Beruflich Dokumente
Kultur Dokumente
QL
666
MISCELLANEOUS
OF KANSAS
JM OF NATURAL HISTORY
DIVERSITY
.L25
F7
PUBLICATION
No. 81
1989
DC
^
of Iguanian Lizards
Squamata)
-J
c
c
By
LAWRENCE
September
28, 1989
f f f
in 1946.
in that series.
Monographs of
the
of Natural History were initiated in 1970. Authors should contact the managing editor
regarding style and submission procedures before manuscript submission. All manuscripts are
subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion
Museum
of the Director.
this
number,
exchanging publications
may
and Miscellaneous Publications by addressing the Exchange Librarian, The University of Kansas
Library, Lawrence, Kansas 66045-2800, USA. Individuals may purchase separate numbers from
the Office of Publications, Museum of Natural History, The University of Kansas, Lawrence, Kansas
66045-2454, USA.
Museum of Comparative
Zoology library
Harvard University
Museum
of Natural History
Taxonomy
By
Barrel R. Frost^ and Richard Etheridge^
'Museum of Natural
Lawrence
1989
of
t,-^
JW^o
MISCELLANEOUS PUBLICATIONS
Managing
MCZ
LIBRARY
OCT
2 4 1989
HARVARD
UNIVERSITY
Miscellaneous Publication No. 81
Pp. 1-65; 24 figures; 3 appendices
ISBN: 0-89338-033^
USA
Printed by
University of Kansas Printing Service
Lawrence,
Kansas
CONTENTS
INTRODUCTION
ACKNOWLEDGMENTS
METHODS
RESULTS
16
18
Anoloids
21
Basiliscines
22
Crotaphytines
23
Iguanines
23
Morunasaurs
23
Oplurines
24
Sceloporines
24
Tropidurines
25
26
27
Summary of Results
29
31
31
32
34
34
34
36
Hoplocercidae
new
family
36
37
39
41
41
43
Leiocephalinae
new subfamily
45
45
47
SUMMARY
47
LITERATURE CITED
48
APPENDIX
53
APPENDIX
57
APPENDIX
62
INTRODUCTION
Iguanidae*^ is a large family (ca. 54 genera
and 546 species) of lizards found in the Americas, the Fiji Island group and the Tongatapu and
Va'vau groups in the Tonga Islands in the southwest Pacific Ocean, and Madagascar and the
Comoro Archipelago in the western Indian
Ocean
The unusual
(Fig. 1).
distribution of the
-f-
[Agamidae*
-i-
Chasister-
family
Squamata
the
1980]
Chamaeleonidae^
-I-
[6 genera,
128 species;
may
to other
[S ukhanov, 1 96 1 ]) the
,
(i.e.,
non-iguanian members of
Even though
known from
the
1983a,b). Thus,
al.,
way
to
used
metataxon convention is
same as the shutter quotation
the
here,
substantively the
[cf.
historical reality of
metataxa
is
questionable, their
Casual collectives
(e.g.,
is arbitrary.
^Because
this
family-group
name
is
based on the
name must be
Chamaeleonidae,
the
Chamaeleontidae.
rather
than
oft-used
Iguanidae* were
first
recognized by Etheridge
PauU
et
>200
are: (1)
anoloids
morunasaurs (3 genera;
25 species);
(5)
era;
182 species).
is to
reevaluate the
onidae, and
MISCELLANEOUS PUBLICATIONS
^G^^
60-
Vy-
Iguanidae*
Agamidae^
Chamaeleonidae
60
60
Fig.
1.
90
90
180
180
ACKNOWLEDGMENTS
For loan of and/or access
care
we thank:
to
specimens
in their
Zweifel, American
(AMNH); Robert
Museum
of Natural History
Norman
National
I.
Museum
of Natural History
University of
The herpetological
collection of San Diego State University (SDSU)
Texas
at
Arlington (UTA).
Museu de Zoologia,
Universidade de Sao Paulo, Brazil (MZUSP);
Gregory K.
(MVZ); Paulo
E. Vanzolini,
Pregill,
METHODS
The
1966;
tree
method of
PAUP), use of the
BLR ANGE (that calculates maximum and minimum branch links) and CSPOSS (that notes
character ambiguity on
Watrous and Wheeler, 1981; Arnold, 1981; Farris, 1982; Maddison et al., 1984; Kluge, 1985;
Brooks and Wiley, 1985; de Queiroz, 1985).
