:
“San
taIsah
elaislan
d(A
lbem
arle)near
Tag
us
Cove.”
Testudo
vandenburghiD
cSola,
1930.N
omen
nud
uin
.
Type
locality:“F
ortym
ilesfrom
Villam
il.atthe
coaston
thesouthern
borderof
Perry
Isthmus
[Albem
arle
Islandi.”(T
hispopulation
nowextinct).
Geochelone
elephantopusguntheri,
Pritchard,
1971a.
Geochelone
elephantopusguentheri,
Pritchard,
1 97lb.
Chelonoidis
elephantopus,B
our,1980.
Chelonoidis
guentheri.B
our,1980.
Geochelone
vicina.C
rumly,
1984.G
eochelonevandenbitrghi,
Cru
mly
.1 984.
There
arefour
possiblydistinct subspecies
forw
hichno
names
areyet
available:
Geochelone
izigrassp.
(Maps
1,5
Fig./8)
PRITCHA
RD—
The
Galapagos
Tortoises
51
Northeastern
Chatham
Island(S
anC
ristóbal). Distinct
fromextinctsouthw
esternand
central Chatham
Islandpopulations.
Geochelone
nigrassp.
(Maps
1,6;
Figs.
28—29)
Northw
esternIndefatigable
Island(S
antaC
ruz).D
istinctfrom
otherIndefatigable
populations,but
may
bean
imported
population,possibly
fromD
uncan.G
eochelonenigra
ssp.(M
ap1)
Barrington
Island(S
antaFe).
Extinct,
possiblyintro-
duced.G
eochelonenigra
ssp.(M
ap1)
JervisIsland
(Rábida).
Extinct,
possiblyintroduced.
Efforts
shouldbe
made
tosecure
typem
aterialfor
thefirst
two
of
theseform
s,at
least.E
ibl-Eibesfeldt
(1959)reported
thathehad
foundskeletal
material
innortheastern
Chatham
in1957,
andthat
thishas
beendeposited
inthe
Senckenburg
Museum
(Frankfurt-am
-Main).
Perhaps
thism
aterialwould
beadequate.T
akinglivingtortoises
fromthe
Galapagos
isnow
prohibited,yetvouchermaterial—
at leasta
holotype,such
asa
salvagedshellfrom
anatural
mortality
isessential
ifa
descriptionof
anew
formis
tohave
legitimacy.
Moreover,
theneed
isnot
purelyacadem
ic.C
onservationinvestm
enttends
toconcentrate
onnam
edtaxa,
andthe
northwestern
Indefatigabletortoises,
althougham
ongthe
mostdistinctive
inthe
archipelago,havereceived
noconservation
attention,despitetheirrarity,presum
ablyin
partbecause
thepopulation
hasnot
beennom
enclaturallyrecognized
asa
subspecies.
SU
RV
IVA
LS
TA
TU
S
The
differentpopulations
ofG
alapagostortoises
haveall
come
undervarying
degreesof
threatto
theirsurvival
duringthe
lastfew
centuries.S
ome
populationshave
be-com
etotally
extinct(e.g.,
Charles
andsouthw
esternC
hatham),
some
arenearly
orprobablyalready
extinct(e.g.,A
bingdonand
Narborough),som
ehave
possiblybeen
savedfrom
extinctiononly
byheroic
conservationefforts
(e.g.,H
oodand
Duncan),
anda
fewretain
relativelyrobustp
opu
lations(e.g., southw
estern1ndefatigable
andV
olcanA
lcedoon
Albem
arle).A
llpopulations,
however,
continueto
beat
riskand
extremely
sensitiveto
thevagaries
ofhuman
explo
itation,
feraland
nativeanim
aldepredation,
andvarious
environmental catastrophies.
Aconstant
vigilis
requiredto
monitor
populationlevels
andtheir
changesthrough
time
ifconservation
managem
enttechniques
areto
beeffective.
The
following
sectionpresents
anhistorical
reviewof
thesurvival
statusof
eachof
theseparate
populationsof
Galapagos
tortoises.from
theearliest
recordedinform
ationup
toas
nearthe
presentas
thepublished
literatureallow
s.T
hisknow
ledgeof
former
populationpatterns
iscritical
toany
understandingand
properinterpretation
ofcurrent
dynamics
asthey
relate
Map
2.Abingdon
Island(P
inta).Shaded
areais
where
tortoiseskeletons
havebeen
foundin
ravines.T
helast
livingtortoise
(Lonesome
George)
was
foundnear
thedot.
toconservation
andm
anagement
issues.O
ngoingcensus
work
bythe
Charles
Darw
inR
esearchS
tationcontinues
tom
onitorand
document
thestatus
ofth
esepopulations.
Histo
ricalR
eviewof
Galap
ago
sT
orto
iseP
op
ulatio
ns
Detailed
accountsof
thestatus
ofthe
variouspopula
tionsof
Galapagos
tortoisesinclude
thoseof
Groom
bridge(1982)
andde
Vries
(1984).M
osto
fthe
dataquoted,
how-
ever,w
erealready
severalyears
old,and
them
ostrecent
accountbasedupon
extensivefirst-hand
information
isthat
ofMacF
arlandet
al.(1974a).
In1989,a
planto
recensusall
ofthe
tortoisepopulations
was
initiatedby
theC
harlesD
arwin
Research
Station
andthe
Galapagos
National
ParkS
ervice(L
.C
ayot, pers.com
m.).
As
ofI995, censuses
havebeen
carriedout for
10ofthe
populationsand
therem
aining5
areplanned
within
thenext
two
years.T
hesestudies
will
helpto
determine
whether the
intensiveconservation
effortsof
thelast
25years
arereflected
inincreased
populations,and
will
providevaluable
information
ongrow
thrates,
survivorship,and
overallpopulation
dynamics,
inthat
use-fulnum
bersof
individualsof
most
ofthe
survivingpopula
tionsw
erem
easuredand
marked
(byshell-notching)
inthe
1960sand
70s.A
bingdon(Pinta). —
This
population. heavilyexploited
byw
halersand
otherm
ariners,underw
entan
overallcol
lapsein
them
id-nineteenthcentury
(Tow
nsend,1925b;
Pritchard,
I 977).The
lastrecordedw
halingcrew
sto
obtainlarge
haulsof
tortoisesfrom
Abingdon
were
thoseof
theA
bigail (l42tortoisesin
I837), theHector(67
in1843),and
RO
CA
SN
ER
US
0
D•
CA
BO
CH
AL
ME
RS
CA
BO
IBB
ET
SON
Figure
33.H
abitatin
thesouthern
uplandsof
Abingdon
Island,1972.
PR
ITC
HA
RD
—T
he
Galap
ago
sT
orto
ises53
Fig
ure
35.B
onycarapace
ofadult fem
aleA
bingdonIsland
tortoiseslaughtered
bym
an.N
otecutedges
ofshell
andabsence
ofplastron.T
hism
ayhave
beenthelastfem
aleofthis
distinctivepopulation. C
arapacepresented
tothe
Charles
Darw
inR
esearchS
tationbythe
author.A
pril1972.
Figure
34.Lonesom
eG
eorge, thelastsurvivingtortoise
fromA
bingdonIsland, photographedjustbeforerem
ovalfromA
bingdonin
1972.
54CH
[LON
IAN
RESEARCHM
ONOGRAPHS,N
umber
1—
I 996
two
otherships
thattogether
collected33
in1848.
Since
then,no
haulof
more
than6
was
recorded.In
I901, R
ollo
Beck
collected2
forthe
Rothschild
collection(N
os.1
and
27, anold
male
andan
allegedlym
aturefem
ale,althoughthe
latterm
ayhave
beenan
imm
aturem
ale).In
1905—06,
the
Academ
yparty
(alsoincluding
Beck)
foundthree
livesp
eci
mens
anda
shell,all
males.
The
statusof
thepopulation
between
1905—06
andthe
late1950s
isunclear.
In1957
M.
Castro
reportedto
Eib
l
Eibesfeldt
(1959)
that tortoisesw
erestillto
befound
“far up
theslope
ofthem
ainvolcano,”
andC
astroand
Eibl-E
ibesfeldt
foundan
oldtrail
thatseem
edto
bea
tortoisetrail
ata
relativelylow
altitudein
1957. How
ever,in1964
Castro
and
Cavagnaro
were
onlyable
tofind
28skeletons
oftortoises,
allw
edgedor
trappedin
ravinesand
creviceson
Abingdon,
with
nosigns
oflivingspecim
ens(S
now,
1964).M
oreover,
becauseof
theabundance
offissures
andholes
inthe
area,
Snow
addedthat “there
must have
beenvery
many
more
that
were
notfound.”W
hilethis
accumulation
ofskeletons
may
haverepresented
normal,
casualm
ortalityover
many
de
cades(tortoise
bonesand
shellsare
stillfound
incaves
on
Charles
Island,150
yearsafter the
demise
ofthepopulation),
itissignificantthatgoats
hadbeen
introducedto
theislands
duringthe
precedingseven
years,and
hadm
ultipliedex
traordinarily.T
heim
plicationis
thusthat
thetortoises,
underunaccustom
edgrazing
pressure,had
beenforced
intoabnorm
alhabitat
(Fig.
33),and
haddied
inacci
dents.The
populationw
asdeem
edextinct,
buta
liveadult
male
was
seenbyJ. V
agvolgyi in197
1 (Pritchard,1977)(Fig.
34),and
in1972
thisindividual
was
relocatedand
trans
ferredto
theD
arwin
Research
Station
forsafekeeping,
where
itstillresides,now
known
asL
onesome
George.
The
circumstances
ofits
rediscoveryin
1972are
describedby
Cruz
(1994).E
ffortsto
locateotherindividuals
haveproved
fruitless.
The
goatson
Abingdon
haveprogressively
beenelim
inated
byshooting
overabout
adecade
(about 26,000w
ereshot in
1972,thefirst year),and
thevegetation
isnow
substantially
recovered(W
helanand
Ham
ann,1989).
Iwas
ableto
finda
carapaceof
anadult
female
onA
bingdonin
1972,the
plastronofw
hichhad
clearlybeen
severedby
human
action
(Fig.35).
This
was
depositedin
theC
harlesD
arwin
Research
Station
collection,and
may
bethe
onlyfem
ale
specimen
inany
collection.A
skull,cervical
vertebra,
andshell
fragments
ofa
largem
ale(P
CH
P50)
were
also
found,again
with
thecharacteristic
signsof
havingbeen
slaughteredw
itha
machete,
andm
orerecently
D.
Green
Figure
36.California
Academ
yofS
ciencesparty
collectingtortoises
onD
uncanIsland,
1906.Photo
byR
H.
Beck,reprinted
fromS
levin
(1931). T
heoriginal
captionread:
“We
hada
luxuriouscam
pon
Duncan
Island.T
hepet
hawk,
standingon
thedead
tree.w
asfed
turtle
meat
everyday
andbecam
equite
tame.”
‘-‘
.:4
.
‘*
‘—
--‘
PR
ITC
hA
RD
—T
heG
alapagosT
ortoises55
(pers.C
Oflirn.)
andcom
panionsfound
avery
largem
aleshell,
with
some
scutesstill
attached;this
toois
nowat
theC
DR
S.In
March
1981
(i.e.,
nineyears
afterthe
removal
ofL
onesome
George,
thesupposed
lastsurviving
Abingdon
tortoise),O
leH
amann
andL
indaC
ayotfound
asingle
olddried
tortoisescaton
Abingdon
Island(A
nonymous,
1982).In
thatthe
scatw
asintact
andlying
ontop
ofdried
grasses(L
.Cay
ot,p
ers.com
m.),and
furthermore
notingthat several
ElN
iño(high
rainfall)seasons
hadintervened
between
1972and
1981,the
possiblyexists
thatL
onesome
George
isnot
thelast.T
ownsend
(1925b)includes
aphotograph
ofaliv
eadult
Abingdon
tortoisethatw
as“am
ongthe
first...broughtto
theN
ewY
orkZ
oologicalP
ark,”but
gaveno
detailsof
collec
tion.B
aur(1889)
alludesto
anold
male
fromA
bingdonthat
hadbeen
collectedby
theA
lbatro
ssexpedition,
andthat
endedup,
“nearlycom
plete,”in
theU
SN
Mcollection.
Perhaps
thisw
asthe
same
one.D
uncan(P
inzón).T
hispopulation
was
reasonablyabundant
inthe
nineteenthcentury,
havingsurvived
theoccasional
onslaughtof
victuallingw
halers,but
sufferedheavy
collectionfor
scientificpurposes
(e.g.,8
collectedby
Baur
in1891,29
bythe
Webster-H
arrisexpedition
in1897,
86by
theA
cademy
expeditionin
1905—06;
Fig.36).
Itw
asfeared
virtuallyextinct
bythe
1920s(B
eebe,1924),
andindeed
Tow
nsend,com
bingthe
slopesof
Duncan
in1928
with
theassistance
ofeight
men,
didnot
finda
singletortoise,and
hesuggested
thattheyw
erealready
extinct. But
asm
allpopulationofaging
individualssurvived
(Figs.1
,2,37,
38),although
poachingalso
continued—
in1957,E
ibl
Eibesfeldt(1959)
collectedtheshellofan
adultmale
thathadbeen
killedabout
ayear
earlier.S
uccessfulreproduction
ofthe
Duncan
tortoiseap
par
entlyceased
inthe
1890sw
henferal
blackrats
reachedthe
island(first
recordedin
1891according
toP
attonet
al.,1975)
andpreyed
uponthe
hatchlingtortoises
with
nearly100%
efficiency.T
heresulting
absenceof
youngw
asal
readynoticed
byB
eck(1903),
andw
asreflected
inthe
abrupttruncationof the
lower-end
sizeclasses
ofthe1905—
06tortoises,
thesm
allestof
which
hadcarapace
lengthsof
about47
cm.
Indeed,in
theyears
imm
ediatelyfollow
ingtheir
introduction,there
isevidence
thattherats,even
inthe
presenceof
numerous
short-earedow
ls,w
ereparticularly
voraciousand
would
evenattack
largetortoises
ifthe
latterw
ereconfined
orcom
promised
insom
ew
ay.T
hus,w
henW
alterR
othschildenlisted
theservices
ofW
ebsterand
Harris
in1898
tocollecttortoises
forhimon
Duncan,H
arrisrecords
inhis
diary(in
M.
Rothschild,
1983)the
following
descriptionof
theparty’s
excursionashore
topick
upeight
tortoisesthat
hadbeen
locatedan
imm
obilizedw
ithheavy
stonestw
odays
earlier:“O
ngetting
tothe
crater,w
efound
onebig
tortoisedead;
oneofthe
bigrocks
thatw
ew
eightedit
with
hadshifted
andfallen
onits
neck,and
shutits
wind
off.Rats
hadgnaw
edoutone
ofitseyes
andhad
alsognaw
eda
pieceoutofone
hindfootofthe
livingsm
allerspecim
en.”S
ince1965,eggs
fromnatural
nestshave
beentaken
tothe
Darw
inS
tationfor
hatchingand
captivehead-starting,
Map
3.D
uncanIslan
d(P
inzón).Shadedarea
isoveralldistribu
tionofnative
andrepatriated
tortoises.D
otsare
nestingareas.
andthe
wild
populationof
150—200
oldadults
(MacF
arlandand
Reeder,
1975)had
beensupplem
entedby
226young
by1985
(Tierney,
1985)and268byD
ecember
1990(C
aporaso,1991),
althoughin
afew
casesthe
youngtortoises,
releasedtoo
early,have
beenkilled
byrats
(deR
oyM
oore,1979;
deV
ries,1984).
