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Herpetologica, 57(4),
2001,
515-523
(?
2001
by
The
Herpetologists' League,
Inc.
A
SEVENTH
SPECIES
OF
MINUTE
SALAMANDER
(THORIUS: PLETHODONTIDAE)
FROM THE
SIERRA DE
JUAREZ,
OAXACA,
MEXICO
JAMES
HANKEN'
AND DAVID B. WAKE2
'Department
of Organismic
and
Evolutionary Biology
and Museum
of
Comparative Zoology,
Harvard University, Cambridge,
MA
02138,
USA
2Department
of
Integrative Biology
and
Museumii
of
Vertebrate
Zoology,
University of California,
Berkeley, CA 94720-3160,
USA
ABSTRACT:
We describe a new
terrestrial species of
minute lungless salamander
of the Mexican
genus
Thorius
(Plethodontidae)
from montane
pine-oak
forests
in
the Sierra
de
Juarez
of north
central Oaxaca, Mexico. The new species
is distinguished
from congeners by a
combination of body
size, external
morphology, osteology,
and dental traits, and
it is well differentiated
genetically from
other named species for which data
are available. This
is the seventh endemic
species of
Thornis
reported
from the Sierra de Juarez,
and known localities
are geographically isolated
from those of
all other
species.
Discovery
of
another
new species of plethodontid
salamander
from Oaxaca en-
hances
the state's standing as a preeminent
center of herpetological
diversity
within both M6xico
and Mesoamerica.
Key words:
Miniaturization; Taxonomy;
Biogeography;
Osteology
MESOAMERICA
iS one
of
the world's bio-
diversity
hotspots;
both the
relative
and
absolute
numbers of
species
and
the
de-
gree
of
endemism are
exceeded
by
few
other
terrestrial habitats
(Myers
et
al.,
2000).
Approximately
40%
of the
nearly
3000
species
of
tetrapod
vertebrates re-
ported from
Mesoamerica
are endemic
to
this
region,
and
they
represent
more
than
4%
of
the total
number of
tetrapod
species
known
worldwide. Within
Mesoamerica,
geographic
patterns
of
species richness
vary
among
taxa. For
amphibians and
rep-
tiles, an
important
center of
biological
di-
versity
is
the
Mexican
state of
Oaxaca.
In-
deed,
the
richness of
Oaxaca's
herpetofau-
na,
which
includes
more
than
one-third
of
all
species
of
amphibians
and
reptiles
found
in
Mexico, is
exceeded
only
by
Cos-
ta
Rica
(Casas-Andreu et
al.,
1996).
One
of
the
groups
that has
helped
to
establish
the
biological
importance
of both
Mesoamerica
and
Oaxaca
is
the
terrestrial
salamander
genus
Thorius
(Plethodonti-
dae).
Thorius
is
endemic to
Mexico,
where
it is
restricted
to
four
states
along
the
southeastern
edge
of
the
Mexican
pla-
teau-Oaxaca,
Veracruz,
Puebla,
and
Guerrero
(Wake and
Lynch,
1976).
Al-
though the
genus
was
first
described
>130
yr
ago
(Cope,
1869),
as
recently
as
1993
it
comprised
only
nine
formally
described
species
(Duellman,
1993).
In
the last
sev-
eral
years,
the number
of
valid,
named
species
has
more than doubled
to
22,
in-
cluding
five new
species described from
Oaxaca
(Hanken
and
Wake,
1994,
1998;
Hanken et
al.,
1999).
There
remain,
how-
ever,
many
populations of
Thorius that
cannot
be
confidently
or
easily
assigned
to
any
known
species,
and which
are
believed
to
represent
undescribed
species
(Hanken,
1980,
1983;
Hanken and
Wake,
1998).
Several of
these
populations exist
at re-
mote
montane
localities in
northern
and
western
Oaxaca,
and are
poorly
represent-
ed
in
most
museum
collections.
Here,
we
formally
describe a new
species
of
Thorius
based on
large series of
specimens
collect-
ed from
adjacent
localities
at the
extreme
northwestern
edge
of
the
Sierra de
Juairez
of
northern
Oaxaca
(Fig.
1).
This
species,
which
is
both
morphologically
and
genet-
ically
distinct
from
all
congeners,
further
enhances
Oaxaca's
standing as
an
impor-
tant
center
of
herpetological
diversity
within
both
Mexico
and
Mesoamerica.
MATERIALS
AND
METHODS
Measurements
were
made
using
digital
or
dial
calipers or
a
dissecting
microscope
fitted
with
an
ocular
micrometer;
standard
515
516 HERPETOLOGICA
[Vol. 57, No. 4
PUEBLA
(
OAXACA
\,VERACRUZ
11 *
schmidti
. tt
Sierra
de
Huautla
o
DOmineog
papaloae
@
67
aureus,
boreas,
.4'9'>
@
arboreus, smithi,
'R,
JF
insperatus,
;
macdougalli
1
p
ON
.]g
<
7-30,
32-45
San
Juan
del
Estado
0
sp. nov.
57,69
'
a
narisovalis
@46
'<
pulmonaris-
52
0
Oaxaca Mexico
40
km
97' 96'30
FIG.