Following the evidence of Camp (1923),
Gauthier et al. (1988), and Estes et al. (1988),
Scleroglossa (=Scincogekkonomorpha = all noniguanian squamates) was selected as the
first
was
to determine,
where pos-
of particular transformation
reversal) output,
(i.e.,
series.
literature (e.g..
ment
tionship.
tions (see
few
traditionally
this analy-
were autapomorphies of
because the characters
within the transformation series suffered from
sis,
the
taxonomic
units, or
i.e.,
PAUP (Phylogenetic
across
all
taxonomic
Analysis Using Parsimony) version 2.4.1 (Swofford, 1985) and, late in the
development of the
Within
distribu-
tion of
.5 (Farris,
1988).
to constitute particular
That
first
we were
merely
how
char-
these cases, the "ancestor" cells in the data matrix were coded as unknown ("9" in PAUP; "?" in
network
ters
tinct trees of a
rela-
regard the
cedure to be performed on
topologically dis-
we
treated as if
all
evidence of
is
is
required
if
we
Additionally, in
some
multiple-step transfor-
mations, additivity of the steps was not maintained because of lack of evidence of order of
was one
step ("unordered"). In
some
in-
MISCELLANEOUS PUBLICATIONS
deduced even
though the polarity of transformations beyond
this initial condition could not be; in this case the
the ancestral condition could be
"ancestor"
the transformation
the initial
some
set.
nomic
we used deduced
ancestors as taxo-
was not
coded as "unknown" for those taxa. The analytical programs allow for this contingency by
making "what if assignments minimally in disagreement with the otherwise most parsimonious
arrangement of the characters on the tree.
We have not employed differential character
weighting. This practice has been argued against
by Patterson (1982) and Novacek (1986). Thus,
the analysis was allowed to proceed as though the
probability of undetected homoplasy was distrib-
all
transformation series.
However, should the assumption that the plasticity of transformation series was historically equal
prove to be wrong, or our a priori assessments of
homology prove to be wrong, our results will
have to be reevaluated.
allowed by: (1) the historical reality (=monophyly) of the terminal taxa; (2) the quality of
duced and the character assignment was "unknown." The terminal taxa that we employed
were:
is
and relative
homology
Our out-group
struc-
good
is
as
al.,
1988). Although
problems remain
al.,
some charac-
in several of the
A. Acrodonts.
its
broad contact
if it is
assumed
that the
is
Igua-
Agami-
mized as much
to termi-
as possible.
With regard
nal taxa,
would mean we might employ genera as our operational taxonomic units. However,
a number of these nominal genera are paraphyletic, and using these with their derivative
Iguania, this
Our
solution
was
to
(Moody,
1980 [unweighted analysis]), is also more easily
treated separately. For purposes of deducing
ancestral characters for agamas we provisionally
accepted the phylogeny of Moody (1980).
for
sis-
is
supported
monophyly
by so many
extremely extensile tongue, failure of the
pterygoid to meet the quadrate (Rieppel, 1981),
Bohme
its
cor-
is
the
and
its
both views.
Additionally,
we have
in order to
Moody, 1984) in
putative agamid would allow a more clear resolution of acrodont phylogeny. At least one fea-
saurs.
this
its
may
rior
mylohyoid foramina
Borsuk-BiaJ'ynicka and
in the splenial,
Moody
(1984), includ-
Williams, unpubl.])
may
Queiroz, 1988, presented some evidence to support this conclusion); however, our analysis
paraphyly of fPriscagaminae*.
B. Anoloids.
(9) "Pristidactylus";
anoles.
The monophyly of
the anoloids
is
monophyletic group;
this is
corroborated by the
it
and Williams,
bone and palatine teeth (Etheridge, 1959). However, unlike Guyer and Savage (1986), we regard
the phylogenetic structure within the remainder
corroborated a priori by the character endolymphatic sacs extending into the nuchal musculature (Etheridge, 1959; Etheridge
feature
is
de Queiroz, 1989).
C. Morunasaurs.
number of
length), but
Polychrus
is
united by a large
regarded as provisional.
Enyalius, "Pristidactylus," Diplolaemus, and
'Usually unjustifiably
emended
to Chamaeleolis.
MISCELLANEOUS PUBLICATIONS
which, in turn,
is
Hoplocercus. "Enyalioides" laticeps is the sistertaxon of "." praestabilis + the remaining mo-
The
tro-
based on com-
is
The a
priori
similar.
The
Liolaemus
group
(13) Basiliscus;
other
al-
Within
number
vomers.
all
{Phymaturus,
supported by
is
(14) Co-
D. Basiliscines.
cally
is due
Etheridge and de Queiroz (1988)
regarded Phymaturus as the sister-taxon of all
others, the latter group now with more than 100
species. Although the majority of these are, and
to
loss.
these are
of the
we have considered
to the
genus Lio-
new and
taxa, although
it is
laro, 1982;
pharys as monotypic
(C.