By
1990,the
nativeadult
populationw
asdow
nto
80—100
(Morillo
andC
ayot,1990),
with
afew
more
dyingeach
year(Figs.
15,39),at least8during
thedroughtof
1984—85.
Indeed,of
the64
females
marked
byshell-notching
from1963—
69,onlyabout sixteen
havebeen
recordedsince
1977,although
thisfinding
may
stem,
inpart,
fromboth
healingand
abrasionof
theperipheral
shellnotches
occurringover
theyears,
sothatthe
codednotch
combinations
canonly
beidentified
inless
than50%
ofindividuals
(L.
Cayot,
pers.co
mm
.).
Márquez
etal.
(1992)
reportthe
numbers
ofD
uncantortoises
hatchedat
CD
RS
from1965—
66to
1986—87,
andC
aporaso(1991)
givesthe
numbers
repatriatedfor
eachyear
from1970
to1990.
Repatriated
tortoisesare
shown
inF
igs.12,
40,and
41,and
totalnum
bersrep
atriated
aregiven
inT
able1.
Inthe
early1970s
around50
nestsw
erefound
eachnesting
season,butbythe
endofthe
decadethe
number
was
closerto
20per
year,and
from1980—
86few
erthan
I0w
erefound
annuallyin
thetraditional
nestingareas
(which
mci-
dentallysuffered
much
erosionby
rainbetw
eenD
ecember
1982and
July1983).
Nevertheless,
thisis
notnecessarilya
signof
actual(adult)
populationcollapse,
inthat
duringseveral
ofthesenesting
seasons,thepark
wardens
involvedin
thesurveys
were
newand
relativelyuntrained,
andalso
severalextreme
droughtyearsunquestionably
depressedthe
nestingproductivity.
Indeed,it
was
extremely
encouragingw
henm
oreexperienced
personnelw
ereable
tolocate
nofew
erthan
45nests
inS
eptember
1987(F
ritts,1988),
the
CA
DE
RA
0FT.
.
L_zI
EN
CA
fAD
A
‘-
BA
NC
OS
‘.‘
c%
•
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises57
majority
inthe
more
western
nestingzone,
andby
theend
of
theI987—
88season
thetotal
hadrisen
to96
nests(M
orillo,1992;
Cayot
etal.,
1993),although
extreme
droughtthe
following
yearreduced
thenum
berto
just
I1
nestsfound.in
1989—90,
29nests
were
found.In
thattortoises
arenot
normally
seenin
thecourse
of
nesting,it
isuncertain
what
proportionof
thenests
inrecent
yearshave
beenm
adeby
nativetortoises,
asopposed
torepatriated
animals,
butL
.C
ayot(pers.
comm
.)is
of
theopinion
thatthe
majority
arestill
made
bynatives.
The
growth
ratesof
therepatriated
animals
havebeen
veryvariable.In
some
years,nogrow
that alloccurred
fortenm
onthsfollow
ingrelease,w
hereasotheryear-classes
showed
uninterruptedgrow
th(M
etzgerand
Marlow
,1986).
The
oldestrepatriated
tortoises,from
1965—66.
havegrow
nat
excellentrates,even
surpassingnative
adultsin
sizew
ithin
two
decades(R
hodinet
al..1983;
Pritchard,
1985),and
Caporaso
(1991) reported
signsofb
reedin
gand
preliminary
nestbuilding.
The
repatriatedtortoises
haveshow
nan
ap
parentsex
ratioof
0.63: 1
males
tofem
ales,contrasting
with
thew
ildsex
ratioo
f2:1
(Metzger
andM
arlow,
I986),although
Morillo
(1992)fo
undth
enativ
etorto
isesex
ratioto
favorfem
ales.A
conservationprojectcan
onlybe
consideredcom
pletew
henthe
causesof
thecrisis
orthe
declinehave
beenaddressed
directly.T
hus,the
head-startingprogram
forthe
youngD
uncantortoises
was
essentialto
preventthe
nativepopulation
fromdisappearing
throughthe
periodof
almost
100%recruitm
entfailure.
Butthe
Charles
Darw
inR
esearchS
tationand
theG
alapagosN
ationalP
arkS
ervicedid
notm
issthe
opportunity,follow
ingthe
extreme
droughtof
1988,to
attempt
toelim
inatethe
feralblack
rats(R
attus
(Figures
onfacingpage)
Figure
37(u
pp
erleft).
Adult
male
nativeD
uncanIsland
tortoise.sheltering
fromthe
sun.Septem
ber1982.
Photoby
A.G
.J.R
hodin.
Figure
38(u
pp
erright).
Very
oldadult
male
tortoiseon
Duncan
Island.T
hisindividual,
known
inthe
islandsas
Onan,has
sincedied.
Note
thepitted
surfaceofthe
carapace,indicativeoffenestration
inthe
bonyshell,and
thegrow
thoflichens
inconcave
partsofthe
carapace.T
hissam
eindividualis
alsodepicted
inthe
paintingon
thefront
cover.A
ugust1986.
Fig
ure
39(low
erleft).
Carapace
ofa
female
tortoiseon
southwestern
Duncan
Island.E
veryyear
seesthe
naturaldeathofa
fewm
oreof
thenative
stockof
veryold
tortoiseson
Duncan.
September
1982.Photo
byA
.G.J.
Rhodin.
Figure
41.H
ealthy,large
subadultrepatriated
tortoiseon
Duncan
Island.N
otethe
contrastin
anteriorshell
profileand
elevationof
thecarapace
with
thenative
adultm
alein
Fig.38.
September
1982.
Figure
40(low
erright).
AtallO
puntiagalapageia
var.inacrocarpacactus
inthe
southernhighlands
ofD
uncanIsland
providesm
inimal
hutadequate
shadefor
ayoung,repatriated
tortoise.Septem
ber1982.
58CH
ELON
IAN
RESEARCH
MO
NO
GRA
PHS,N
umber
I—
1996
rattus),w
hosedepredations
uponthe
hatchlingtortoises
constitutedthe
primary
threatto
thepopulation.
The
time
seemed
especiallyopportune
inthat
thedry
conditionshad
notonly
reducedthe
ratpopulation
tothe
lowest
levelin
many
years, butalsothe
presumed
extreme
hungerand
thirst
ofthe
survivorsm
ightpredispose
themto
beattracted
toappropriate
bait.T
hecam
paignw
asdescribed
byC
ayotet
al.(1993).
Ittook
placeinitially
ona
localizedscale,
inthe
criticallyim
portantw
esterntortoise
nestingzone.
Subsequently
it
was
expandedinto
fivew
orkzones
thatcovered
thew
holeisland.
Total
personnelnum
bered47,
andbait
station
sw
ereestab
lished
at50
mquadrants
overthe
entireisland.
Poison
usedincluded
thecom
mercial
rodenticide
Kierat,
asw
ellas
theanticoagulant
Racum
inm
ixedw
ithrice.
The
campaign
clearlyreduced
therats
toa
verylow
levelindeed,
andin
thecourse
of
1989the
onlyevidence
of
survivingrats
was
foundat
asingle
siteon
thehighest
partof
theisland.
Nonetheless,
successcannot
yetbe
claimed,
ratshaving
extraordinarypow
ersof
reproductiononce
favorable
conditionsreturn.
Butthe
virtualelim
inationofthe
ratsprom
otedhealthy
increasesin
populationsof
small
nativevertebrates,
includinglava
lizards,m
arineiguanas.
Galapagos
doves,and
othersm
allbird
species.S
ign
ificantly,
however,
anum
berof
deadhatchling
tortoisesw
erefound,
l2in
March
1988an
d9in
late1989, w
ithsm
allholesin
thecarapace
suggestiveof
predationby
nativehaw
ks(B
uteogalap
ago
ensis)
ratherthan
byintroduced
rats(C
ayotet
al.,1993).
Italw
ayspresents
afrustration
to
Table
1.N
umbers
ofrepatriatedD
uncanIsland
tortoises.
Seasonof
Num
berY
earof
hatchingrepatriated
repatriation
1965—66
201970
1965—66
91971
1966—67
201971
1967—68
231971
1968—69
101972
1969—70
291973
1970—71
111974
1971—72
171976
1972—73
181976
1973—74
141977
1974—75
111979
1975—76
101980
1976—77
31981
1977—78
41983
1978—79
11983
1979—80
261984
1980—81
131985
1981—82
71985
1982—83
——
1983—84
111987
1984—85
111988
1985—86
—
1986—87
23
conserv
ationists
when
onerare
species
attacks
ano
ther,
andeven
more
sow
henit
ispossible
thatthe
predationupon
therare
nativespecies
may
havebeen
aresultofthe
virtualelim
inationof
theprevious
preythe
harmful
feralspecies.
Follow
ingthe
intensivefield
work
onD
uncanin
thelate
1980s,C
ayotet
al.(1993)
reportedthat
258tortoises
hadbeen
observedin
thecourse
ofthe
census,out
ofan
estimated
totalof
292anim
als,the
totalpresum
ablyincluding
bothnative
andhead-started
orrepatriated
animals,
butnot
hatchlingsor
post-hatchlings.O
fthe
I00native
tortoisesthat
hadbeen
marked
inthe
I960s,73
were
found
,alth
oug
hin
man
ycases
theo
rigin
alsh
ell-notch
combinations
were
nolonger
readable.H
ood(E
spañola).—
The
whalers
reducedthese
tor
toisesdrastically.
Hood
isa
small,
low-lying
island,and
isthe
firstoftheislands
tobe
encounteredby
mariners
arrivingfrom
aroundC
apeH
orn,and
thetortoises,being
among
thesm
allestin
thearchipelago,
were
easyto
carry.T
ownsend
(1925b)found
recordsof
atleast
1698tortoises
removed
fromH
oodfrom
1831to
1868, thelargehauls
beingtow
ardsthe
beginningof
thisperiod.
The
Isabella,for
example,
collected335
infive
daysin
1831,and
theL
operobtained
237in
fourdays
in1 834.B
utbyI 842
thenew
sw
asbad. T
heM
asterof
theG
eorgeW
ashingtonreported
that“T
errapin
[were]
veryscars”
onH
oodin
1842,and
in1847
theP
ersia“g
ot
but
fewtu
rpin
,”alth
ou
gh
the
Ch
arlesF
rederick
was
ableto
collect
67fro
mH
ood
inth
atyear.
But
thereafterthe
takew
asnever
more
than7
ona
givenv
isit.By
1905—06
thew
halersw
erelong
gone,but
insteadferalgoats, presentin
thousands, haddenuded
thelandscape,
virtuallyelim
inatingfood
orprotection
foryoung
tortoisesand
competing
dangerouslyw
iththe
survivingadults
forfood.
The
Acadeniv
expeditionin
thoseyears
onlyobtained
3live
tortoiseson
Hood
despiteprotracted
searches,and
thelaconic
conclusionw
as“evidently
theyhave
beenw
ellcleaned
out.”N
onetheless,som
esurvived.
Banning
(1933)reported
thatthe
Pinchot
expeditionof
1929had
recalledthe
“extinct”
Hood
tortoiseto
thestatus
of”rarity,”although
thetw
ospecim
ensencountered
were
bothrem
oved,running
ase
rious
risko
fpro
vo
kin
ga
returnto
theform
erstatus
(Thornton,
1991’?M
ap4.H
oodIsland
(Española). Shaded
areais
overalldistribu
tionof
repatriatedtortoises.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises59
Map
5.Chath
amIslan
d(S
anC
ristóbal).Northeastern
population:Shaded
areasrepresentlow
densitytortoise
distribution,stripedarea
isa
regionofhigh
tortoisedensity.Southw
esternand
(entralpopulation:The
blackdotis
theapproxim
atelocation
ofthelastlive
tortoisecollected
in1906,the
lightlyshaded
opencircle
isa
cavew
hereI7
skeletonsw
erefound
in1906,the
opencircle
marks
theapproxim
atelocation
where
thelastsurviving
specimen
may
havebeen
killedin
1933.
1971).
There
was
some
levelof
US
military
presenceon
Hood
duringW
WI1,
butrecords
arenot
availableas
tow
hethertortoises
were
encountered.B
utE
ibl-Eibesfeldt
( I959)stated
thata
shellw
ithscutes
intact,that
hefound
in1957,
was
froma
tortoisekilled
in1942.
The
tortoiseslocated
onthe
islandduring
theI950s
and60s
were
sofew
anddispersed
thatit
isassum
edthat
reproductiondid
notoccur
(Corley
Sm
ith,1977),
althoughitis
alsopossible
thathatchlingsw
ereproduced
but thattheystarved.
baked,or
failedto
survivethe
onslaughtsofhaw
ksand
otherpredators
ina
goat-clearedlandscape.
The
firstanim
allocated
inrecent
decadesw
asan
apparentm
ale,56
cmin
carapacelength,
foundfeeding
ona
fallenO
puntiacactus
incom
petitionw
ithfifteen
goatsby
M.
Castro
on31
August
1963(S
now,
1964).On
24Novernber
1963,anotherindividual,
59cm
inlength,
was
located,and
thefirst
specimen
was
re-sightedin
lateJuly
1964.Snow
estimated
thatthe
totalpopulation
couldnot
beabove
about20
mdi-
viduals,and
hew
asalm
ostcertainly
correct.T
hedecision
was
made
totransfer
alllivetortoises
thatcouldbe
foundto
acaptive
breedingcenter
onIndefatigable
Island,and
ulti
mately
agroup
of3m
ales(one
along-term
captiveat the
SanD
iegoZ
oo)and
I2fem
alesw
ereaccum
ulatedatthe
Darw
inS
tation.R
eproductionhas
beenexcellent.
By
1982,113
younghad
beenreturned
toH
oodIsland
(Reynolds,
1982),and
theferal
goatshad
beenelim
inatedby
shooting.B
y1986.
184had
beenrepatriated,
ofw
hichonly
7had
beenfound
dead(A
nonymous,
1986).F
iguresgiven
byM
árquezet
al.(1992)
indicatethat,
between
1980—81
and1986—
87,the
captivecolony
laidbetw
een24
and32
nestsannually
(138—204
eggs).P
ercent-age
hatchingw
aslow
(9.3—33%
)butatotalof206
indiv
idu
alsfrom
theyear-classes
1970—71
to1984—
85had
beenreleased
onH
oodIsland
asof
1992,w
ith53
animals
from1984—
85and
I985—86
stillin
captivity.The
earlierreleased
animals
havereached
adultsize
(pers.ohs.),
anda
recent
report(C
aporaso,1991)
indicatedthat
328H
oodIsland
tortoiseshad
beenrepatriated
asof
January199
1.U
pdatedfigures
aregiven
byC
ayotand
Morillo
(inpress).T
heyreported
thatreleaseshad
occurredattw
osites
(El
Caco
andL
asT
unas)in
theinterior
ofH
oodIsland.
By
theend
of1993,
575tortoises
hadbeen
released,w
ith288
more
stillheld
incaptivity
atthe
Breeding
andR
earingC
enterat the
Charles
Darw
inR
esearchS
tation.The
survivalrate
ofthevery
youngtortoises
(releasedat
1 2—18
months
ofage)
was
poorP
iños(1987)
indicateda
mortality
rateof
56%w
ithinthe
firstyear. One
tortoisew
asapparently
killedby
anative
hawk
theday
following
itsrelease.
Butbetw
eenI988
and1992,
them
inimum
survivorshipw
as55%
,and
atotal
of202
differenttortoises
was
observedon
theisland
(averageof
89individuals
observedper
trip).F
ournest
attempts
were
foundin
1 990(M
árquezetal.,
I991), allatEl
Caco,
two
carcassesof
hatchlingskilled
byhaw
ksw
erefound
inlate
1990,and
alive
hatchlingw
asfound
inJune
1991.S
ixteennests
were
foundin
Novem
ber199
1,andI2
nests(and
two
livehatchlings)
in1993.