1.-Map
of north-central
Oaxaca,
M6xico,
showing
the
type
localities
(closed
circles,
bracket)
of
Thorius
papaloae
and its
geographically
closest
con-
geners.
Thorius
papaloae
is known from several lo-
calities near the
village
of
Concepcion
Papalo.
All
lie
within montane
pine-oak
forest at elevations
ranging
between 2500 and 2850
m.
The
open
circle denotes
the
locality
of two
sympatric
populations
of Thorius
from 15
km
northeast of the
village
of
San
Juan
del
Estado,
2550 m
elevation.
Neither of these
popula-
tions can be referred to
any
named
species;
each
like-
ly
represents
an
additional,
undescribed
species.
Lo-
cation of main
map
area
in
southeastern
M6xico is
depicted
in inset at lower
right.
Populations
are
num-
bered
according
to Hanken
(1983).
length
(SL)
was measured from the ante-
rior
tip
of the snout to the
posterior
angle
of the vent. Limb interval
equals
the num-
ber of costal
interspaces
between the
tips
of
appressed
fore- and hind
limbs,
mea-
sured
in
one-half increments
(e.g., 3,
4.5).
Whole-mount
skeletal
preparations
were
stained for bone and
cartilage
using
aliza-
rin red S and Alcian blue
8GX, respective-
ly
(Klymkowsky
and
Hanken,
1991).
Os-
teological
descriptions
use the cranial
character states and
mesopodial
patterns
described and illustrated
by
Hanken
(1982,
1984,
1985),
Hanken and Wake
(1994,
1998),
and
Hanken et al.
(1999);
A
-
B
-
FIG.
2.-Photographs
of Thorius
papaloae.
(A)
Ho-
lotype,
MVZ
183468,
an adult female. Photo
by
D.
Wake.
(B)
Live
specimen,
collected
at the
type
lo-
cality by J.
Hanken and H. B.
Shaffer,
15
February
1976
(museum
number
unavailable).
Photo
by J.
Hanken. Scale
bars
=
0.5 cm.
see Wake and Elias
(1983)
for
comparisons
with other
tropical genera.
Counts of
pre-
sacral
(trunk)
vertebrae do
not include the
first,
or
atlas,
vertebra.
Except
for the ho-
lotype,
tooth counts
are based on cleared-
and-stained
specimens;
all alcoholic
spec-
imens were examined for
the
presence
of
maxillary
teeth.
Numbers
of vomerine
teeth
in
the
holotype
are
provided sepa-
rately
for
right
and left
sides;
these
counts
are summed for other
individuals.
Insti-
tutional abbreviations are as
listed in Lev-
iton et
al.
(1985)
except
for MZFC
(Museo
de
Zoologfia,
Facultad
de
Ciencias,
Univ-
ersidad Nacional Autonoma
de
M
dxico,
M
exico,
D.
F.).
DESCRIPTION
OF NEW
SPECIES
Thorius
papaloae
sp.
nov.
Paipalo
Minute Salamander
Holotype.-MVZ
183468,
an adult fe-
male from
8
km
(road)
NE
of
Concepci6-n
Paipalo,
Gaxaca, Medxico,
elevation
2670
m,-
collected
15
February
1976
by J.
Hanken
and
H. B. Shaffer
(Fig.
2A).
Paratypes.-All
from
Oaxaca,
Medxico:
MVZ
183470, 183473,
183475,
187052-69
(18
specimens),
same data as
the
holotype;
December
2001j
HERPETOLOGICA
517
MVZ
183479,
183483,
183485-6
(two
specimens),
183493-94
(two
specimens),
183496,
183504-5
(two
specimens),
183508,
183510,
MCZ A-134202-3
(two
specimens),
MZFC 12630-31
(two
speci-
mens),
same data
as the
holotype,
except
11 km
(road)
NE of
Concepcion
Papalo,
elevation
2820
m;
MVZ
183518,
same
data
as the
holotype,
except
15
km
(road)
NE
of
Concepcion
Paipalo,
elevation
2850
m;
LACM 121728-29
(two
specimens),
121731,
121733,
121737-38
(two
speci-
mens),
approximately
14 mi E of
Santos
Reyes
Paipalo
(likely
identified
incorrectly,
should instead be
Concepcion
Papalo-
see
Remarks),
elevation 2500
m,
collected
3
August
1975
by
T. W
Taylor
and A.
D.
Lau.
Some
specimens are
cleared and
stained or
have had tissue removed for
protein
comparisons.
Diagnosis.-This
is
a small
species
of
Thorius with
large,
elliptical
nostrils,
no
maxillary
teeth,
and a
relatively
long
tail
(Fig.
2B).
Most
specimens have
prominent
paratoid
glands.
It
is
distinguished
from
all
other small
species
of
Thorius from the Si-
erra de
Juairez
that lack
maxillary
teeth,
as
follows:
from
T
macdougalli
in
having
somewhat
larger hands
and
feet,
a
slightly
narrower
head,
more vomerine
teeth,
and
premaxillary teeth
in
most
females,
and
in
lacking
a
mottled dorsal
coloration;
from
T
insperatus
by
having
narrower,
more
elongate
nostrils;
from
T
arboreus
in
hav-
ing
larger
adult
size, a
larger, more elon-
gate
nostril,
and
a more
obscure
dorsal
coloration;
and from T
pulmonaris
in hav-
ing
smaller
adult
size and
a
relatively
shorter tail.