{Uma, Callisaurus, and Holbrookia). The sceloporines have their monophyly supported by the
cies
E. Sceloporines.
1948),
which involves an elongated septomaxilla. Additionally, they have unique hemipenes (Frost,
1987). Although thecladogram of Presch (1969)
was supported by Etheridge and de Queiroz
(1988),
it is
is
to
make
in this
we
Urosaurus, Uta,
we
we synonymize all of the
all
species
(i.e.,
laemus.
Leiocephalus
is
monophyletic (Etheridge,
and ante-
features that,
we have
Gambelia and Crotaphytus as terminal
treated
(Etheridge, 1967).
taxa.
H. Oplurines.
The oplurines
form a monophyletic
don.
Because '"Stenocercus"
is
may be
poly-
likely
clear that
it
is
reasonably
Chalarodon and Oplurus are sistertaxa (but see caveat in Etheridge and de Queiroz,
1988) we treat them separately because deducing
I.
Iguanines.
(33) Dipsosaurus;
(34) Bra-
monophyly of the iguanines, the large, herbivorous iguanids, seems unassailable. Although
shared with Uromastyx and Hydrosaurus in
Agamidae* (Iverson, 1980, 1982), the presence
of colic septa are likely a synapomorphy of this
Uracentron) has
rated by a
was unclear
its
group, as
Uranoscodon.
G. Crotaphytines. (29) Crotaphytus; (30)
Gambelia. The monophyly of the crotaphytines
is not supported by any descriptively unique
features, although they share a combination of
is
TRANSFORMATION SERIES
Sources of the various transformation series
apomorphic charac-
are given.
ter,
is
arbitrary.
which
A char-
The apparently
signment
similar xiphisternal bars in Tapinurus
major, which
is
ambiguous
myocommata
of the m. pectoralis
(character matrix).
shown
in
as-
assignments
Appendix
MISCELLANEOUS PUBLICATIONS
1.
is coded
"unknown,"
2. Maxillae (Cope, 1864; Estes et al.,
1988).
(0) do not meet, separated by premaxilla; (1) meet broadly anteromedially behind
as
(0) an-
at,
or
Our
plesiomorphic condition {contra Borsuk-Bialynicka and Moody, 984). Chameleons are coded as
"unknown" because Brookesia and some Chamaeleo have condition "0."
4. Vomers (Borsuk-BiaZynicka and Moody,
1
1984).
(0) flat or
interspecific variability.
pothesized.
Parietal
11.
frontal.
Laemanctus
is
ered to be "unknown."
cave.
1988).
"unknown" because of
12.
mostly on the
2). (0)
of the
foramen.
Queiroz, 1988).
7.
(0) lacrimal
1988).
is
interspecific
Lang, 1989).
posterior extremity
but
its
and
lateral view.
is
As an individual
variant in a
is
few
located
this set
of
variation.
"0"
is
plesiomorphic)
a third,
frontal
8.
we add
(1) extensive
scales,
authors,
is
coded as
Queiroz, 1988).
^vvCvKW*''-'
Fig. 2.
on
lateral side of
B: supratemporal substantially on medial side of supratemporal process of parietal (extent noted by dashed line). C:
supratemporal
fits in
Queiroz, 1988).
(0)
extend
parietal.
16. Dentary,
unpolarized.
17.
(Pregill, 1984;
fic variation
sia
is
difficult to charac-
all
(i.e., it is
beyond
level of superior
apex of coronoid;
(1)
is
coded as
this transformation.
18.
Coronoid
MISCELLANEOUS PUBLICATIONS
10
is
ambiguous because of
coded as "un-
known,"
21. Splenial, anterior extent (Fig. 4).
(0)
more than
Vfe
Because rhynchocephalians
polarized.
1988)
determined the identity of the angular (or anguloLabial view of mandibles. Top: P hysignathus
KU 69303 (scale=10 mm). Bottom: Leiocephalus
149104 (scale=10 mm). Lettered arrows
carinatus,
Fig.
3.
lesueuri,
UMMZ
show:
(a) dentary
Oplurus and
splenial?) in rhynchocephalians.
Liolaemus are sufficiently variable interspecifically that we have coded them as "unknown."
(0)
mandibular fossa;
edge of mandibular fossa. See comment under previous transformation series. Be-
at anterior
tion series
"Enyalioides"
larized.
in this
ation.
as
19.
(0)
foramen ventral
to posterior ex-
"unknown"
for the
4).
on ventral or
splenial
tion series
this
transforma-
be treated as unpolarized.
20. Meckel's groove (Etheridge and de
on
in C.
condition, out-
also coded
this
is
labial face.