Chatharn
(San
Cristóbal).
—T
hetortoises
ofsouth-
western
andcentral
Chatham
Islandw
ereespecially
heavilycollected
byw
halersin
them
id-nineteenthcen
tury
—T
ow
nsen
d(1925b)
fou
ndth
at,of
13,013tortoises
reportedto
havebeen
collected
byw
halersin
theG
alapagosfrom
I831
to1868, 4798
were
fromC
hatham.
Am
inuterem
nantof
thispopulation
survivedto
1905—06
(Van
Denburgh,
1914),
butit
isnow
extinct.P
ossiblythe
lastw
askilled
bysettlers
nearP
rogresoin
1933(B
anning,1933).
Nevertheless,
severalhundred
tortoisesstill
livein
thearid
northeasternpart
ofC
hatharnIsland,
althoughthey
areapparently
adifferentsubspecies.
Feral
dogshave
menaced
thispopulation
inthe
past,but
may
nowhave
beenelim
inated.
Inaddition,
captivehatching
andhead-starting
pro-gram
shave
resultedin
therelease
of13
animals
fromthe
C(O
BU
JOC
A8O
Nor
;:
60C
HE
LO
NIA
NR
ES
EA
RC
HM
oN
ouR
Aplis,
Num
ber1
—1996
1972—73
year-class,6
from1973—
74,23
from1974—
75,5
from1975—
76, and8
from1978—
79(M
árquezet
al.,1992).
Gordillo
andK
endrick(1989)
pronouncedthat
thetortoise
populationofnortheastern
Chatham
(Fig.18)w
asone
of them
ostsecure
inthe
archipelago,follow
inga
decadeof
diii-gent
researchand
managem
entactivities
byN
ationalP
arkpersonnel.
Indefatigable(S
antaC
ruz).—
This
large,central
is-land,
todayinhabited
byseveral
thousandpeople
andthe
headquartersofthe
Charles
Darw
inR
esearchS
tationand
theG
alapagosN
ationalPark
Service,
was
settledon
atem
po-rary
basisbefore
1846,w
henhuts
occupiedby
gatherersof
archil(a
lichenused
inthe
dyeindustry)
were
built atWhale
Bay
onthe
west
coast,w
itha
trailleading
inlandto
small
citrusgroves
atS
antaR
osaand
Salasaca
(Perry,
1972).T
hesehuts
were
finallyabandoned
(althoughstill to
beseen
attheturn
ofthe
century),and
Indefatigabledid
not receiveperm
anent inhabitantsuntil the
Norw
egiancolonizing
expe
ditionof
1926.T
ortoisesfrom
Indefatigablefeatured
fromtim
eto
time
inthe
logbooksof
Am
ericanw
halersT
ownsend
(1925b)reported
that 44w
erecollected
bythe
Pacific
in1833,
140by
theA
bigailin
1834,12
bythe
Benezet
in1834,
2pIus
“many”
bythe
Pio
neer
in1836,
45by
theA
lfredT
ylerin
1 845,and
36by
theR
oman
in1848.
But
ingeneral
thecollecting
pressurew
asslight
—
of13,013
tortoisesreported
collectedin
theG
alapagosbetw
een
1831and
1868,only
279w
erefrom
indefati
gable.Cookson
( 1876)claim
edthat
tortoisesw
ereso
rareon
Indefatigablein
I875that
theyw
ereno
longerhunted,
andR
othschild( 1903
)incorrectly
claimed
that,since
theI 812
visitof
David
Porter
tothe
Galapagos,
“noone
hasfound
tortoiseson
Indefatigable, anditw
assupposed
theyhad
beenexterm
inated.“
But
hegoes
onto
reportthat
Beck,
undercontract
toR
othschild,had
founda
youngtortoise
nearthe
shoreof
Indefatigablein
I901,and
in1902, after
along
andw
earisome
hunt,B
eckcollected
7m
ore,ranging
upto
verylarge
size. TheA
cadem.y
expeditionin
1905—06
collected23.
andconcluded
that theindefatigable
tortoisew
asstill
fairlycom
mon
(Fig.42).
Inthe
yearsfollow
ingperm
anentsettlem
ent,slaughter
oftortoiseson
indefatigablew
asw
idespread.A
1930ex
pe
ditionto
thehighlands
ofIndefatigable,
describedby
Tow
nsend(1930)
andR
oosevelt(1930),
foundjusttw
oof
the35
Norw
egiancolonists
who
hadarrived
fouryears
beforestill
inresidence
Ofl
theisland,
andthey
andsom
eE
cuad
orian
settlersreg
ularly
huntedthe
torto
ises.T
ownsend
partyfound
asingle
animal
weighing
anesti
mated
250—300
pounds.L
ocalpeople
broughtin
8sm
allerones,
40—80
poundsin
weight,
andthese
were
takento
the
Fig
ure
42.Until trails
were
made
intothe
tortoisearea
onIndefatigable
Island,access
was
verydifficult, and
fewtortoises
were
removed.
Photoby
RH
.B
eck,reprintedfrom
Slevin(1931). T
heoriginal caption
was:
“Packingtortoises
throughthejungle
ofindefatigableis
nochild’s
play.”
p
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises61
Map
6.In
defatig
able
Island
(San
taC
ruz). L
ighter shadedareas
arethe
distributionsofthe
dome-shelled
southerntortoise
populations,the
darkershadedarea
isa nesting
zone. The
stripedarea
atCerro
Montura
isthe
distributionofthe
saddlebackednorthw
esternpopulation.
New
York
Zoological
Park.T
heparty
alsolearned
thatnatives
ofC
hathamIsland
hadcom
eto
Indefatigableeight
months
earlierand
killeda
number
oftortoises.
Tow
nsendw
rote:“W
hilein
thetortoise
countryw
em
easuredthe
bleachedshells
oflarge
tortoiseskilled
yearsbefore.
Som
eof
themw
erenearly
fivefeet
inlength
oftop
shellas
measured
overthe
curve.T
heym
usthave
weighed
consid
erablyover
500pounds... T
heN
orwegian
settlement on
thisisland,
recentlyabandoned, m
usthavekilled
many
tortoisesas
we
foundnum
erouslarge
skeletonsduring
ourbrief
stayin
thetortoise
country..
. Tortoise
meatw
asstillan
important
partof
theirthod
supplyand
tortoiseoil
servedthem
asbutter.
They
informed
usthat
youngtortoises
were
seldomfound.
Afive-pound
specimen
representsthe
onlysm
alltortoise
we
obtained.T
heG
alapagoshaw
k,w
hichis
more
abundant onIndefatigable
thanelsew
here,was
creditedw
ithdestroying
newly
hatchedtortoises.
Itis
possible,since
tortoisesare
nolonger
abundant, thathawks
serveto
hindertheir
increaseon
Indefatigable.”T
odaythe
hawk
isalm
ostextinct
onIndefatigable.
afew
individualssurviving
inthe
extreme
northwest.
Ram
bech(pers.
comm
.in
MacF
arlandet
al.,l974a)
estimated
thatat
leastone
totw
othousand
tortoisesw
erekilled
onthe
islandby
oilhunters
inthe
1930s.B
ym
id-
century,Indefatigable
hadbeen
settledcontinuously
forseveraldecades, and
thereports
ofseveralvisitorssuggested
thattortoise
huntingw
asstill
aproblem
.E
ibI-Eibesfeldt
( 1960),visiting
thegalapaguera
insouthw
esternin
defati
gablein
I957,reported
evidenceof
intensiveslaughter:
“.
..w
hatwe
foundw
asa
graveyard. The
shellsof slaughtered
tortoiseslay
scatteredeveryw
here.M
ostof
thesecarcasses
were
oldand
half-rottedaw
ay,though
some
were
quitefresh.
Sothis
areaw
asstill
ravagedby
present-daysettlers,
despiteall
theprohibitions.”
Eibl-E
ibesfeldt(1959)
inter-view
edsettlers
onIndefatigable,
andw
asadvised
that,25
yearsearlier,
tortoiseshad
beenfound
frmthe
aridzone
upto
thetransition
zone,although
theyw
ereless
abundanton
thenorthern
slopesthan
thesouthern.
Today
theyare
con-fined
tothe
southernslopes
(apart fromthe
disjunct colonyin
theextrem
enorthw
est).H
ealso
foundhones
andshell
fragments
ina
caveabouttw
om
ilesnorth-w
est ofAcadem
yB
ay,an
areafrom
which
theyhave
sincedisappeared.
And
hem
entioneda
fewrecords
fromthe
northernand
easternparts
ofthe
island.H
econcluded
thatthe
speciesw
as“in
rapidretreat.”
Hendrickson
( 1965)illustrated
theshells
ofslaughteredtortoises
foundon
Indefatigable,but
gaveno
details.M
acFarland
etal.
(1974a)indicated
that15—
25tortoises
PTA
CA
AR
ION
eAH
IAIO
PR
ER
OB
AH
IA1O
RT
UG
AN
EG
RA
0
ISLA
SP
LA
BA
HIA
CO
NW
AY
FICERROM
ON
TUR
A
CE
RR
OQ
LO
RA
DO
I.E
DE
N
CA
SCA
JOG
PISK1—
CA
CH
AC
HO
NT
LL
O
CFA
TA
L
CE
A%
CE
RR
OM
E
.
CR
RO
CR
KE
R
:::.,-
3:::
ED
1
EB
O
PTR
Ff
BE
1A
VIA
CE
RR
ON
UN
EZ
:‘\
P1JERN
EZ
II
—
TA
CIO
NIO
LO
GIcA
CH
AR
LE
SD
AR
WIN
AN
UN
EZ
PUE
RT
OA
VO
RA
L.0
iC
AA
MA
NO
PTA
EST
RA
DA
RA
HIA
TO
RT
UG
A
62CHELONIAN
RESEARCHM
ONOGRAPHS,N
umber
1—
1996
were
slaughteredannually
onIndefatigable
during1965—
68;15
in1969;6
in1970; an
d2
in1971.T
hereaftertherehas
beenlittle
killingofthese
tortoises,theresultoflaw
enforce-
ment
andeducational
effortsby
theC
harlesD
arwin
Research
Station
andthe
Galapagos
National
Park,
but
variousother
stressesrem
ain.In
certainyears,
forex
ample,
therehas
beenheavy
predationupon
thenests
of
Indefatigabletortoises
byferal
pigs(subsequently
brought
undercontrol).
Black
(1973)noted
thathe
hadencountered
carapaces
of11
tortoiseskilled
bym
anin
theeastern
sectionof
IndefatigableIsland,and
heestim
atedthatabout20
tortoises
were
slaughteredannually
inthe
yearsup
to197
1. P
riorto
theestablishm
entof
theN
ationalP
ark,a
Tortoise
Reserve
hadbeen
declaredin
southwestern
Indefatigable,inthe
area
southofthe
villageof S
antaR
osaand
eastof Puerto
Ayora,
adjacentto
thehighland
agriculturaland
pastureareas.
The
slaughteredanim
alsw
erefound
bothinside
and
outsidethe
Reserve.
Black
describedhow
thetortoises
were
brutallychopped
openw
hilestill
alive.Interest-
ingly,how
ever,the
shellsof
reportedlyslaughtered
an
i
mals
inthe
accompanying
photographw
erecom
plete
(i.e.,w
ithplastron
stillattached,
andno
obviousm
achete
marks
visible).A
searly
asI960—
61,
tortoiseson
Indefatigablew
erebeing
marked
with
paintednum
bersby
alocal
worker
answerable
tothe
Director
oftheC
harlesD
arwin
Research
Station,
andabout
1000anim
alsw
erecensused
inthis
way.
But
paintw
asunlikely
tolast
form
orethan
afew
months,
andsubsequently
paintednum
bersw
ereused
onlythr
short-
termstudies
where
thebehavior
ofspecific
tortoisesneeded
tobe
observedfrom
adistance.
Hendrickson
( 1965)reported
that700
Indefatigable
tortoiseshad
beenm
arkedby
carapace-notching,and
that
thepopulation
was
estimated
tonum
berbetw
een1500
and
2000individuals.
De
Vries
( 1968)indicated
that37
1m
ales,
367fem
ales,and
254juveniles(992
total)had
beenm
arked,
andhe
estimated
theactual
populationto
number
2000—
3000individuals.
Groom
bridge( 1982)
estimated
3000—
4000,as
didC
ayot(1985),
who
indicatedthat
more
than
2000had
beenm
arked.T
hevast m
ajorityof the
population
isconcentrated
inthe
southwestern
quadrantof
Indefati
gab
le,but
thereis
ano
utly
ing
pop
ulatio
n,
nowsep
arated
fromthe
main
populationby
asettled
agriculturalzone,
tothe
east,as
well
asan
enigmatic
saddlebackedp
op
u
lation
.very
differen
tin
extern
alm
orp
holo
gy
,in
thefar
no
rthw
est.T
hetortoises
arestrongly
migratory.
Generally
speak-
ing,large
tortoisesm
oveto
thehighlands,
nearthe
villageof
Santa
Rosa,
at drytim
esof
theyear,
andsom
erem
ainthere
year-round(F
ig.43).
Migrations
tothe
lowland
nesting
zonestake
placegenerally
inJanuary
toM
ay(R
odhouseet
al.,1975),and
veryyoung
tortoisesare
thususually
foundat
lowaltitudes.
How
ever,extraordinary
climatic
eventscan
Fig
ure
43.Adultm
aletortoises
ina
dryingpond
inthe
southeasternuplands
ofIndefatigable
Island.July
1986.
PRITCHARD—
The
Galapagos
Tortoises
63
precipitateextraordinary
movem
ents,and
Cayot
(1985)docum
enteda
dramatic
exodusof
tortoisesfrom
thehighlands
tothe
lowlands
duringthe
El
Niflo
eventof
1982—83,
when
torrentialrivers
flowed
innorm
allyarid
country,and
when
eventhe
lowland
nestingareas
were
sow
etthat
many
nestsw
ereabandoned
priorto
com
ple
tion.Snow
(1964)drew
attentionto
thedanger
ofextinctionof
thepopulation
ofsaddlebacked
tortoisesin
thearea
ofS
addlebackH
ill(C
erroM
ontura)in
northwestern
Ind
efatigable.Intensive
searchessince
thediscovery
ofthispopula
tionin
1962had
revealedonly
7individuals
(anadultm
aleand
sixjuveniles,
lessthan
fiveyears
old),plusa
deadvery
youngspecim
en.T
hepopulation
survivedin
the1970s
and1980s
—I
encountereda
youngtortoise
in1972
(Fig.29),
andreasonably
freshdroppings
in1982
(Fig.30),
andR
.R
eynolds(pers.
comm
.)reports
havingencountered
two
individualsof
medium
size.Study
andconservation
ofthis
populationis
nowa
priorityforthe
Galapagos
NationalP
arkS
ervice.T
heorigins
andtaxonom
icstatus
ofthis
populationrem
ainobscure.O
netantalizing
scrapofdata
isprovided
bya
photograph,published
byF
rittsand
Fritts
( 1982),of
anunidentified
mem
beroftheAcadem
yexpedition
of1905—
06(possibly
Slevin)
carryingw
hatappears
tobe
asm
all,sm
ooth-shelledsaddlebacked
tortoiseon
hisshoulders.T
hecaption
suggeststhat
thephotograph
was
takenon
Duncan
orHood,presum
ablybecause
bothofthese
islandshad
small
saddlebackedtortoises.