The
predominant
tarsal
pat-
tern is
different from
the
one
found
in
most other
species,
and there
are
signifi-
cant
protein
(allozyme)
differences.
Description.-This
is a
small
species;
adult
standard
length
averages 20.4
mm in
10
males
(range
19.0-23.2)
and 21.6
mm
in 10
females
(20.1-23.4).
The head
is rel-
atively broad;
SL
averages 7.2
times
head
width in
males
(range
6.5-7.5) and
7.4 in
females
(6.7-8.4).
Snouts
are
pointed.
Nostrils
are
large
and
elliptical; the
mean
ratio of
major
to
minor
axes
equals 1.7
(range
1.5-2.0) in
males and
females.
Eyes
are
moderately large
and
protrude
slightly
beyond
the
margin
of the
jaw
in
dorsal
view.
A
suborbital
groove
intersects
the
lip
on each side of the head. There
are
1-2
premaxillary
teeth
in
adult males
(mean
1.8) and 0-5
teeth
in
females
(mean
2.2).
There
are no
maxillary
teeth. Vomerine
teeth
average
6.1 in
both males
(range
4-
10)
and females
(4-8).
Limbs are moder-
ately
long;
limb
interval
averages
3.6
in
males
(range
3-4)
and 4.5
in
females
(4-
6).
Hands
and feet are
relatively
well de-
veloped and
moderately
broad;
foot width
averages
1.0 in
males
(0.9-1.1)
and
1.1
in
females
(0.9-1.2). Digits
1 and
4
(hand)
and
1 and
5
(foot)
are
short,
but the cen-
tral
digits are
relatively
long
and
have
rounded
tips.
Fingers,
in
order of decreas-
ing
length, are
3-2-4-1;
toes
are 3-2-4-5-1.
The tail
is
moderately
long (exceeds
stan-
dard
length)
and
tapered;
mean SL divid-
ed
by
tail
length
equals
0.85
in
two
males
(range
0.83-0.86) and 0.92 in
six females
(0.75-1.06).
A round and
relatively
prom-
inent
mental gland is
present
in
most adult
males
(maximum
dimensions: 1.2
mm
wide,
1.0 mm
long).
The
postiliac
gland
is
small and
pale,
and it is
relatively
incon-
spicuous
externally
in
some adults. Para-
toid
glands
are
prominent
in
most
speci-
mens,
but less
so
in
a few others.
This is
a
relatively
dark
species,
with
some indication
of an
obscure dorsal band
in
most
individuals.
Specimens with
the
lightest band have
a
herringbone
pattern
middorsally.
The
venter,
while
dark, is
lighter
than the
flanks,
and
the
underside
of
the tail
is
especially
light.
The
gular
re-
gion
is
marked
with
numerous white
spots.
Many
individuals
have a
light nuchal
spot,
and
some
have
a pair of
light
streaks over
the
shoulders.
Measurements
of
the
holotype (in
milli-
meters).-Head
width
3.0;
snout to
gular
fold
(head
length) 3.9;
head
depth
at pos-
terior
angle
of
jaw
1.8;
eyelid
width 0.7;
eyelid
length
1.5;
anterior
rim of
orbit to
snout
1.0;
horizontal
orbit
diameter 1.0;
interorbital
distance
1.7;
distance
separat-
ing
external
nares 0.8;
major
axis of
nostril
0.5;
minor
axis of
nostril
0.3;
snout
projec-
tion
beyond
mandible
0.5;
snout
to pos-
terior
angle of
vent
(standard
length) 22.1;
snout to
anterior
angle
of
vent 20.8;
snout
518
HERPETOLOGICA
[Vol. 57, No.
4
to
forelimb
6.0;
axilla to
groin
13.0;
limb
interval
6.0; shoulder
width
2.0;
tail
length
29.3;
tail
width
at base
2.0;
tail
depth at
base
2.2;
forelimb
length
(to
tip
of
longest
toe)
3.7;
hind limb
length
4.7;
hand
width
0.7;
foot
width 0.9. Numbers of
teeth:
pre-
maxillary
2;
maxillary
0;
vomerine 6-8.
The
right
foot is
partly
regenerated.
Coloration
of
the
holotype
(in alco-
hol).
-The
ground color of
the
head,
body,
and tail
is dark
blackish-brown.
It is dark-
est
along
the flanks
of the trunk
and
tail.
An
obscure
brown
dorsal
stripe
with
in-
distinct
borders
begins
on
the
nape
and
extends
posteriorly
onto the
proximal
por-
tion
of
the
tail. The
venter
is
pale
brown
with
scattered white
spots
in
the
gular
re-
gion;
the
ventral
spots become
indistinct
in
the
trunk. Limbs
are a
slightly
lighter
brown
than the
rest of
the
animal,
and the
hands
and
feet
are
even less
densely
pig-
mented.
Costal
grooves,
the
gular
fold,
and
the
extension
of the
fold
onto
the neck
all
stand
out
because of their
lack
of
pigment.
Otherwise,
there are
no
distinguishing
marks of
any kind. The
paratoid
gland
is
prominent.
Coloration in
life.-Based on
field
notes
by
J.