How
ever,in
thebackground
ofthe
photorises
what
appearsto
bethe
characteristicprofile
ofS
addlebackH
ill,thechieflandm
arkin
thetortoise
habitatofnorthw
esternIndefatigable,
andthe
presenceof
what
ap
pearsto
bea
white
sandybeach
inthe
rightforeground
iscom
patiblew
iththis
conclusion.M
anyspeculations
arepossible,
onebeing
thatthis
tortoise(w
ithothers?)
was
takenby
theA
cademy
partyfrom
nearbyD
uncanIsland,
andreleased
nearS
addlebackH
ill,possibly
constitutingthe
maternal
parentof
thetortoises
foundthere
today.T
hephotographs
(Snow
,1964)
(Fig.28)
ofalarge
male
(King
Gustav)
seenby
Gusch
Angerm
eyerin
1962suggest
alarger
formthan
isfound
onD
uncantoday,
butthiscould
bea
resultofbetternutrition
onthis
largerand
higherisland,where
evenlow
landtortoise
populationshave
accessto
humid
uplandsduring
droughtperiods.
Narborough
(Fernandina).
—T
hesurviving
recordsof
thew
halersand
buccaneersare
essentiallyfree
ofrecordsof
tortoiseson
Narborough.
The
singleearly
mention
was
thatof
Porter
(1822),
who
reportedtortoises
ingreater
orlesser
abundanceon
allthe
largerislands
ofthe
groupthat
hevisited.nam
ely“H
ood’s,Marlborough.Jam
es,Charles,and
Porter’s.”H
ealso
foundtortoise
boneson
Chatham
,butwas
onA
lbemarle
fortooshorta
time
tofind
tortoises,andhe
didnotgo
toB
indloe.Abingdon,D
owne,
orB
arrington.In
thatPorter’s
Islandis
asynonym
forIndefatigable,
andD
owne
anapparentsynonym
forD
uncanor
Tow
er(W
oram,
1989).it
appearsthat
allthe
largerislands
areaccounted
for,and
Marlborough
must
bea
misspelling
ofN
arborough,the
Map
7.Narborough
Island(F
ernandina).The
dotrepresentsthe
approximatelocation
ofthesingle
tortoisecollected
byR
olloB
eckin
1906.
name
givento
theisland
byA
mbrose
Cow
leyin
1685.T
heisland
hasnever
beeninhabited,
andrem
ainsfree
offeral
mam
mals.
Yet
thesingle
adultm
aletortoise
collectedby
Rollo
Beck
in1905
(Fig.20)
remains
theonly
tortoiseever
seenon
thisisland.
Eibl-E
ibesfelt(1959),
ina
1957w
alkclear
acrossthe
islandfrom
threem
ilesw
estof
Cabo
Douglas
(hem
usthave
meant
east—
Cabo
Douglas
isthe
westernm
ostpoint
ofN
arborough)up
tothe
calderaand
thencedow
nto
Punta
Espinoza,
foundno
traceof
tortoises.In
1964H
endrickson(1965),
with
theadvantage
ofa
heli
copterfor
explorationof
remote
areas,located
alarge
tortoisedropping
andan
Opuntia
cactusw
ithbite
marks
seemingly
made
bya
tortoise.Possibly
oneortw
oin
div
idu
alssurvive.T
hereason
fortheextrem
erarity
orextinctionof
thisform
cannotbe
ascribedto
human
activities.C
on
ceivably
thesporadic
major
eruptiveactivity
oftheN
arboroughvolcano
hasdestroyed
nestingareas
orseparated
themfrom
feedingareas.
Many
ofthe
largenum
bersof
landiguanas
(Conolophus)
onthis
islandtoday
nestinash
depositsatthe
bottomofthe
vastcaldera(W
erner,1982),a
habitatinacces
sibleto
tortoises.C
harles(S
antaM
ariaor
Floreana).
—T
hereis
noevidence
thatthis
populationsurvived
beyondthe
1840s.H
eller(1903)
estimated
thatextinction
occurredin
1840;B
room(1929)
estimated
1850.S
teadman
(1986)review
edall
availableinform
ation,and
thoughtthat1850
was
avery
reasonableestim
ate,follow
inga
decadeof
extreme
rarity.B
ythen
theisland
hadbeen
sporadicallyinhabited
forsom
etim
e—
aw
ildIrishm
anlived
therealone
atthebeginning
ofthe
century,and
ahighland
settlement
was
establishedby
GeneralV
illamilin
1829,althoughtroubles
beganw
henthis
was
convertedto
apenal
colony,and
itw
aseventually
abandoned.L
atera
newpenal
colonyin
thesam
earea
alsofailed.C
harlesD
arwin,on
avisitto
Charles
Islandin
1835, sawno
livetortoises,
onlyem
ptycarapaces
(Sullow
ay,1984).
Three
yearslater,
thefrigate
La
Venus
visited,butfound
notortoises,and
B.
Seem
annofFIM
SH
erald,visitingin
1846,reported
thetortoises
tobe
extinct.B
anning(1
933)m
en-tioned
thata
Charles
Islandsettler
(oneof
onlyeighton
the
PTA
.E
SPINO
SA
PTA
MA
NG
LE
CA
BO
DO
UG
LA
S
C.
HA
MM
ON
D
64C
HE
LO
NIA
NR
ESE
AR
CH
MO
NO
GR
APH
S,1”Juinber
1—
I 996
island)claim
edto
havediscovered
asm
alltortoise
thathad
sincedied,
andthis
claimw
asupheld
byneighbors.
Yet,
at
thattim
e,the
settlerson
Charles
Islandw
erea
groupof
celebrated—
or
inso
me
case
snoto
rious
—settlers
who
had
arriv
ed
from
Germ
any
in1929,
who
would
surelyhave
mentioned
tortoisesin
theirm
emoirs
ifthey
hadfound
any
(Strauch,
1936;W
ittmer,
1960).O
urknow
ledgeof
the
morp
holo
gy
of
thetortoises
of
Charles
Islandis
derivedfrom
shellsand
otherbones
collectedin
cavesnear
PostO
fficeB
ay,e.g., by
Tow
nsend(see
Broom
,1929).
Inad
di
tion,S
hurcliff(1930)
reportedthe
collectionof
severaldozen
tortoiseshells
forthe
Field
Museum
byhim
selfand
K.P
.S
chmidt
ina
cavernadjoining
thatvisited
byT
ownsend.
And
Steadm
an(1986)
foundabundant
frag
—---—
---
__
__
__
__
--—
-
__
__
__
Fig
ure
44.M
ou
nted
specim
enalleg
edly
represen
ting
the
extin
ctC
harles
Island
(San
taM
ariaor
Flo
reana)
torto
isein
theB
ritishM
useum.
PTA.
LUZ
DE
LD
IAPL
AY
APIC
ON
A
0.
\t)
CE
RR
OL
UZ
••.
(,
BA
HIA
DE
LAS
CU
EV
AS
CE
RR
OB
AL
LE
N
1A
SC
OIB
AR
RA
L)
—R
’OU
ES
RO
LA
GU
NA
LA
VA
PRIE
TA
LAB
OT
EL
LA
MO
NJA
‘\
ASIL
OD
ELA
PAZ
PTA.
EN
CIL
LA
DA
(SAD
DL
E)
,4?p
PTA.
SUR
Map
8.C
harlesIsland
(Santa
Maria
orF
loreana).T
hedots
represen
tcav
esw
here
suhfo
ssilto
rtoise
remain
sh
ave
been
foun
d.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises65
Fig
ure
45.B
onycarapace
ofthe
extinctC
harlesIsland
tortoise(adult
fema’e)
collectedin
acave
onthe
island(Field
Museum
,C
hicago).
ments
infour
cavesystem
s:F
inchC
ave,B
arnO
wl
Cave,
Cueva
deP
ostO
ffice(S
uperior),and
Cueva
deP
ostO
ffice(Inferior).
He
alsoreported
small
numbers
oftortoise
fragments
inother,
shallowcaves
inthe
Bahia
delas
Cuevas
region.A
partfrom
thesesubrecent
cavernspecim
ensand
fragments,
thereis
question—
orab
solute
doubt—
aboutall
ofthe
otherspecim
ensalleged
tobe
fromC
harlesIsland
(e.g., those
illustratedby
Garm
an,1917)
(Figs.
44—45).
The
seemingly
abruptextinction
ofthe
Charles
Islandtortoise
iscurious.
On
islandsof
comparable
sizelike
Abingdon
andH
ood,analysis
ofthe
whalers’
collectingrecords
(compiled
byT
ownsend,
1925b)indicates
quiteprecisely
theyear
inw
hichthese
tortoisepopulations
col
lapsedto
alow
level,but
actualextinction
was
avoidedby
theextrem
edifficulty
oflocating
thefew
,scattered
surv
ivors.
Yet
therecords
forC
harlesIsland
showsubstantial
collectionsin
theyears
imm
ediatelypreceding
apparenttotal
extinction.In
1831
theM
agnoliatook
155and
theF
rancestook
179;the
Hector
collected226+
in1832;
theO
ctavia235
in1833;
theB
engalcollected
100,the
Moss
350,and
theB
enezet120
in1834;
theB
arclaytook
50and
theB
enezet 40in
1835;and
theE
lizaA
dams
obtained24
in1837
(Tow
nsend,1925b;S
teadman,
1986).Moreover,these
recordsrepresent
onlya
fractionof
thevisits
byw
halersto
Charles
Islandduring
thisperiod
—there
were
31
inju
sttheten-m
onthperiodbeginningO
ctoberl832(R
eynolds,1835).
Subsequent
tothis,
theonly
recordsare
acouple
ofunquantitative
onesfor
thelate
1840s—
theC
ongareelogged
“some
Terrapin”
fromC
harlesin
1847,and
theA
urora“gotafew
Turrapin”
in1848,and
Tow
nsend(1925b)concluded
thatthesem
usthavebeen
tortoisesim
portedfrom
otherislands
forsale
tom
ariners.The
entryforthe
Fabius
inI858
(“boatsashore
afterturtle”),
inits
avoidanceof
some
formofthe
usualvernacular“terrapin,”
probablyreferred
tothe
East
Pacific
greenturtles
(Chelonia
agassizi)that
stillnest
onC
harlesIsland.
Doubtless
thetotal
extinctionof
theC
harlesIsland
tortoisew
ashastened
bythe
presenceof
thepenal
colony,the
relativelysm
allsize
ofthe
island,the
conversionof
thehighlands
tofarm
ingand
fruitplantations,and
theinflux
ofnum
erousferal
mam
mals,
includingpigs
anddogs.
Pigs,
cats,dogs,
goats,burros,
andcattle
were
allintroduced
around1832
(Hoeck,
1984);the
blackrats
andhouse
mice
may
havearrived
aroundthe
same
time.
Jervis(R
ábida).—
As
hasbeen
discussedabove,
onlyone
livetortoise
isknow
nto
havebeen
collectedon
thisisland
(Van
Denburgh,
1914),and
itm
ayhave
beenintro-
duced.A
lthoughcertain
authorshave
associatedthe
name
Testudo
wallacei
(orsubsequent
combinations,
includingG
eocheloneelephantopus
wallacei)
with
thealleged
tor
toiseof
JervisIsland,
theargum
entsin
favorof
thisare
hardlyacceptable.
Nevertheless,
theconclusion
thatJervis
neverhad
nativetortoises
hasto
takeinto
accountthe
comm
entsof
Steadm
anand
Zousm
er(1988),
mentioned
above,frustratingin
theirlack
ofcitation
orany
otherbasis
forcertainty,
butneverthelessalluding
withoutany
expres
sionof
doubtto
aform
erpopulation
ofboth
tortoisesand
landiguanas
onJervis
Island.B
arrington(S
antaFe).
—T
hetortoises
ofthis
islandare
somew
hatof
am
ystery.T
heisland
todayhas
nothad
nativetortoises
inliving
mem
ory,yet
itis
freeof
feralm
amm
als,and
suchvulnerable
speciesas
theendem
icland
iguanaand
therice
rathave
survivedin
goodnum
bers.T
ownsend
( 1925b)found
onlytw
oaccounts
ofw
halershaving
foundtortoises
onB
arrington—22
were
takenby
theG
eorgeand
Susan
in1839,and
1by
theH
enryH
.C
rapoin
1853.V
anD
enburgh(1914)
reportedthat
norecent
visitorto
thearchipelago
hadm
adenote
oftortoiseson
Barrington,
althoughhe
thencited
two
informants, one
ofwhom
claimed
tohave
takentortoises
offB
arrington16
yearsearlier
(i.e.,around
1890),and
theother,
acaptain
ofan
inter-island
schooner,w
hoclaim
edthat
tortoiseshad
beenfound
allover
Barrington
30years
earlier(i.e.,
aroundI876),
some
ofw
hichhe
hadcollected,
buthe
doubtedif
anyw
ereleft.
Possibly
theseobservations
were
accurate,but
ifthey
were
itis
odd,in
viewof
theexcellent
naturalharbor
andrelatively
easyterrain
ofBarrington,thatthere
was
not more
mention
ofcollectionoftortoises
byw
halersand
buccaneersa
centuryor
two
ago.T
heA
cademy
expeditionfound
oldbones
—pelvises
andlim
bs—
estimated
tohave
come
from14
individuals,but
noshells.
Som
eof
thebones
were
fromvery
largetortoises,
andsom
efrom
small
butapparently
66CHELONJAN
RESEARCHM
oNooR
PHs,
Num
ber1
—1996
fullym
atureones.
As
Van
Denburgh
said,“It
would
thusappear
that thetortoises
ofBarrington
Islandvaried
con
siderably
insize.”
The
disappearanceof
anative
tortoisepopulation
froman
islandthe
sizeofB
arrington, neverinhabitedby
man,and
without introducedpredators
(theonly
introducedm
amm
alsw
eregoats,
nowextirpated)
seems
unlikely,in
viewof
thepersistence
ofsmallpopulations
oftortoiseselsew
herein
thearchipelago
inthe
faceof
greater,m
ultiplestresses.
An
alternativeexplanation
isthat
thefew
comm
entatorsw
horeported
tortoisesfrom
Barrington
were
eitherunreliable
orhad
confusedB
arringtonw
ithanother
island.T
hen
om
enclature
ofthe
Galapagos
Islandsover
thecenturies
hasbeen
exceedinglyconfusing
(Woram
,1989), and
it issignificant,
forexample, thatthe
Whalers’
logbooksstudied
byT
ownsend
( I 925)m
aderegular
mention
oftortoises
collectedon
Crossm
anIsland,
asatellite
ofA
lbemarle
fartoo
small
tohave
anative
tortoisepopulation.
As
forV
anD
enburgh’ saccount
of thebony
fragments
foundon
Barrington, itis
significant that noshells
(not evenfragm
entaryones)
were
found, eventhough
theshellis
oftenthe
most durable
component
ofa
deadtortoise,
andthat
thelim
bbones
seemed
tocorrespond
tom
atureindividuals
ofvery
divergentsizes.
Apossible
explanationis
thatthe
excellentanchorage
onB
arringtonencouraged
overnightcam
pingby
whalers
andother
mariners,
who
may
havetaken
ashoreparts
ofbutchered
tortoises(i.e.
,entirelim
bs)from
theiron-hoard
stocksof
tortoisescollected
onvarious
otherislands,
toserve
ascam
pvictuals.
Eibl-E
ibesfeldt( 1959’) reportedthatsom
etortoises
hadbeen
releasedon
Barrington
inI956,and
hehad
seenone
ofthese,
ingood
health,in
thesum
mer
of1957.