Hanken
(18
February
1976):
In
ju-
veniles,
the
solid
dorsal
stripe
is
interrupt-
ed
by
a
median
dark
line.
In
adults,
it
is
reduced
to a
dark,
herringbone
pattern.
Osteology.-This
description
is
based
on
data
from 18
adult
skeletons.
The
skull
is
very
poorly
ossified,
even
relative
to
most
other
species
of
Thorius.
Some cra-
nial
characters
are
sexually
dimorphic,
with
males
typically
less
developed
than
females.
The
degree
of
contact
between
ascending
processes of
the
premaxillary
bone is
highly
variable.
The
processes
re-
main
separate
from
one
another in
half
of
the
specimens
(character
1, state
a),
but
in
the
other
specimens
they
range
from
bare-
ly
articulating
to
fusing
along
more
than
half
of
their
length
(states
b-d).
Dental
processes
of the
premaxilla
are
separate
from
the
maxilla
in
all
males
and
most
fe-
males
(character
2,
state
a). In
a
few
fe-
males, the
two
elements
overlap
slightly
in
ventral
view
but
do
not
articulate
(state
b).
The
premaxilla
bears
teeth
in
all
speci-
mens
but
one
(character
8,
state
b).
The
nasal
bone
is
thin,
rod-like,
and
confined
to
the
posterior
edge
of
the nasal
capsule,
especially
in
males
(character
3,
state
b).
In
some males
and
many
females,
it
is
slightly
broader
and extends somewhat
an-
teriorly
over
the nasal
capsule
(state
c).
The
nasal and the maxilla
barely
articulate
in
most
specimens
(character
4,
state
b),
but
they
are
separate
in
a few
specimens
(state
a)
and
fuse to one another on
one
side of one
specimen
(state
c).
The
pre-
frontal
bone is
slender.
It remains
separate
from the
nasal in
nearly
all
females
(char-
acter
5, state
b)
but contacts the
nasal
in
all
males
(state
c).
In
all
specimens
but
one,
the
prefrontal
does
not extend
ventral
to the
dorsal
border of
the
nasolacrimal
foramen;
thus,
it
remains
well
separated
from
the
maxilla
(character
6,
state
a).
In
one
specimen,
the
prefrontal
extends
pos-
teriorly
beyond
the
nasolacrimal
foramen
to
articulate with
the
maxilla
(state
b).
Sep-
tomaxillary
bones are
absent in
nearly all
specimens
(character
7,
state
a).
In
one
specimen,
they are
represented
by
a
tiny
sliver of
bone at
the
edge
of
each
external
naris
(state
c).
The
maxillary bone
is
delicate-long
and
slender-and lacks
teeth
(character
9,
state
a).
In
one
specimen
(MVZ
187064),
the
right maxilla
appears to
have
been
bro-
ken and
partly
repaired;
a
cartilaginous
callus
bridges
the
fracture
site.
The
vomer
is
reasonably
well
developed.
A
short
to
very
short
preorbital
process
is
present
in
all
specimens
but one
and
bears
teeth.
There
are
relatively
few
vomerine
teeth,
which
are
arranged
in
a
short
row.
The
frontal
fontanelle
is
relatively
narrow
for
Thorius;
the
parietal
fontanelle
is
wide
(its
breadth
equals
0.44-0.67,
mean
0.52,
times
the
maximum
skull
width
across
the
parietals).
Otic
crests
are
lacking,
and
there is
no
columellar
process
on
the
oper-
culum.
The
postsquamosal
process
is
well
developed.
Hyobranchial
cartilages
are
mineralized
in
only
three
specimens
(and
only the
basibranchial).
All
specimens
have
14
presacral
verte-
brae.
Typically,
all
trunk
vertebrae
but
the
last
bear
ribs,
but
in
a
few
specimens
the
last
trunk
vertebra
has
only
a
partial
rib.
The
limbs
are
slender
but
well
developed.
December
2001]
HERPETOLOGICA
519
The
tibial
spur is
present as
an attached
crest
in
most
specimens,
but it
ranges
from
well
developed to absent
in
those remain-
ing.
Mesopodial
morphology is
variable,
al-
though
the
frequency
of
non-modal vari-
ants is somewhat lower than in
most other
species
of Thorius.
Asymmetry,
however,
is
common:
nearly
40%
of the
specimens
have a different
carpal
or tarsal
pattern
be-
tween
right
and
left sides.
Carpal pattern
I
predominates in the
forelimb
(94%
of
adult
carpi
examined).
This
pattern
con-
tains six
separate
elements,
with two de-
rived
states
in
relation to
outgroup genera:
fused intermedium
plus
ulnare,
and fused
distal
carpal
4
plus
centrale. It
is the most
generalized
pattern
observed in
Thorius
and is the
likely
ancestral state for
the
ge-
nus. Two other
carpal
patterns,
each
with
additional or
different
fusions
relative to
pattern
I,
are each found
in
a
single
adult
(one
side
only):
II
(fused
distal
carpals
1-
2
and 3),
3%, and VIII
(like
II, but with
the centrale
fused to the intermedium-ul-
nare instead of
distal
carpal 4,
and
centrale
1
fused to the first
metatarsal),
3%.