There
isa
skullof
atortoise,
collectedon
Barrington
Islandby
Tjitte
deV
ries,in
thecollection
ofthe
Charles
Darw
inR
esearchS
tation.C
ayot(pers.
comm
.)reports
thata
largetortoise
was
founddead
onB
arringtonin
the1980s,
andthat
alive
onehad
appearedat
thevisitor
siteas
recentlyas
1988.Jam
es(San
Salvador
orS
antiago).—
James
isa
largeisland,
andtoday
thetortoises
arerestricted
tothe
farinterior.
How
ever,this
may
notalw
ayshave
beenthe
case.M
cBirney
andW
illiams
( 1969)presented
adetailed
map
oflava
flows
onJam
esIsland.
andindicated
thatthe
coastallow
landsalong
abouttw
o-thirdsofthe
shoreline,including
theentire
southerncoastal
zone,w
erecom
posedof
fresh,unvegetated
lava.T
hew
esternsections
ofthese
lavaflow
sw
erealready
inplace
onD
arwin’s
1835visit,but, in
that theflow
engulfedceram
icm
armalade
potsstashed
bybucca
neersin
1683(H
eyerdahl,1963),
it must be
concludedthat
them
ajorflow
soccurred
between
1683and
1835(Sim
kin,1984). A
ctually. majorfiow
sm
ayhave
occurredonly
ashort
time
beforeD
arwin’s
visit.Porter
(1815)
collectedno
fewer
than14
tonsof
tortoises(about
500,averaging
about60
pounds)neara
bayin
thenortheast, observing
that onlythree
ofthe
tortoisesw
erejadulti
males,
andall
thefem
alescollected
were,w
ithout exception, full ofeggs, ofwhich
10—14
were
“hard”(i.e.,
shelledand
readyfor
oviposition).T
heonly
bayof
anyconsequence
innortheastern
James
Islandis
Sullivan
Bay,
which
todayis
inthe
midst
ofthe
enormous
wasteland
offresh
lava.N
otortoises
arefound
anywhere
nearthe
baytoday.
On
ahike
inlandfrom
Sullivan
Bay
in1973,
1walked
about 40%o
fthe
width
ofthe
islandbefore
encounteringa
singleadult
female
tortoise(see
easternnesting
zonein
Map
9).T
heyoung
Charles
Darw
in,on
anovernight
visitto
theinterior
ofJam
esin
1835,found
aparty
ofS
panishtortoise
PTA
.C
OR
DO
VA
CA
LE
TA
BU
CA
NE
RO
C.C
OW
AN
BA
HIA
JAM
ES
0’
CER
RO
NEG
RO
SUG
ARLO
AF
a,
cb.
PTA
.•
BA
QU
ER
IZO
sc::?PA
RA
ISOPOZA
FELICID
AD
CPO
ZATR
AG
ICA
CER
RO
CO
LO
RA
DO
PTA
.C
.N
EP
EA
Nc:
c3M
AR
TIN
EZ
‘<
CA
BO
TR
EN
TO
N
Map
9. James
Island(San
Salvador). Lightershaded
areais
thedistribution
of tortoises, darkershaded
areasare
nestingzones. T
hedot
representsan
outlyingrecord.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises67
huntersencam
pedsix
miles
inland,at
analtitude
ofnearly
2000feet.
During
hisvisit
Darw
insubsisted
entirelyupon
tortoisem
eat(D
arwin,
1845).D
uringthis
same
period,T
ownsend
(1925b)reported
that whalers
regularlyobtained
tortoisesfrom
James.
The
Hector
obtained23+
in1834
and124
in1835.
The
George
andS
usancollected
68and
theB
enezet35+
in1835.
In1836
theH
espertook
13“plus
many”
andthe
Lim
aobtained
118in
1836and
224in
1837.T
heP
hoenixcollected
“12plus
sevenboatloads”
in1838
andthe
Chili
93in
1841.T
heE
quatortook
20in
1845,but
subsequent logbooksup
to1868
indicateno
furthertortoises
collectedon
James.
By
1905—06
thetortoises
were
rare,even
thoughthe
feralpigs
thatsubsequently
plaguedthe
islandhad
notyet
beenintroduced
(thedate
ofintroduction
was
estimated
as“before
1930”by
Hoeck
[1984]).G
oatshad
alreadybeen
therea
longtim
e,although
theprecise
historyof
theirpresence
remains
conjectural.It
isreported
thatgoats
were
introducedto
James
byC
aptainP
orterinI812, and
althoughthere
isno
reasonto
doubt this, thesem
aynot have
beenthe
goatsthat
producedthe
currentpopulation
(L.
Cayot, pers.
comm
.). Ultim
atelythegoats
reachedapopulation
estimated
at100,000
(Calvopina,
inH
oeck,1984),
butthe
Academ
yexpedition
(1905—06)
neverm
entionedgoats
onJam
es, andeven
inthe
1980sthe
goatsw
erereported
tobe
progressivelym
ovinginto
newhabitat, w
hichw
ouldsurely
not havebeen
thecase
ifthey
hadbeen
onthe
islandfor
nearlytw
ocenturies.
InI906
Beck
sawrem
ainsof
theS
panishtortoise
hunters’cam
p,still
litteredw
ithtortoise
bones,but
foundonly
5live
tortoises(3
male,
2fem
ale).A
saltm
inew
asin
operationat
James
Bay
fromabout
I924to
1930,and
thew
orkersapparently
conductedan
extensivetortoise
oilbusinessin
thew
esternand
southwest-
emparts
oftheinterior
ofJames
(A.
Ram
bech,pers.com
m.
inM
acFarland
etal.,
1974a).L
ongafter
them
inew
entbankrupt,
heavyequipm
entrem
ainedon
theisland,
anda
caretaker,Sr.
Leonardo
Apolo,
was
stillpresent
aslate
as1970.
Apolo
hada
sympathetic
interestin
thetortoises
andother
fauna,and
Leveque
(1963)reported
thata
partyfrom
theD
arwin
Station, escorted
byA
polo, hadfound
two
adultm
aletortoises,
about4
kminland,
inA
pril1963.
Apolo
showed
me
what
may
havebeen
thesam
eindividuals
in1970. A
t thattime
Ialso
sawa
smallbut
oldfem
aletortoise
entrappedin
asteep-w
alledcoastaltuffcone
southofJam
esB
ay(Fig.
26),and
foundthe
skeletonof
anotherfem
aleinland
fromJam
esB
ay.L
aterin1970, a
majorexpedition
todeterm
inethe
statusof
theJam
esIsland
tortoisew
asundertaken
(Villa
etal.,
1971).T
heseauthors
marked
atotal
of196
tortoises(129
males,
64fem
ales,and
3juveniles),
andre-encountered
allbut
29of
153anim
alsm
arkedon
aprevious
expedition.F
romthese
datatheyconcluded
thatthepopulation
couldnot
number
more
than500
animals.
Moreover,
inall
foursub-
regionsm
alesoutnum
beredfem
alesand
therew
asan
ex
treme
shortageof
juveniles.A
bundantnests
were
founddistributed
overfour
distinctnesting
zones,and
theco
nd
i
tionof the
eggsw
asgenerally
excellent. Butthe
presenceof
verylarge
numbers
offeral
pigsresulted
inthe
destructionofessentially
allhatchlings. This
observationis
ofinterest inthat, in
thenesting
zoneoflndefatigable
Island, feral pigsare
primarily
amenace
tothe
eggsratherthan
thehatchlings, and
akey
conservationm
easurehas
beento
erectheavy
rockcairs
overknow
nnests,
with
intersticessufficiently
wide
toallow
emerging
hatchlingsto
escape.V
illaet
al.(1971)
estimatedthatrecruitm
ent inthe
James
Islandtortoise
po
pu
lationhad
startedto
declineabout
50years
earlier,and
hadreached
zeroabout
20years
afterthat.
Villa’ s
partykilled
about1500
pigsduring
thefour
month
stay.but
theyesti
mated
thattw
icethat
number
survived.In
1982, Enriquez
(1983)spenttw
elvem
onthson
James
Island,and
observedthat
pigscontinued
tobe
athreat
totortoise
nests,and
alsothat
thenative
hawks
(especiallyabundant
onJam
es)occasionally
preyedupon
hatchlingtortoises. T
ortoisesw
ereconcentrated
inthe
highlandsfrom
Augustthrough
January, butfromM
archuntilm
id-May,the
adults(m
alesas
well
asfem
ales)w
ereconcentrated
inthe
lower,nesting
zones.(Thiscontrastsinterestingly
withP
orter’ s(1815)
observationin
thenesting
groundsof
northeasternJam
esIsland,
where
onlyfem
alesand
subadulttortoisesw
erefound;
thethree
males
capturedw
erefound
much
furtherintothe
interior).T
hreeshells
ofdead
tortoisesw
erefound.
Márquez
et al.(1992)
reportedthat
158young
tortoiseshad
beenrepatriated
fromeggs
collectedon
James
Island—
50from
1970—71,
16from
1971—72,
15from
1972—73,
20from
1973—74,
14from
1974—75,
12from
1975—76,
18from
1976—77,
10from
1981—82,
and3
from1982—
83.In
addi
tion,a
totalof
49tortoises
fromsubsequent
yearsw
erestill
incaptivity.
Albem
arle(Isabela):
Volcan
Wolf
—T
heearly
historyofthe
tortoisesofthis
precipitous,arid
mountain
isobscure.
The
accountsof
tortoisecollection
onA
lbemarle
bynine-
teenthcentury
whalers
compiled
byT
ownsend
(l925b)did
notdifferentiate
between
thevarious
Albernarle
popula
tions,and
itis
probablethat
them
ajorityw
erecollected
eitherin
thehum
idareas
inthe
south.w
iththeir
enormous
tortoisepopulations, or from
Volcan
Darw
in.with
theexcel-
lentanchorage
offeredby
Tagus
Cove.
The
lackof
fresh
Fig
ure
46.H
arshterrain
onthe
northernslopes
ofV
olcanW
olfnear
PuntaA
lbemarle,
Albem
arleIsland.
68C
FIoN
IAN
RESEARCH
MO
NO
GRA
PHS,
Num
ber1
—1996
water
onV
olcanW
olf preventedany
settlement or
agricu
l
tureand,
while
tortoisesare
foundquite
closeto
thesea
at
bothP
untaA
lbemarle
tothe
northand,
toa
lesserextent,
at
Bank’s
Bay
tothe
southwest.
theform
eris
anunstable
anchorageand
bothpresent
anextraordinarily
ruggedlit-
toral zonethat m
ustbetraversed
with
great difficultybefore
tortoiseareas
arereached
(Fig.46).
Thus,
itw
asnot
untilthe
twentieth
centurythat
the
Volcan
Wolf
tortoisesw
eredocum
ented.T
heym
aynever
havebeen
abundant—
therew
ereneither feral m
amm
alsnor
ahistory
of collectionby
man
when
Beck
collectedthe
type
specimen
in190
1and
fiveothers,
alsofor
Rothschild,
in
1902(R
othschild,1901,
1903).B
eckalso
collectedfor
Van
Denburgh
inA
pril1906.
Going
ashoreat
Bank’s
Bay,
this
partyfound
sevenindividuals
aftera
week’s
search,but
concludedthat,
inview
ofthe
dryw
eather,m
ostof
the
tortoiseshad
gonehigher.
None
were
foundin
acoastal
wooded
areanorth
ofB
ank’sB
ay.
Subsequent
surveydata
arefew
. Dow
ling(1962)
found
severalyoung
individualsat
analtitude
of
about800
feet,
andS
now(1964),
with
two
companions.
thundtw
elvein
a
singleday’s
search.They
were
foundat altitudes
of between
230and
600feet,
onebeing
aju
ven
ile(22.5
cm)
andthe
remainder
between
54and
83cm
incarapace
length.S
now
alsonoted
thepresence
of
feralcats.
Snow
(1964)also
comm
entedthat,
althoughthe
po
pu
lationprobably
numbered
many
hundredsof
individuals,
theyoccurred
soclose
tothe
seaat P
untaA
lbemarle
that this
was
oneof
thechief
areasfor
theillegal
collectionof
live
tortoisesfor
exportto
thecontinent.
This
supp
ositio
nis
supportedby
them
orphologyof
many
of theconfiscated
liveto
rtoises
of
unk
no
wn
orig
inhoused
atthe
Charles
Darw
inR
esearchS
tation; m
anyof
theseshow
thebroad,
reverted
anterio
rasp
ectch
aracteristico
fthe
Volcan
Wolf
tortoises.M
yow
nobservations
aresim
ilarto
thoseo
fS
now.
In
tripsof
afew
hourseach
toB
ank’s
Bay
in1971
andto
Punta
Albem
arlein
both1972
and1989, I found
small num
bersof
tortoiseson
eachoccasion.
The
sevenfound
nearB
ank’s
Bay
on2
1January
1971
were
allsm
allto
medium
insize
CA
BO
MA
RS
HA
LL
PTA
.A
LB
EM
AR
LE
RA
NC
AS
—.
,ç,/j
V
CA
BO
BE
RK
EL
EY1V
.EC
UA
DR
PUERTO
BR
AV
O)
BAN
KS
BAY
A”-
V.
DA
RW
IN
PTA
.T
PTA
.G
AR
CIA
TAG
US
BE:G
LECRA
TER
ISLA
FE
RN
AN
DIN
AC
EO
)Ic
A.
AL
F::o
ISTM
OPE
RR
YI.]
30
F’
BA
HIA
EL
IZA
BE
TH
PTA.
MO
RE
NO
A’
PTA
.SA
NJU
AN
PTA.
CR
ISTO
BA
LA
’/-‘
A.
LO
BE
RIA
RA
ZC
AL
ET
AIG
UA
NA
RPTO
VIL
LA
MIL
PTA
.E
SSEX
-
CA
BO
RO
SA
Map
10.Albem
arleIsland
(Isabela).L
ightershaded
areasare
thedistributions
ofv
ariou
spopulations
oftortoises,darker
shadedareas
arenesting
zones.Stripedarea
isthe
approximate
former distribution
oftheextinct populationreported
byD
eSola
(1930),flO
Wunder lava.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises69
(Pritchard,
1971b),and
includedsaddlebacked
asw
ellas
dome-shelled
individuals. At P
untaA
lbemarle,I only
foundone
in1989,
atan
altitudeof
over1000
feet.In
theearly
1970sI
alsofound
severalold
skeletonsof
animals
whose
plastrahad
clearlybeen
separatedby
human
action, sothese
tortoiseshave
beenutilized
bysettlers
forfood
untilquite
recenttim
es.S
omew
hatm
orerecently,
nofew
erthan
27carcasses
of female
tortoisesw
erediscovered
nearP
untaA
lbemarle
(Anonym
ous,1980).
They
hadpresum
ablybeen
slaughteredw
henthey
came
down
tothe
coastto
laytheir
eggs.F
rom1980
tothe
presenttim
e,V
olcanW
olfhas
beenlittle
visitedby
scientificparties,
althoughthere
hasbeen
increasingillegal
visitationand
camping
byfisherm
enand
especiallyby
seacucum
ber(pepino)
collectors.C
ayotand
Louis
(1995)reported
thatsix
fieldtrip
reportsw
ereon
filefor
thelast
fifteenyears
forboth
theP
iedrasB
lancasand
Puerto
Bravo
areas.In
fourof
thosefrom
Piedras
Blancas
slaughteredtortoises
were
reported,as
theyw
erein
allsix
reportsfrom
Puerto
Bravo.
The
totalsize
ofthe
Volcan
Wolf
tortoisepopulation
remains
conjectural.S
now’s
(1964)estim
ateof
many
hun-dreds
may
havebeen
toohigh
—he
assumed
thatthedensity
heobserved
inan
areaof
aboutI
sq. kmw
assustained
overa
verylarge
area, butthism
aynotbe
thecase.N
evertheless,the
higherregions
ofV
olcanW
olfare
almost
unexplored,and
althoughm
uchis
freshlava
orvery
steepterrain,
therem
aybe
many
tortoisesup
there,and
therelative
abundanceofjuvenile
specimens
at thelow
eraltitudes
isencouraging.