Pattern
VIII
is
unique
to
T
papaloae.
The modal tarsal
pattern is
V,
75%.
This
pattern
contains
seven
separate
elements,
with two
derived states
in
relation to out-
group genera:
fused
intermedium
and
fi-
bulare,
and fused
distal tarsals
4
and 5.
Tarsal
pattern
VII,
with
one additional fu-
sion
relative to
pattern
V
(fused
distal tar-
sal 4-5
and
centrale),
is
present
at a mod-
erate
frequency,
19%.
Pattern
I
(like V,
but
with
separate
intermedium and
fibulare),
the
presumed
ancestral
pattern
for Thor-
ius that
predominates
in
many
species,
is
found
on
only
one
side of one
adult,
3%.
The tarsal
pattern
could not be
scored
in
one abnormal
foot that had
only
four toes.
The
digital skeleton also is
highly
vari-
able,
especially
in
the hind
limb,
including
several
instances of
phalangeal
loss
and
gain.
The
predominant
phalangeal
formula
in
the
hand is 1-2-3-2
(86%
of adult
carpi).
1-2-3-1
and 2-2-3-2
are rare
variants,
3%
each;
the formula could
not be counted in
two
hands that were
damaged
during
preparation,
3%.
The
modal formula
in
the
foot is
1-2-3-3-2,
53%;
1-2-3-3-1
is a com-
FIG.
3.-Concepci6n
Papalo, Oaxaca, 15
Februaxy
1976. The
type
series
of
Thorius
papaloae was col-
lected from
terrestrial microhabitats
within
forested
slopes like those
visible
in
the
background, between
8
and 15
km
by road
NE of
the
village.
Photo by
J.
Hanken.
mon
variant,
22%. Three
other formulae
(1-2-3-2-2,
1-2-3-2-1,
and
1-2-3-4-1)
are
rare,
3-6%
each; a few
other
feet
were
damaged
during
preparation,
14%. Limb
bone
epiphyses
and
mesopodial
elements
are
mineralized
in
most adults.
Habitat
and
range.-Thorius
papaloae
is
known
only
from the
vicinity
of the
type
locality,
which lies at the
northwestern
edge
of
the Sierra de
Juarez
in
north-cen-
tral
Oaxaca,
Mexico,
northeast
of
the
vil-
lage
of
Concepcion Papalo
(Figs.
1, 3).
Re-
corded
elevations
range
from 2500
to
2850
m.
The
dominant natural habitat is
pine
or
pine-oak
forest
(Binford,
1989).
Goldman
(1951:220-221) discussed the local
terrain,
vegetation and climate of this
region,
which
he visited with E. W
Nelson
15-24
October 1894.
According
to field notes of
J.
Hanken
(15
and 18
February 1976),
most
salamanders
were collected under
cover
objects
on the
forest floor.
Speci-
mens collected at 15 km NE
of
Concep-
cion
PaTpalo
were
found beneath the bark
on
fallen
pine
logs
in
a small
clearing
alongside
the road. While this
occurred
during
the winter
dry
season,
moist
micro-
habitats
were common on
north-facing
wooded
slopes
with
abundant cover. Sev-
eral
specimens
of
Pseudoeurycea
are the
only
other
salamanders collected at these
localities
(e.g.,
MVZ
137002,
LACM
121722-27;
tentatively
assigned
to
P.
smi-
thi).
520
HERPETOLOGICA
[Vol.
57, No. 4
Etymology.
-The
species
name
refers
to the
village
of
Concepcion
Papalo,
which
is
very
near the
type
locality.
Papalo
(Span.)
is a
pungent
digestive
herb
eaten
raw by
the
sprig
with
many
Mexican
foods.
The word is
derived
from
papalotl,
which
means
"butterfly"
in
Nahuatl,
the
language
of
the
Aztecs.
Remarks.-Genetic variation in T
pa-
paloae and
relationships
to
congeners
were
examined
using protein
electrophoresis
by
Hanken
[1980,
1983;
populations
67-68,
listed
as
T
sp.
G
(part].
Evolutionary
con-
sequences of
miniaturization of
adult
body
size for
appendicular
morphology
were
ex-
amined
by
Hanken
(1982,
1985;
T
sp.
G).
Original
data
for the
type
specimens
from
MVZ,
MCZ,
and MZFC
misidentifies
the
village
of
Concepcion
Paipalo as
"Santos
Reyes
Palpalo,"
a
different
village
that
lies
only
a
few
kilometers
away,
due to
a
faulty
road
map
for
Oaxaca
State
in
wide use
at
the
time of
collection.
The
revised
(cor-
rect)
village name
used
in
locality data
above
is derived
from newer
and
improved
topographic
maps
for this
region,
and
was
confirmed
during
field work
by J.
Hanken,
D.
Wake,
G.
Parra-Olea,
and M.
Garcia-
Paris
in
summer
1999.
We
believe that
the
same
error
applies
to the
remaining
type
specimens from
LACM,
which
were col-
lected
within
a
year
of
the
former
speci-
mens "on
[the] road to
Santa Maria
PaTpalo
...
approximately
14
mi. E
of
Santos
Reyes
Papalo"
(letter from
A.
Brame
to
D.
Wake,
22
October
1975).