Occasionally,
eggshave
beencollected
andyoung
tortoiseshead-started
andrepatriated
14from
1969—70,
22from
1978—79,
and1
from1979—
80(M
árquezet
al.,1992).
Unfortunately,
significantslaughter
ofV
olcanW
olfto
rtoises
continuesup
tothe
presenttime.
Louis
(1994)reported
ona
visittoPiedras
Blancas
andL
osN
idos,on
thenorthern
sideof
Volcan
Wolf,
on11—
15A
ugust1994,
when
hefound
10live
and12
deadtortoises. H
ecom
mented
onthe
extremely
easyaccess
tothe
areaandthe
closeproxim
ityofillegalfishing
boats. Atanothersite, Puerto
Bravo, w
est-southwest ofV
olcanW
olf,18
liveand
12dead
tortoisesw
erefound.
Some
ofthe
carapacesw
erefound
atanillegal
camp
occupieda
coupleof
months
earlierby
seacucum
berfisherm
en.A
lbemarle
(Isabela):
Volcan
Darw
in.—
Th
ism
ountainprovided
thetortoises
that, forcenturies,have
beenrem
ovedthrough
theexcellent
deep-water
anchorageatT
agusC
ove.T
heintensity
ofth
iscollection
duringthe
nineteenthcentury
Fig
ure
47.V
iewof
Volcan
Darw
in(A
lbemarle
Island)from
theeastern
foot ofTagus
Cove
Mountain,February
1975.The
closestarea
offorest,beyondthe
lavaflow
, isan
importanttortoise
habitatand
breedingarea.
Fig
ure
48.V
olcanD
arwin
(Albem
arleIsland)
tortoiseresting
atthe
western
footof
them
ountain.N
otethe
landiguana
(Conolophus
subcristatus)living
inm
icrosympatry
with
thetortoise.
April
1972.
70C
HE
LO
NIA
NR
ES
EA
RC
HM
ON
OG
RA
PH
S,
Num
ber1
—1996
Fig
ure
50(u
pp
erright).Post-hatchling
tortoise(carapace
lengthca. 10
cm) entering
a shallowburrow
orretreat. inner slopeofthe
southernflank
ofVolcan
Alcedo.
April
1985.
Fig
ure
51(low
erright). A
dult female
tortoisesin
aw
ell-grazedarea
onthe
easternrim
ofV
olcanA
lcedo. August
1983.
isunclear,
inthat
thew
halers’records
donot
differentiatebetw
eenthe
differentparts
ofthe
verylarge
islandof
Albem
arle,butB
eck(1903)
was
probablycorrect
when
heobserved
thatthe
Volcan
Darw
intortoise
(“Testu
do
inicrophes”[sic
i)had
beenused
asfood
forw
halersm
orethan
anyother.
Beck
foundlarge
numbers
oftortoise
trails,indicating
aform
erpopulationofhundreds
oftortoises, but,he
reported,“today
it isa
hardm
atterto
findone.”
But
ona
latertrip
(March
—A
pril1906),
Beck’s
partycollected
13m
alesand
ayoung
female.H
isdesignation
ofthelocality
asT
agusC
oveis
confusing,in
thatthis
drowned
craterand
associatedcoastal hill
represent theaccess
routerather
thanthe
collectingsite. T
hetortoises
were
collectedat the
footofV
oicanD
arwin
itself,acrossa
zoneofrough
lavathatw
ouldbe
impassable
totortoises
(Figs.47—
48).O
ccasionallya
tortoiseis
foundnear
Tagus
Cove
itself,but
theredoes
notappear
tobe
apopulation
there,and
theorigin
ofsuch
individualsis
unclear.T
wentieth
centuryauthors
includingB
eebe(1924),
Eibl-E
ibesfeldt(l959),
andB
owm
an(1960)
foundno
live
tortoisesat
Tagus
Cove
atthe
western
footof
Volcan
Darw
in,alth
oug
hth
eirdisco
very
of
deadyoung
tor
tois
es
didin
dicate
that
repro
ductio
nw
asoccu
rring.
Snow
(I964)
climbed
toan
altitudeof3000
feetonw
esternV
olcanD
arwin,
findingfresh
tortoisedroppings
andtrails
butno
actualanim
als.B
utM
acFarland
etal.
(1974a)re
portedan
estimatedpopulation
of500—1000
animals. A
totalof
65tortoises
hadbeen
marked
inO
ctober1965,
andthe
sizeand
sexdistribution
ofthese
(6adult
males,
26adult
females,
2young
males,
1young
female,
8non-sexed,
22sm
all)w
asencouraging.
This
isan
aridarea,
andm
uchof
western
Volcan
Darw
inas
well
asm
ostof
thew
esternlow
landsare
unvegetatedfresh
lava,unsuitablefortortoises.
The
populationapparently
descendsto
asm
allarea
ofthe
western
lowlands
duringrainy
periods,but
otherwise
isdispersed
highup
onthe
mountain.
On
anascent
of
Volcan
Darw
inon
22January
1971
, when
itw
asvery
dry,Ireached
thelow
landtortoise
zonew
ithinan
hour,hut foundonly
olddroppings
andtw
ocrum
blingskeletons.
Asingle
juvenileto
rtoise
.abou
t40
cmin
length,w
asfound
halfw
ayup
the
Figure
49(left).
Adult
male
tortoisesin
anarea
ofthe
Volcan
Alcedo,
Albem
arleIsland,
calderarim
dominated
bytree
ferns(C
yatheas’eatherhyiana).
July1986.
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises71
Fig
ure
52.Adultfem
aletortoise
fromthe
Volcan
Alcedo
calderafloor,show
ingextensive
carapacialscarring,possiblyas
aresultofcontactw
ithhotvolcanic
ejecta.A
ugust1983.
mountain,
andfive
largerones,
includingan
adultfem
ale,w
erefound
closetogether
nearthe
calderarim
(Pritchard,
197ib).
Yet
ona
latertrip,
in1972,
Iencountered
17tortoises
ofa
wide
rangeof
sizesin
thesm
allvegetated
lowland
area.T
hispopulation
would
seemto
havegood
prospects,unless
theferal
catsbecom
ea
problemfor
thehatchlings.
But
itis
likelyto
belim
itedfrom
reachinghigh
densityby
thelow
rainfall,rugged
terrain,and
relativelym
odest
extentof
thearea
suitablefor
tortoises.L
ouis(1994)
visitedthe
Tagus
Cove
areafrom
16—18
August
1994.and
was
ableto
collectblood
samples
from25
livetortoises.
He
foundno
deadtortoises,
butdid
observethat
goats,previously
consideredto
berestricted
tothe
southernparts
ofIsabela,hadreached
thearea,the
rimof
Beagle
Crater
beingcarpeted
with
goatdroppings.
One
tortoisew
asfound
killedatU
rbinaB
ay(V
olcanD
arwin)
in1984
(Cayot
andL
ouis,1994).
Albem
arle(Isabela):
VolcanA
lcedo.—
The
nineteenthcentury
literaturem
akesno
mention
ofthe
tortoisesof
Volcan
Alcedo,
anditis
probablethatthey
occurredtoo
farinland
tobe
convenientvictualling
forw
halers.M
oreoverthe
shorelineon
theeastern
sideof
them
ountainoffers
nogood
anchorage.N
evertheless,V
anD
enburgh(1914)
reported
thatBeck’sparty
hadencountered
remains
ofacam
pon
theupper
easternslopes
ofA
lcedo,at
2200to
2500feet
elevation,w
herethe
remains
ofabout
70old
tortoisesk
eletons
indicatedthatthe
tortoiseshad
beensubjected
toheavy
exploitation.T
hecam
pw
aspresum
ablylong
abandonedand
thetortoise
shellsalready
verydeteriorated,
inthat
nocollectible
carapacesw
erefound.
But
theferal
burrosthat
hadbeen
usedto
transportkegs
oftortoise
oilto
thecoast
andthat
now,
asin
1905—06,
runfree
onthis
mountain
arepresum
ablya
permanent
legacyof
theoperation.
Fig
ure
53.Group
of subadulttortoisesin
apond
(sincedried
up)beside
thefum
arole,innerslope
ofsouthwestern
partofVolcan
Alcedo
caldera,Albem
arleIsland.M
arch1970.
72CHELONIAN
RESEARCHM
ONOGRAPHS,N
umber
1—
1996
Beck’s
partyfound
andcollected
asingle
livefem
aletortoise.
This
specimen
hadm
atureovaries
buthad
acara
pacelength
ofonly67.75
cm.Itw
asvery
steeplydom
edand
may
havebeen
slightlykyphotic
—certainly
itw
asan
unfortunatespecim
enon
which
tobase
conclusionsabout
them
orphologyof
thepopulation.
Beck’s
reportof
anessentially
vanishedtortoise
popu
lation,andgrass
sohigh
andthickthatascentto
therim
ofthecaldera
was
allbut im
possible,apparently
discouragedo
thers
fordecades
tocom
e.Butin
August
1963, M.
Castro
was
ableto
climb
Alcedo
fromthe
northeast,reaching
therim
andspending
two
nightshigh
upon
them
ountain(S
now,
1964).H
elocated
anapparent
tortoisenesting
area5
kmnorth
ofthe
craterrim
andfound
many
oldtortoise
drop-pings,
butsaw
nolive
animals.
The
following
year,in
February
1964,m
embers
ofthe
Galapagos
InternationalS
cientificP
roject,flyingby
helicopteroverthevolcano,saw
many
tortoisesfeeding
inthe
grassyareas
ofthe
calderafloor.F
inally,Eric
Shipton
(therenow
nedH
imalayan
moun
taineer)reached
thisarea
byfoot
in1965,
andw
asable
toconfirm
theexistence
ofavery
largetortoise
population(de
Vries,
1984).In
thefollow
ingyears,
aroute
was
developedw
herebythe
craterrim
andfloor
couldbe
reachedw
ithoutgreat
difficulty,andsubsequently,num
erousparties
evenrela
livelylarge
groupsoftourists
haveascended
Alcedo
and
observedthe
largesttortoise
populationin
theG
alapagostoday,
andthe
onlyone
ina
semblance
ofprim
ordialab
un
dance(Figs.
5,49—
55).T
hepopulation
was
estimated
at3000—
5000individuals
byM
acFarland
etal.
(1974),al
thoughF
owler
(1983)observed
thatthem
ethodologyused
bythese
authorsw
asunstated,and
thattheactualpopulation
was
probablycloser
tothe
lower
figure.V
ariousaspects
ofthis
populationhave
beenstudiedin
depthby
thosew
hohave
managed
tosolve
thew
atersupply
problem;
Fow
ler(1983),
forexam
ple.cam
pedin
thecrater
forover
ayear
while
studyingthe
feedingecology
ofthe
tortoisesand
theferal
bu
rros.
Today
thetortoise
populationis
certainlylarge
andincludes
asatisfactory
proportionofyoung
animals
(indicat
inggood
reproduction)interm
ingledw
ithsom
eofth
elarg
esttortoises
inthe
world
today(indicating
goodadult
Ion-gevity).
Hum
anexploitation
hasnot
beensignificant
sincethe
earlypart
ofthis
century.C
ayotand
Louis
(1994)reported
thatonlythree
slaughteredtortoises
were
foundon
Alcedo
duringthe
1970s,and
thesew
ereestim
atedto
havebeen
killedduring
thepreceding
decade.N
evertheless,the
presenceofferal
burros,cats,
andrats
underscoresthe
needfor
carefulm
onitoring.T
heim
pactof
thefirst
ofthese
hasbeen
studiedby
Fow
ler(1983), F
owler
andR
oe(1984),and
Fow
lerand
Johnson( 1985).
These
studiesshow
that18.2%
of88
nestsin
thecaldera
floorw
eredisturbed
by
Figure
54.Tortoises
inthe
mist. C
oncentrationofm
ostlyadulttortoises
ina
cloud-catchmentdrip
zoneon
southernrim
ofVolcan
Alcedo.
July1986.
Also
seefigure
andpoem
inthe
editorialprologue.
PRITCHARD—
The
Galapagos
Tortoises
73
thetram
plingof
burros.T
hepercentage
was
higheron
thesouthern
nestingsite
(32.1%)
thanon
thenorthern
( I I.7%).
Inaddition
tothese
problems, in
recent yearsgoats
havesucceeded
incrossing
Perry
Isthmus
andare
noww
ide-spread
onA
lcedo.T
hethreat
thattheypose
tothe
tortoisesw
asgraphically
describedby
Merlen
(1992). Mufloz
(1993)reported
thatgoatshad
colonizedA
lcedow
ithinthe
lastfouryears.
Astudy
was
conductedin
1991to
determine
theircurrent
distribution,population
structure,hom
erange,
reproductive
cycles,and
feedingbehavior.
The
goatsw
erefound
onthe
southern,eastern,
andsouthw
esternflanks
ofthe
volcano,w
iththree
groupsaveraging
18.2individuals
eachw
ithinthe
calderaitself.
Seasonal
movem
entsw
erefound,
asthe
goatsm
igratednorthw
ardsor
soughtfood
orw
aterduring
thegaru
aseason.B
y1995,the
goat populationhad
increasedcatastrophically,
to20—
30,000individuals,
andthe
tortoisepopulation
mustnow
belisted
asthreatened
(L.
Cayot,
pers.com
m.).
Volcan
Alcedo
isnow
closedto
tourism,
andthe
Charles
Darw
inR
esearchS
tationand
theG
alapagosN
ationalP
arkS
ervicenow
plana
major
goatkilling
campaign
(Cayot
andSnell,
1996b).V
olcanA
lcedohas
permanent
hotfum
aroles,sulfur
vents,and
otherhot
spots,and
thesem
ayoccasionally
killtortoises.
Individualsare
sometim
esseen
with
heavyscar-
ringof
thedorsal
scutes(Fig.
52),possibly
asa
resultof
exposureto
volcanicm
aterials.H
ayesand
Beam
an(1985)
founda
deformed
adult female
tortoisethat,they
speculated,m
ayhave
beena
survivorof
anearly
burroinjury,
althoughother
causesofthe
deformity, including
geneticor
teratoge
nicones,
were
alsoconsidered.
Itisim
portant tonote
that, althoughvery
highdensities
of tortoisescan
beseen
onA
lcedo, theactual
areaof suitable
tortoisehabitat
isnotparticularly
extensive.T
ortoisesm
aybe
foundon
most
ofthe
rimof
thecaldera
(anessentially
lineardistribution)
andare
foundin
scatteredfashion
onthe
uppereastern
flanksofthe
volcano.B
utmuch
ofthecaldera
flooris
precipitouslava,the
lower
easternslopes
arelargely
almost
unvegetatedpum
ice,and
theentire
western
sideof
thevolcano
isdensely
vegetated,interlaced
with
vertical-sided
ravines,and
hasvery
fewtortoises.
The
highestnu
mbers
arefound
inthe
fiator rolling,grassyorlightly
wooded
areasbelow
thefum
aroleon
thesouthern
sideofthe
calderafloor (Fig.53). P
opulationm
ovements
areextensive. D
uringdry
times, the
tortoisesconcentrate
alongareas
ofthecaldera
rimw
herem
oisturefrom
passingclouds
isentrapped
bylow
treesand
forms
heavym
orningdew
oreven
puddles(Figs.