According to
newer
maps,
the
village of
Santa
Maria
Pa-
palo
is
accessible
by road
only
from
Con-
cepcion
PaTpalo
and
not from
Santos
Reyes
Paipalo.
The
earliest
record
of
T
papaloae
is a
single
specimen
(USNM
047797)
col-
lected
by E.
W
Nelson
and E.
A.
Goldman
from
"near
Reyes,
Oaxaca,"
elevation
3048
m,
on 20
October
1894.
"Reyes,
Oaxaca"
has
been
identified
as
Santos
Reyes
PaTpalo
(Binford,
1989;
Goldman,
1951),
and
is
thus
very
near
the
type
locality.
DISCUSSION
Recognition of
ThorTius
papaloae
as
a
new
species
is
justified by
its
large
degree
of
genetic
differentiation
from
other
named
species,
by
the
substantial
geo-
graphic distance of
known
localities
from
populations
of its
nearest
congeners,
and
by
subtle
but characteristic
morphological
features.
Thorius
papaloae
is
genetically
distinct
from
all
named
species
of
Thorius
in
northern Oaxaca and
adjacent
Puebla
for
which
allozyme data are available
(Table
1;
Fig.
4).
Levels of
genetic
differentiation
(Nei
genetic
distance,
DN;
Nei,
1972)
are
comparable
to or
exceed those that
are
typically
seen
among
congeneric
species
of
plethodontid
salamanders
(e.g.,
Highton,
2000).
In
contrast,
two
populations
of T
papaloae from
8
and 11 km
NE of
Con-
cepcion
Paipalo
are
virtually
identical
ge-
netically;
DN
equals
0.014
(n
equals
20
specimens
each).
Thortius
boreas
from the
Sierra de
Juairez
in
Oaxaca
is
the most sim-
ilar to
T
papaloae,
but even
this
distance
is
very
large-DN
equals 0.64-and
there
are
seven fixed
allelic
differences.
Adults
of
ThorTius
boreas also are
much
larger
and
more
robust,
and
they
are
osteologically
distinct. The
next
most similar
species
are
T
macdougalli,
also
from
the
Sierra
de
Juarez,
and T
pulmonaris
from
Cerro San
Felipe
in
the
Sierra
Aloapaneca.
Each of
these
species
has
a
DN
to
T
papaloae of
0.68, which
includes six or
seven
fixed
dif-
ferences
and
major
frequency
differences
at
2-3
additional
loci.
Thorius
pulmonaris
is
a
larger
species,
and
whereas T
mac-
dougalli resembles T
papaloae
in
some
morphological
features,
the two
latter
spe-
cies
differ in
dental
characters and
external
coloration.
Three
other
Oaxacan
species
(T
aureus,
T
narisovalis,
and T
arboreus)
each
have
a
DN
to T
papaloae of
around
0.8.
Thorius
aureus
and
T
narisovalis
are
much
larger
species;
T
aureus also
has
maxillary
teeth
and
a
distinctive
cranial
morphology.
Thorius
arboreus
is
smaller
and
has
a
different
external
coloration
than
T
papaloae,
and
it
occurs at
much
lower
elevations.
Finally,
T
schmidti,
from
southeastern
Puebla
and
extreme
northern
Oaxaca
(Hanken
and
Wake,
1998;
Garcia-
Paris
and
Parra-Olea,
1999),
has
a
DN of
1.24,
with 10
fixed
allelic
differences.
It
is
a
much
larger
species
and
has
maxillary
teeth.
Genetic
distances
from T
papaloae
to
December 2001]
HERPETOLOGICA
521
TABLE
1
-Genetic differentiation
between Thorius papaloae
and several of its geographically
closest con-
geners (Fig. 1). DN,
Nei
(1972)
genetic
distance between
populations;
Fixed, numbers of protein
loci showing
fixed differences between species
(boldface) or nearly fixed
differences, i.e., variant alleles
are shared by two
or more populations at frequencies
of
0.25 or less; n, numbers
of individuals sampled per population.
Allozyme
data
(Hanken, 1980, 1983)
are
unavailable
for
T
smithi and
T
insperatus,
two additional species from the
Sierra de Juarez, Oaxaca (Hanken
and Wake,
1994).
DN
to
Species
Locality
Population'
n T
papaloae
Fixed
Uncertain
San
Juan
del
Estado, Oaxaca
69 9 0.41 4, 32
Uncertain San Juan del Estado, Oaxaca
57 10 0.50 5,
23
T
boreas
Sierra de Juarez, Oaxaca
36 10 0.64
7, 04
T
pulmonaris Cerro
San Felipe, Oaxaca
52 3
0.68
6, 35
T
m-acdougalli
Sierra de Juarez, Oaxaca
30 17 0.68 7,
26
T
aureus Sierra
de Juarez, Oaxaca
37 9 0.80
7, 27
T
narisovalis
Cerro San Felipe, Oaxaca
46 20 0.80 8,
08
T
arboreus
Sierra de Juarez, Oaxaca
41
8
0.81 7,
29
T schmidti Zoquitlan,
Puebla
11 6 1.24
10,
210
' Populations
are numbered according
to
Hanken
(1983).
The reference
population
for T
papaloae (67)
is
from the
type
locality (n
=
20).
CAP, Ldh-1, Mdh-2, PEP, Aat-1,
Gp-2,
Idh-2.
Enzymes
are abbreviated
according
to
Murphy
et al.
(1996).
3
GAPDH, GPI, G3PDH, Ldh-1,
PEP, Aat-1,
Mdh-2.
CAP, Gp-2, GPI, G3PDH,
Ldh-1, Mdh-1,
PEP.
CAP, GAPDH, GPI, Ldh-1,
MPI, PEP, Aat-1, Idh-1,
Mdh-1.
6
CAP, GAPDH, GPI, G3PDH,
Ldh-1, MPI, PEP, Aat-1,
Mdh-1.
7Aat-1, CAP, GAPDH, GPI,
G3PDH, Mdh-1, Mdh-2, Idh-2,
PEP.
5Aat-1, CAP, Gp-2, G3PDH,
Ldh-1, Mdh-1, Mdh-2,
PEP.
CAP, GAPDH, GPI, G3PDH,
Ldh-1, Mdh-1, PEP, Aat-1,
Mdh-2.
CAIP,
GAPDH, Gp-2, GPI,
G3PDH, Ldh-1, Mdh-1, Mdh-2, MPI, PEP,
Idh-1, Ldh-2.
schmidti
pulmonaris
macdougalli
arboreus
SJDE
1
papaloae
SJDE
2
boreas
aureus
narisovalis
munificus
FIG.
4.-Neighbor-joining
phylogram
among
Thor-
ius
papaloae and the nine
geographically
closest
con-
geners
in
north-central Oaxaca and
adjacent
Puebla
for which
allozyme
data are
available
(Fig.
1;
Table
1). The
dendrogram
was
produced
by
analyzing
a ma-
trix of Nei
genetic distances
(DN)
with
the
program
Neighbor
in
PHYLIP, version 3.57c
(J.
Felsenstein,
University
of
Washington,
Seattle,
unpublished
1995). It
is
intended
to
depict
only the relative
mag-
nitude of
genetic
differences
among
taxa,
not
phylo-
genetic
relationships.
SJDE
1
and
SJDE
2
are
two
sympatric, undescribed
species from
near San
Juan
del
Estado,
in
the Sierra
Aloapaneca, Oaxaca
(Han-
ken
1980,
1983;
populations 57 and
69,
respectively).
Thorius
muniicus
from
northern
Veracruz is
includ-
ed as
an
outgroup,
although
the overall
branching
sequence is
unrooted.
Alternate
branching
schemes
among
these
and
additional
populations
based
on al-
lozyme data
are
presented
in
Hanken
(1983).
remaining
species
in
Oaxaca and
Puebla,
and
to
all
species
in
Veracruz and Guer-
rero,
are
similarly
large;
they range
from
0.41-1.63
(Hanken,
1980,
1983).
Thorius
papaloae
also is
morphologically
distinct
from
two
diminutive,
poorly
known
spe-
cies from
the Sierra de
Juarez
for which
allozyme
data are not available
(Hanken
and
Wake,
1994).
It differs
from
T
smithi
in
lacking
maxillary
teeth,
and from both
T
smithi and
T
insperatus
in
having
nar-
rower and
more
elongate
nostrils.
More-
over,
neither of
the latter two
species
is
known from
elevations
above 1550
m,
whereas
T
papaloae
has not
been taken
below 2550
m.
Hanken
(1980,
1983) used
allozyme
data to
assign
tentatively
one of two
sym-
patric
populations from
near
San Juan del
Estado,
in
the
Sierra
Aloapaneca,
Oaxaca
(T
sp. G,
population
69), to
the
same spe-
cies as
populations from
near
Concepcion
Paipalo, which
are
herein
described as T
papaloae.
Indeed,
each
of
the two
popu-
lations from
San
Juan del
Estado is
more
similar
genetically to
T
papaloae
than is
any
other
species of
Thorius,
named
or un-
named
(Table 1,
Fig. 4).
Based on
prelim-
inary
morphological
analysis
(Hanken
and
522
HERPETOLOGICA
[Vol.
57,
No.
4
Wake, unpublished),
we
regard
both
pop-
ulations
from San
Juan
del Estado as
dis-
tinct from those at Concepcion Papalo, so
at
this time
the known
range
of
T
papa-
loae
is
confined to
the
vicinity
of the
type
locality.
Neither
population
from San
Juan
del Estado
can be
confidently assigned
to
any
named
species
of
Thorius.
Their
iden-
tities, along
with
those of several remain-
ing enigmatic
and
geographically disparate
populations
from southern
and western
Oaxaca,
are
currently
under
investigation
and
will be considered
in
a
subsequent pa-
per.
Acknowledgments.
H.
Bradley
Shaffer
helped
collect
specimens
in
the field. B. Stein
(MVZ),
K.
Beaman (LACM),
0. Flores-Villela
(MZFC), J.
Ro-
sado
(MCZ),
and
A.
Wynn (USNM) kindly provided
access to and valuable
information
concerning spec-
imens
in
their care.
K.
Klitz and
T. Carl
helped pre-
pare
the
figures.
Research
support
was
provided by
NSF
(IBN-9419407
to
J. Hanken,
BSR-9019810
and
DEB-9408347 to D.
B.