54—55;
seealso
figureand
poemin
theeditorial
prologue).B
utduring
wetter
times,
thetortoises
migrate
tolow
eraltitudes,
many
ofthem
concentratingin
lushgrassy
mead
ows
onthe
calderafloor,
where
seasonalpools
may
attractparticulary
largenum
bers.
Fig
ure
55. More
tortoisesin
them
ist.A
groupof m
ostlyadulttortoises
onthe
rimofV
olcanA
lcedoA
ugust1986.
74C
HE
LO
NIA
NR
ES
EA
RC
HM
ON
OG
RA
PH
S,
Num
ber1
—1996
Albem
arle(Isabela):
Sierra
J\/egraand
Cerro
Azul.
—
Originally,
thisarea
ofsouthern
Albem
arlem
usthave
har
boredenorm
ouspopulations
oftortoises,
judgingby
thegreat
areasof
what
appearto
thisday
tobe
suitablehabitat.
But the
populationsare
severelyreduced
andtortoises
havebeen
totallyextirpated
fromvast
areason
thesouthern
uplandsof
Sierra
Negra
where
theyonce
abounded.T
heim
pactof
thew
halerson
thisarea
issom
ewhat
conjectural,in
thatfew
ofthe
whalers
specifiedthe
partof
Albem
arlefrom
which
theirtortoises
were
obtained(see
backcover
engraving,w
ithfigure
captionon
p.4).
Insom
ecases
thereports
indicatenot the
main
islandof
Albem
arle,but
satelliteislets
fartoo
small
tohave
tortoisepopulations
(e.g.,
Cow
leyIslet,
orC
rossman).
Itcan
probablybe
as-sum
edthat
most
ofthe
2493tortoises
collectedfrom
Albem
arleby
whalers
whose
logbooksw
ereexam
inedby
Tow
nsend(1925b)
were
collectedeither
fromT
agusC
ovein
thenorth, orfrom
Villam
ilorIguanaC
oveinthe
south;butthe
actualdistribution
ofthe
collectingeffort
will
neverbe
known
forsure.
The
tortoisesw
erestill
abundantin1902
(Fig.56), but
theirprospectsw
erebleak,w
henB
eckspent tw
ow
eeksata
newly-established
ranchten
miles
fromthe
shoreon
south-em
Albem
arle, presumably
onS
ierraN
egra.C
attleroam
edthe
mountain
ingreat num
bers,as
didferal
dogs,and
while
theform
erm
aynothave
presentedserious
problems
tothe
tortoises,thedogs
were
reportedto
eat youngtortoises
asfast
asthey
were
hatched,and
Beck
doubtedif
more
thanone
inI0,000
escaped.T
hedogs
evenkilled
adulttortoises,
primarily
thesm
allerfem
ales,but
occasionallyeven
largem
ales.A
sbad
asthe
dogs, orevenw
orse, were
the50
orsofield
handsem
ployedon
theranch,
who
killedtortoises
notonly
forfood,butalso
fortheiroil
—a
valuableexport co
mm
od
ity.A
lthoughthe
proprietorof
theranch
claimed
thatonly
male
tortoisesw
erekilled,
infact
females
asw
ellas
males
were
slaughteredw
ithoutrestraint,
andsom
etimes
onlya
fewpounds
ofm
eatw
asrem
ovedfrom
anadult
tortoise.B
eckfoundabout150
tortoiseskeletons
at asingle
waterhole
(Fig.57),
andabout
100at
another.H
efound
7tortoises
ina
pondI0
miles
fromthe
ranch(Fig.56), but feared
thattheyw
eredoom
edw
ithinthe
coming
months.
On
areturn
triponly
fouryears
later, Beck
was
ableto
reviewthe
impact
ofthe
massacre
(Fig.57).
He
was
ableto
collectabout
40tortoises,
butnearly
allw
erefem
ales,and
only2
males
more
than28
inchesin
carapacelength
were
included.H
undredsof
tortoisebones
were
foundalong
thetrails
andatthe
water holes. T
henatives
saidthat all tortoises
hadbeen
killedoffbelow
thesettlem
entand
that theyw
ereno
longerabundant
anywhere
onthe
mountain.
Settlem
entson
southernA
lbemarle
havecontinued
throughthis
century,including
am
ajorpenal
colonyin
theinterior
from1946—
59,a
military
presenceduring
WW
II,and
theexisting
coastalvillageofV
illamil,founded
in1895,
connectedto
thehighland
comm
unityof
Santo
Tom
ásby
foottrail.
In1957,
youngtortoises
couldstill
bepurchased
fromV
illamil
residents(E
ibl-Eibesfeldt,
1959).E
xp
loita
tionof
thetortoises
bysettlers
continuedinto
the1960s
Map
11.South
ernA
lbemarle
Island(Isabela):
Sierra
Negra
and
Cerro
Azul.N
amed
tortoiseregions
designated.Lightershaded
areasare
thedistributions
ofvarious
populationsof
tortoises,darker
shadedareas
arenesting
zones(galapagueras).
Stripedarea
isthe
approximate
former
distributionof
theextinctpopulation
reportedby
De
Sola(1
930),now
underlava.
I
-
:t1/
ISAN
TOTO
MA
SI
\‘
IGU
AN
A
CA
BO
SAN
PED
RO
_________
PR
ITC
HA
RD
The
Galapagos
Tortoises
75
(Snow
,1964),
andeven
later.M
acFarland
etal.
(1974a)reported
thatsettlers
stillkilled
tortoiseson
Sierra
Negra,
andthat,
asrecently
as197
1,
remains
of60
slaughteredtortoises
hadbeen
foundin
anarea
of0.9
sq.km
,although
only8
appearedto
havebeen
slaughteredrecently,
therem
ainder2—
Syears
earlier.M
acFarland
etal.
(1974a)also
documented
thevirtual
extirpationof
two
ofthe
pocketsof
tortoiseabundance
onsouthern
Albem
arlethathad
survivedthe
onslaughtsofthe
settlersand
oilhunters.
Tortoises
hadbeen
almost
com
pletelyelim
inatedfrom
120sq. km
ofthe
uplandsof
Sierra
Negra
byoilhunters
fromthe
prisoncolony
inthe
1940sand
SOs,
with
operationsextending
evento
Cerro
Azul.
And,
between
1959to
1969,tortoises
were
decimated
inan
areaof
about5
sq.km
inlandfrom
IguanaC
ove,in
theextrem
esouthw
est,byem
ployeesofcattle
companies.T
helatterarea
hadpreviously
beenan
important
tortoisearea,
andhad
providedthe
vastm
ajorityof
the180
mostly
juvenileto
rtoises
collectedby
Tow
nsend’sparty
duringthree
weeks
in1931.H
owever,L
.C
ayot(pers.
comm
.)reports
thatIguanaC
ovestill
hada
goodtortoise
populationin
1995.T
ortoisessurvive
inlowdensity
inthree
metapopulations
onsouthern
Albem
arle.A
nestim
ated300
individualssu
rvive
ina
dryarea
nearC
erroG
rande,east
ofthe
calderaof
Sierra
Negra
andinland
fromC
erroB
allena.M
osttortoisesfound
hereare
young,suggesting
goodnaturalreproduction
andrecruitm
ent.V
illa(1972)
reportedthat
217in
div
idu
alsof
thispopulation
hadbeen
marked.
This
colonyis
widely
separatedby
settlements
andcultivation
froma
lowland
southernpopulation
ofperhaps
200individuals
inlandfrom
Cabo
Rosa.
And
aroundC
erroA
zul,to
rtoises
may
befound
fromlow
altitudes(the
humid
zoneextending
down
tosea
levelon
thism
ountain)allthe
way
tothe
calderarim
.T
hispopulation
isestim
atedat
500—700
individuals,of
which
203had
beenm
arkedby
1971(V
illa,1972).
Afew
individualsalso
surviveon
theeastern
sideof
along
lavaflow
eastof
Cerro
Azul
(Fig.
58).T
hisflow
reportedlyoccurred
in1926
(J.G
ordillo,pers.
comm
.),nearly
reachingthe
sea,and
dividingthe
Cerro
Azul
tortoisepopulation.
The
Cerro
Azul
populationutilizes
fourw
ell-definednesting
zones:Las
Tablas
(westofC
erroA
zul),Los
Crateres
orP
egas(east
ofC
erroA
zul),P
lanchonadade
losC
incoC
erros(w
est ofSanP
edro),andL
osG
avilanesnorthw
estofC
erroA
zul(Villa,
1972).Inmostoftheselocalities,pressure
fromferal
mam
mals
(pigs,dogs,
andpossibly
cats)w
asintense,
pigsraiding
thenests
anddogs
killingyoung
andeven
half-grown
tortoisesup
to55
cmin
curvedcarapace
Fig
ure
56.Scene
onsouthern
uplandsof
SierraN
egra,A
lbemarle
Island,in
1901
.Tortoises
onthis
mountain
arenow
veryfew
anddispersed.Photo
byR
H.B
eck,reprintedfrom
Beck
(1903)and
Slevin(1959).T
he1959
captionread:
“Atypicalw
aterholescene
of1901
ontop
ofVillam
ilMountain,A
lbemarle
Island,thesite
ofprobablythe
largesttortoisecolony
onthe
islandsuntilthe
oilhuntersbegan
operations.”
76C
HELO
NIA
NRESEA
RCEEM
ON
OG
RA
PHS,
Num
berI
—1996
length.F
eralcattle
were
alsoabundant
andconstituted
an
important
foodsource
forthe
dogs.C
onservationefforts
inrecent
yearshave
concentrated
oneducating
thelocal
population,field
patrolsand
law
enforcement,control offeraldogs
bypoisoning
(Moore
and
Moore,
I982),protectionofnatural
nestsfrom
predationby
erectingrock
cairns, andsporadic
head-startingofeggs
from
wild
nests.M
árquezet
al.(1991)
reported25
tortoises
repatriatedto
Cerro
Azul
from1971—
72;25
from1972—
73;14
from1973—
74;37
from1975—
76;17
from1977—
78;
16from
1978—79;
17from
1979—1980;
14from
1980—
81;
14from
1981—82;
and3
from1986—
87;and
33m
ore
were
stillheld
incaptivity.
On
Sierra
Negra,
only13
youngtortoises
hadbeen
repatriatedfrom
artificially
incubatedeggs
laidbetw
een1977
and1982,
although53
animals
from1983—
84and
subsequentseasons
were
still
heldin
captivity.T
hew
idehiatus
between
thehighland
tortoisepopula
tionssoutheast
ofS
ierraN
egraand
eastof
Cerro
Azul
is
anthropogenicin
nature,theresultof the
intensiveslaughter
oftortoises
andconversion
ofnative
vegetationto
pasturelandsas
describedby
Beck
(1903).O
nthe
other
hand,recent
maps
ofthe
geologyof
southernA
lbemarle
showvirtually
continuouslava
flows
overthe
entirenorth-
emhalf
ofthisenorm
ousblock
ofland,
reachingfrom
(and
joining)the
two
calderas,and
extendingnorth
toengulf
Perry
Isthmus.
No
tortoisescould
survivein
thislava
field
today,buttherew
asa
populouscolony
inthe
lowlands
ofthe
easternside
ofP
erryIsthm
usuntil
comparatively
recent
times.
To
wn
send
(1925)reported
thata
tortoiseof
extrao
r
dinarysize,
famous
among
visitingw
halers,lived
inthis
area,an
dw
asseen
byG
eorgeA
.Grant ofN
antucketinI8 8
1.
The
animalw
asknow
nas
PortR
oyalTom
,andhad
datesand
nam
escarv
edin
toits
carapace,
the
earliestd
atebein
g179
1.
Itw
asin
thissam
earea
thatD
eS
ola(1930)
observed
numerous
tortoises,som
eof
which
hecollected
alive,and
which
henam
edT
estudovandenburghi.
McB
irneyand
William
s(1969)
reportedthree
lavaflow
ssince
1930from
ven
tsn
ot
farfro
mthe
north
ernrim
oftheSierra
Negra
caldera,
in1948.
1953—54,
and1963—
64.T
helast
flowed
primarily
towards
Punta
Moreno
andthe
western
sideofP
erryIsthm
us,
buteither
ofthe
two
earlierones
may
havedestroyed
the
torto
isehab
itaton
the
easternside
of
theIsth
mu
s.
Anew
threatflaredup
inF
ebruary1985,
when
agreat
fire,apparently
startedby
settlers,burned
alarge
areaof
southernA
lbemarle.
Tortoises
were
roundedup
andp
repa
rationsw
erem
adeto
evacuatethem
byhelicopter,
although
thistum
edouttobeunnecessary.A
ninternationalfirefighting
effortw
asm
ountedand
eventuallya
firebreakline
40km
longw
ascleared.
Tortoises
were
regularlyencountered I
PR
ITC
HA
RD
—T
heG
alapagosT
ortoises77
alongthis
firebreak,but nonew
asreported
killedby
thefire,
although175
sq. kmof
wilderness
was
devastated.and
two
subadultsdied
fromthe
extreme
heatw
hilebeing
carriedto
acorral
preparedfor
themin
acoastal
areanear
SanP
edro.T
hefire
was
notfinallyextinguished
untilrainscam
ein
July(M
árquez,1986;
Now
akand
Law
esson,1988).A
nu
nfo
rtunate
side-effectof thisfire
was
thenew
accessitcreated
forpoachers
toreach,
andto
kill,tortoises
(Waliner,
1995).A
notherw
asthat
anaggressive
exoticplant,
theguayabo,
rapidlycolonized
areasw
herenative
vegetationhad
beendestroyed.
Because
oftheir
severelyreduced
populationsand
ongoingstresses,
tortoisesfrom
theC
erroP
aloma
andC
azuelapopulations
(west
andeast
ofthe
Sierra
Negra
caldera,respectively),were
transferredto
acaptive
breed-ing
facility,or
crianza,
atV
illamil.
Currently,
oneadult
female,
fiveadult
males,
andfive
juvenilesfrom
Cerro
Palom
areside
atthisfacility,
asw
ellas
approximately
75tortoises
fromC
azuela(only
15adults).
There
arestill
fiveadult
males
(noknow
nadult
females)
inthe
wild
atC
erroP
aloma,
anda
feww
ildtortoises
atC
azuela(L
.C
ayot,pers.
comm
.; Cayot
andS
nell,1996a).
The
most
recentsum
mary
ofthe
statusof
southernA
lbemarle
tortoisepopulations
isthatofC
ayotandM
árquez(1994).T
heseauthors
estimated
that therew
ere1000—
2000tortoises
onC
erroA
zul,with
subpopulationsof
100—300
atL
asT
ablas,500-1000
atL
osC
rateres,less
than100
atL
os
Gavilanes,
and100—
300at
Cinco
Cerros.
Tortoises
were
considerablyscarcer(less
than500)
onS
ierraN
egra, with
anestim
ated100—
200at
Cabo
Rosa,
lessthan
100at
Roca
Union,
lessthan
20at
Ceno
Palom
a,and
lessthan
100at
Cazuela.
Recent
reportsindicate
thatproblem
sfor
thetortoise
populationsof
southernIsabela
arefar
fromover.
Asecond
fire,apparently
startedw
hena
camp
cookingfire
was
incompletely
extinguished,occurred
inA
pril1994
inthe
southwestern
sectorof
Sierra
Negra
(Cruz
et al.,1994). T
hefire
was
notnearlyas
disastrousas
the1985
fire,butit
was
notextinguishedfor
almosttw
om
onths.Furtherm
ore,Cruz
Márquez
andcolleagues
were
ableto
travelwith
theh
elicop
terassigned
toassist
with
firecontrol,
andalthough
theyfound
notortoises
killedby
firein
thenearby
Roca
Union
subpopulation,they
foundthe
remains
ofeight
tortoises(3
males,
3fem
ales,2
juveniles)thathad
clearlybeen
killedby
man
fortheir
meat (M
árquezetal.,
1994).AtC
erroP
aloma,
remains
offurther
tortoisesw
erefound,
buttheseappeared
tohave
beenkilled
before1988.