Wake), by
the Council on
Research and Creative
Work, University
of
Colorado
at
Boulder, by
the Museum of
Vertebrate
Zoology,
the Center
for
Latin American
Studies,
and
Sigma
Xi
(Alpha chapter), University
of California at
Berkeley,
and
by
the Putnam
Expeditionary
Fund of the Mu-
seum of
Comparative Zoology,
Harvard
University.
The Direccion General de la Fauna
Silvestre,
M6xi-
co, provided collecting permits.
RESUMEN
Se describe una
especie
nueva de sala-
mandra de la
familia Plethodontidae del
genero
Thorius
del
bosque
de
Pino-Enci-
no de la Sierra
de Juairez en el Estado de
Oaxaca,
Mexico.
Esta salamandra se difer-
encia de sus
congeneres por una combi-
nacion
de
caracteres
morfologicos
que in-
cluyen tamanio
total
del cuerpo, osteologia
y
caracteristicas de la
denticion. Asimismo
esta
especie esta bien diferenciada
gene-
ticamente.
Esta es
la
septima especie en-
demica
de
Thorius
de la Sierra de
Juairez,
aunque
esta
se
encuentra
geograficamente
aislada de
las otras
especies de la
region.
El
descubrimiento de una especie nueva
de
salamandra de
la
familia Plethodonti-
dae
para
el
Estado de
Oaxaca,
incrementa
la
relevancia
del Estado como un centro
importante de diversidad
herpetologica
para
Mexico
y Mesoamerica.
LITERATURE CITED
BINFORD,
L. C. 1989.
A distributional
survey
of
the
birds of the Mexican
state of Oaxaca.
Ornithologi-
cal
Monographs
43:1-418.
CASAS-ANDREu, G.,
F. R.
MENDEZ-DE
LA
CRUZ,
AND
J. L. CAMARILLO.
1996. Anfibios
y
reptiles
de
Oa-
xaca. Lista,
distribucion
y
conservacion.
Acta Zool-
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Herpetologica,
57(4), 2001,
523-531
?
2001
by
The Herpetologists' League,
Inc.
A NEW TOAD OF
THE
BUFO MARGARITIFER
COMPLEX
(AMPHIBIA:
BUFONIDAE)
FROM NORTHWESTERN
VENEZUELA
ABRAHAM
MIJARES-URRUTIA' AND
ALEXIS ARENDS2
'Colecci6n Herpetol6gica Regional,
Centro cle
Investigaciones
en
Ecologfa y
Zonas Aridas
(CIEZA),
Universidad Francisco de
Miranda, Apartado 7559,
Santa
Ana
de
Coro
4101-A,
Venezuela
2Divisidn
de
Ecologia
Animal,
Centro dle
Investigaciones
en
Ecologia y
Zonas Aridas
(CIEZA),
Un
iversidad
Francisco de Miranda,
Apartado 7506,
Santa Ana
de Coro 4101-A, Venezuiela
ABSTRACT: We
describe a
new
species
of
toad
of
the
Bufo margaritifer complex
from
the state
of
Falc6n, Venezuela. This new
species differs
from
the other members of the
complex by the
following combination of
characters: medium-size (snout-vent length
mean
in 11
males 61.2 mm,
in
two females 72.9 mm),
presence of a bony knob at the angle of the
mouth,
heel
reaching the
posterior margin
of the
tympanum
when
hindlimbs are adpressed forward,
tibia length/snout-vent
length
ratio
ranging
from
0.40-0.45, toes almost fully webbed, tarsal fold
absent, cephalic crests
protuberant and thick,
supratymipanic
crest visible and projecting slightly obliquely
from body
(slightly
more
pronounced
in
females), and neural spines generally distinct (except in
some males).
Key
words:
Amphibia; Anura;
Bulfo
onargaritifer
complex; Bufonidae; Taxonomy;
Venezuela
AT
PRESENT,
one of
the
most
complex
taxonomic
problems
within
Neotropical
Bufonidae
involves
the
members
of
the
Bufo
margaritifer
(Laurenti,
1768)
com-
plex
(as
identified
by
Hass et
al.,
1995).
Hoogmoed
(1989a) stated:
(1)
the
name
Bufo
'typhonius'
(Linnaeus,
1758)
is
not
applicable to
the
species it
is
associated
with at
the
moment,
(2)
the
name
proba-
bly was
originally
based
on a
specimen
currently
assigned
to
the
genus
Rana (R.
tigerina
Daudin,
1802),
(3)
Bufo
margari-
tifer
would be the
correct
name
for
the
taxon
with
large
cephalic
crest,
occurring
in
the
Guianas,
and (4)
the
name
'typhon-
ius' should
be
suppressed. In
spite
of
Hoogmoed's
(1989a)
arguments,
some
herpetologists
still
consider
Bufo
'typhon-
ius'
valid
(Duellman
and
Schulte,
1992;
Velez,
1995)
perhaps
because
of a
lack of
a
review of
the
B.
mnargaritifer
complex
(Kohler
and
Lotters,
1999,
and
references
cited
therein)
and/or
type
series
for B.
inargaritifer.
Hoogmoed
(1986,
1989a,b,
1990)
removed
or
synonymized 13
of
the
17
species
names
which
had
been
associ-