Southern
Albem
arleis
anenorm
ousarea,conservation
staff arefew
,comprehensive
patrolsare
almostim
possible,and
theE
cuadoriansettlers
areresentful
ofincreasingly
restrictiveconservation
regulations.In
1986,theG
alapagosM
arineR
esourcesR
eservew
asestablished,
andin
August
1992a
marine
managem
entplan
forG
alapagosw
atersw
assigned
intolaw
(Cam
hi,1994).
The
regulationsw
erede
L.
‘
Figure
58.Adultm
alesouthern
Albem
arletortoise,east
ofCerro
Azul.M
arch1971.
78CHELONIAN
RESEARCHM
ONOGRAPHS, Num
ber1
—1996
signedto
restrictforeign,
industrialscale
fisheriesin
Galapagos
waters
but tom
akeconcessions
tothe
economic
needsof local
fishers. Nevertheless,
thesettlers
onsouthern
Albem
arlehave
beenvociferous
intheir
objectionsto
thefishery
regulationsand,
onand
offsince
July1993,
theyhave
protestedand
picketedthe
Charles
Darw
inR
e-search
Station,
threateningto
killgiant
tortoises(even
Lonesom
eG
eorge,the
lastsurvivor
ofthe
Abingdon
Islandpopulation),
andto
introducenon-native
speciesto
thefew
remaining
pristineislands.
InS
eptember
1995,the
Park
Headquarters
andthe
Darw
inS
tationw
ereoccupied
fortw
ow
eeksby
localprotesters
(Lem
onick,1995;
Merlen,
1995).S
laughteredtortoises
were
indeedfound,
althoughin
placesw
herethere
isno
certaintythatthe
slaughter hadbeen
doneby
thesam
eindividuals
who
hadm
adethe
threats,or
evenby
fishermen
atall.
Over
80tortoise
mortalities
were
confirmed
onA
lbemarle
alonein
1994(C
amhi,
1994; Cayot
andL
ouis,1994).
The
government
capitulated,rescinding
theban
onlarge-scale
comm
ercialfishing
ofsharks,
seacucum
bers,lobsters,
andgroupers.
Louis
(1994)docu
mented
anum
berof
thetortoise
mortalities
with
photo-graphs,
andreported
onthe
tortoiseshe
foundon
south-em
Albem
arlebetw
een20
Julyand
30A
ugust1994,
asfollow
s:1)
Roca
Union.
22July.
Tw
enty-onelive
tortoisesfound,
asw
ellas
7previously
reportedcarcasses
and5
unreportedones. P
lastronhad
beenrem
ovedby
choppingin
all12
cases.2)
Cazuela.
23July.
Only
onecarapace
found,but
approximately
39dead
tortoiseshad
beenfound
herein
March
1994.3)
Cinco
Cerros.
24—25
July.F
ourteenlive
tortoisesfound.
Later
(22—26
August)
inthe
same
area,7
deadtortoises
foundclose
tothe
beach,and
4m
oreinland.
In8
of
thesecases, the
tortoiseshad
beenbutchered.
Forty-six
livetortoises
foundon
thesecond
visit.4)
Las
Tablas.
28—30
July.T
wenty-five
livetortoises
found,all
females
andjuveniles.
5)C
erroP
aloma,
Los
Crateres,
andL
osG
avilanes.1—
6A
ugust.Five
adult males
foundat C
erroP
aloma, and
aboutfour
deadtortoises.
Tw
enty-fivelive
tortoisesfound
at Los
Crateres,
and8
atL
osG
avilanes.N
amed
tortoiseareas
orgalapagueras
onsouthern
Albem
arleare
locatedas
follows,
inrelationship
tothe
two
calderas(C
erroA
zulinthe
west and
SierraN
egrain
theeast):
1.L
aC
azuelais
east of Sierra
Negra, tow
ardsthe
coastalhillC
erroB
allena, andis
surroundedon
allsidesbutthe
west
bylava
flows.
2. Los
Gavilanes
isnorth
andnorthw
est ofC
erroA
zul.3.
Los
Crateres
iseast
andnortheast
ofC
erroA
zul.4.
Cerro
Palom
ais
midw
aybetw
eenthe
calderasof
Cerro
Azul
andS
ierraN
egra, just eastofthe
longtongue
ofunvegetated
lavaleft
bya
1926eruption.
5.L
asT
ablasis
southwest
ofC
erroA
zul,extending
fromthe
calderato
thesea,
andincluding
IguanaC
oveand
Cerro
Manzanilla.
6.C
incoC
errosis
locatedsouthw
estof
Cerro
Azul,
extendingfrom
thecaldera
tothe
sea, andencom
passingthe
coastal areasbetw
eenC
aboS
anP
edroand
thelava
flowdue
southof
Cerro
Azul.
The
Cinco
Cerros
(=five
hills)are
isolatedcones
arrangedin
anorthw
est-southeast radial fashion
between
thecaldera
andC
aboSan
Pedro.
7.C
aboR
osais
locatedon
thesouthern
coast ofsouth-
emA
lbemarle,
inthe
lowlands
oppositethe
saddlebetw
eenC
erroA
zuland
Sieffa
Negra.
8.R
ocaU
nionis
arock
islandin
theocean
dueeast
ofC
aboR
osaand
duesouth
ofS
antoT
omás;
inthe
aboveaccounts,
thenam
erefers
tothe
lowlands
ofsouthern
Albem
arledirectly
facingR
ocaU
nion.
CO
NC
LU
SIO
NS
Neither
thenornenclatural
northe
survivalstatus
ofG
alapagostortoises
canbe
consideredsettled.
Norn
enclatu
ralS
tatus.—
Inorder
tostabilize
thenom
enclature,it
isnecessary
tom
akepetition
tothe
ICZ
Nto
suppressseveral
names
thatcould,
underpre
vi
0U
5editions
ofthe
Code
ofN
omenclature,
havebeen
declared
no
min
aoblita
(e.g., T
estudocalifo
rnian
aQ
uoyand
Gaim
ard,1824a),
orthathave
beenw
idelyused
inan
incorrectcontext
(e.g.,
Testudo
eph
ipp
ium
GU
nther,I875), or
whose
typelocalities,
andthus
whose
islandsof
origin,are
uncertain(e.g.,
Testudo
nig
ritaD
uméril
andB
ibron,1835).
Som
enew
subspeciescould
potentiallybe
described(e.g.
, onnorthw
esternIndefatigable
Island,or
northeasternC
hathamIsland),
hutw
illrequire
typem
aterial(presum
ablysalvaged)
beforenam
escan
beproposed
anddescriptions
drawn
up.F
urthermore.
afew
recomm
endationscan
bem
adeto
developa
degreeof
sophisticationin
Galapagos
tor
toisenom
enclaturethat
isH
OW
lacking:1)
Morphological
comparisons
needto
bem
adebetw
eenthe
abundantm
useumm
aterialof
Galapagos
tortoises,collected
inthe
nineteenthand
earlytw
entiethcentury,
andthe
current(living)
generationof
tortoisesO
ilall
islandsw
herethey
stillsurvive.
This
will
generateinvaluable
insightinto
thepossibility
ofm
icroevolutionoccurring
evenw
ithina
fewgenerations,
asoccurs
with
thegeospizine
finchesin
thearchipelago.
2)G
alapagostortoises
arelong-lived
butthey
arenot
imm
ortal,and
opportunitiesshould
notbe
lostfor
collectingany
skeletalm
aterialin
reasonablecondition
oftortoises
thathave
died(or
beenkilled)
inthe
wild.
Using
Global
Positioning
System
s.the
localitydata
of
suchspecim
enscan
herecorded
with
precision.C
om
pre
hensiveosteological
collectionsw
illnot
onlyfacilitate
thefirstrecom
mendation
above,hut
will
counterbalancethe
strongbias
towards
entire,m
ountedspecim
ensin
extantm
useumcollections.
3)T
heholotypes
ofthe
severalcurrently
indeterm
inate
“species”ofG
alapagostortoise
(T.
calijorn
iana
andT.
nig
ra,T.
elephan
topus,
T.n
igrita,
T.p
lanicep
s,T
i’nicrophyes,T
wallacel,
T.cliio
sa,and
T.ty
pica)
needto
here-exam
ined.In
thosecases
where
driedsoft
tissuesare
stillpresent
onthe
specimen,
evenin
minuscule
amounts,
itis
possiblethat
mitochondrial
DN
Aanalysis
andappropriate
comparisons
with
specimens
ofknow
norigin
may
allowthese
names
tobe
allocatedd
efini
tively.G
eneticstudies
onall
extantpopulations
are
currentlybeing
conductedby
Edw
ardL
ouis,and
theresults
ofthese
may
well
alsohave
aninfluence
uponnom
enclature.S
urv
ival
Statu
s.—
Eibl-E
ibesfeldt( 1959)
notedthat
about2000
peoplelived
inthe
Galapagos
Islands,and
concluded:“O
nthe
whole
colonizationconflicts
with
natureprotection.
How
canboth
interestsbe
reconciledand
what
arethe
prospects?”T
oday,alm
ost40
yearslater,
we
would
haveto
agreew
iththe
statement,
andconfess
thatw
ehave
nodefinitive
answers,
buta
gen
erally
pessimistic
response,to
thequestions.
For
aw
hilethings
lookedbetter,
buttoday
thereis
much
bleaknew
sto
reportabout
thesurvival
prospectsof
thevarious
populationsof
tortoisesin
theG
alapagos.D
espite35
yearsof
hard,dedicated
work
bythe
staffof
theC
harlesD
arwin
Research
Station
andthe
Galapagos
National
Park
Service,
newproblem
skeep
arising,and
feralm
amm
alscontinue
toreach,
orbe
takento,
newareas.
Tourism
,initially
thecash
cowthat,
throughtour-
isttaxes,financed
and
justified
theN
ationalPark
andthe
variousconservation
programs
inthe
archipelago,nowis
revealingits
disadvantages.F
lightsto
theislands
nowoccur
almost
daily,and
money
ism
adeby
supplyingtourists
with
accomm
odations,transport,
guides,m
eals,and
souvenirs.N
oone
starvesin
theG
alapagosnow
adays,
andthe
archipelago,despite
itsrugged
terrainand
reputationas
aplace
ofshipw
reckand
hardshipat
leastin
former
years,now
hasthe
highestoverall
standardof
livingof
anyprovince
ofE
cuador.T
hishas
prompted
massive
increasesin
thehum
anpopulation
ofthe
islands.U
nderthe
Constitution
ofthe
Republic
ofE
cuador,this
migration
cannotlegally
bestopped.
Yet
many
ofthe
imm
igrantsfind
thatthey
cannotlive
directlyfrom
thetourist
trade,and
areforced
toextracta
livingfrom
theG
alapagosenvironm
entitself,by
agricultureor
fishing,tosurvive.
lnevitablythis
leadsto
conflictsin
anarea
where
95%of
theland
surfaceis
National
Park,
andw
herethere
isalso
aM
arineC
on
servation
Zone
aroundthe
islands.O
flate
theseconflicts
havebecom
eso
severethat
thevery
futureof
theG
alapagos,its
ecosystems,
andits
endemic
species,is
atstake.T
oday,the
habitatof
thetortoises
onIndefatigable
overlapssubstantially
with
privateagricultural property,
andstray
dogsand
feralpigsw
anderthrough
theT
ortoiseR
eserve.O
nD
uncanIsland,
thetortoises
havepossibly
beensaved
throughthe
head-startingprogram
,but
rats
CE
IoN
1N
RESEARCH
MO
NO
GRA
PHS,
Num
ber1
—1996
stillexist,
andit
isfar
tooearly
todeclare
avictory
andretreat.
On
Aihem
arle,tortoises
arethreatened
bydirect
human
actionon
allthe
volcanoesexceptA
lcedo.A
lcedoitself,
home
tothe
largesttortoise
populationin
thearchipelago,
seemed
secureuntil
recently;but
now,
tensof
thousandsof
goatshave
join
edthe
long-establishedferal
burros,and
thevegetative
coverof
them
ountainw
illbecleared
within
afew
yearsunless
anex
traord
inar
ilyam
bitiousgoat
controlprogram
ism
ounted.T
hetortoises
ofH
oodIsland
may
survive.Indeed,
itis
vastly
enco
urag
ing
tosee
ap
op
ulatio
no
fsev
eralhundred
tortoisesw
herenone
existeda
fewyears
ago.B
utone
cannotignore
thepossibility
ofinadequate
geneticdiversity
ina
populationderived
fromju
stthree
males
anda
do
zenfem
ales.T
orto
iseson
James
Island
stillhave
tocom
petew
ithtens
ofthousands
ofgoats
andtheir
eggsand
youngare
stillexposed
todepredation
bypigs.
Perh
aps
onC
hath
am,
nowth
atthe
feraldogs
hav
e
beencontrolled,
thetortoises
may
besafe.
But
onecan
neverrelax
andunrem
ittingvigilance
will
henecessary.
Major
neweducational program
sw
illhaveto
belaunched,
forboth
theyoung
peopleand
theadults
ofth
eG
alapagosIslan
ds.T
hep
ossib
ilityexists
that captiv
ebreed
ing
groupsfor
allsubspecies
may
benecessary,
atleast
untilthe
troubledsociological
andpolitical
situationin
theis-
land
scalm
sdow
n.A
very
firmhan
dw
illhav
eto
betak
enw
iththose
who
breachpark
regulations,and
especiallyw
iththose
who
setup
illegalcam
ps,poach
protectedsp
ecies,or
delib
eratelyor
acciden
tallyin
trod
uce
newan
imal
(and
plan
t)sp
eciesto
pristin
eareas.
The
torto
iseso
fC
harles,
Barrin
gto
n,
Jervis.
Narborough,
Abingdon,
andsouthw
estC
hathamIslands
hav
ebeen
exterm
inated
ina
time
spanth
at,g
eolo
gically
speak
ing,
isa
blin
kof
aneye.
Su
relyth
eirsu
rviv
ing
relativescannotbe
farbehind
unlessdrastic
remedies
areunleash
edpro
mptly
.
LIT
ER
AT
UR
EC
ITE
D
AD
LE
R,K
.K.
1989. Herpetologists
ofthepast.
In:A
dler,K.K
.(E
d.).C
ontributionsto
theH
istoryof
Herpetology.
Soc.Study
Rept.
Am
ph.A
GA
SSIZ,L.
1857.C
ontributionsto
theN
atural History
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nitedStates,
Vol.
1.B
oston:L
ittle,B
rown,
andC
o.A
NoN
ivio
us.
1894.E
ditorial.N
ovitatesZ
oologicae1:1.
AN
oN
yio
us.
1980. New
sfrom
Acaderny
Bay.N
oticiasde
Galapagos
33:1-6.A
NO
NY
MO
US.1982.Is
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forLonesom
eG
eorge?N
oticiasde
Galapagos
35:2.A
No
Nio
us.
1986.T
heE
spañolatortoises
—a
veryspecial
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oticiasde
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44:5.A
Tim
soN
,G
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renchliterature.
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rom1700
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Paris,C
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FF
EN
BE
RG
,W.
1967.Notes
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IJFFEN
BE
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1.A
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1971:106-117.
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e.Z
oonooz51(12):4-7.
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N,J.P.
1980.Som
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urphyand
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uctiv
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iolo
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ISBN
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0-9653540-1-6(soft
cover)