Because rhyn-
this
transforma-
tion
specific variation.
(0) anterior to or
unpolarized.
dition in
24. Posterior
mylohyoid foramen
approximately
(Fig. 4).
at the level
of
is
is
11
earliest rhyn-
homologous.
known" because of
interspecific variability.
icles; (1)
Queiroz, 1988).
are considered
flat,
clavicular flange
arrows show:
variability.
Fig. 4. Lingual
(a)
splenial extending to or
beyond
Vt length of tooth
'/e
row
(b)
(Char.
length tooth
row
mylohyoid foramen
(Char. 24).
of this foramen
rus and
is
Opiums
"unknown" be-
Crowns
is
Because the
32. Interclavicle
except for
25.
this transformation.
Although Brachy-
3).
treated as
it
1988) (Fig.
anterior process,
is
1987).
and
lateral
MISCELLANEOUS PUBLICATIONS
12
absence of a scapular fenestra, although rhynchocephalians clearly lack them. For this reason,
transformation
this
is
unpolarized. Polychrus
is
because of interspecific
des"
is
considered to be
coded as "unknown"
variability. "Enyalioi-
"0" in other
KU
and
ambiguous
(i.e.,
Fig. 5.
plica,
M. A.
is
many
some
the
way
is
the plesiomorphic
and
lateral processes
is
we lack any clear justification for the pobetween conditions "1" and "2," we have
considered this transformation to be unordered.
37. Median enlarged sternal fontanelle(s)
(Etheridge, 1964; Moody, 1980; Etheridge and
fracture plane
de Queiroz, 1988).
34.
its
length.
(0) Scelop-
between paired
cause
larity
(Etheridge, 1967;
be
not paired; (2) present, large and paired. Although "0" clearly is the plesiomorphic condition, "1" and "2" are arguably polarized. There-
(transverse processes,
if
trans-
Moody,
1988). (0)
present, posterior to
unordered
on vertebra number
vertebra
number
4;
on
on vertebra
number
"0" or "3."
This transformation
is
considered
"unknown" because of
Chameleons are coded
"unknown" because numerical homology of
interspecific variability.
Queiroz, 1988).
5.
group comparison
is
Out-
ambiguous; that
is,
as
39.
be determined.
IGUANIAN LIZARD PHYLOGENY
13
interspecific variation,
"unknown."
fewer. Liolaemus
is
40. Postxiphisternal inscriptional ribs (Etheridge, 1965; Etheridge and de Queiroz, 1988).
(0) all attached proximally to dorsal ribs and
none are confluent midventrally; (1) one or more
pairs attached to dorsal ribs and are confluent
midventrally; (2) none attached to dorsal ribs or
continuous midventrally; present as pairs of isolated elements. Although "0" is clearly plesiomorphic, the polarity of "1" to "2" cannot be
determined. Therefore, this transformation series
is
considered unordered.
45.
variability.
lizards
(i.e.,
Uma
has a
anatomical grounds
it
is
questionable whether
is
ho-
(0) at least
considered as unpolarized
patagonicus; "1" in
P.
palluma).
row
(Etheridge and de
is
treated as
unpolarized.
Sand
coded as
41. Tail
is
Queiroz, 1988).
in P.
Phymaturus
47.
1988).
Gular
(0)
(e.g.,
oralis
is fre-
is
and
Ambiguous
out-group
Queiroz, 1988).
apomorphic condition
is
at the level
(e.g.,
Sceloporus nelsoni),
squamation
in
quently obvious.
comparison
though
this feature
has
(Sphenodon has condition "1," but scleroglossans are not really comparable) requires use of
Because of
MISCELLANEOUS PUBLICATIONS
14
scales,
it
apomorphic condition
is
from
major deviations
under subsequent
smooth. Out-
Queiroz, 1988).
digital scales
primitive condition by
variation.
condition.
The
more
(which
is
sup-
dorsal end.
of interspecific variation.
obviate
53. Nasal
1948).
54. Nasal
1948).
is
nasal cavity. In
*1
condition otherwise
of following discussion).
we
use
all
genetically,
we
is
shaped
like
unknown
a plow-share, a
in lizards.
Phylo-
anterodorsal position.
own
work
is
most
di-
observations.
been hypertrophied
to
is
by a "tongue" of
which
is
divided sagit-
tissue, the
concha, that
More
or
communicate with
the oral cavity. Behind the internal choana and
the concha is a blind cavity, the antorbital chamber. This condition obtains in Sphenodon and in
cha, the slit-like internal nares
55. Nasal
to roof of nasal
is
is
chamber (condition
fused
*3
of
some
is
slightly elongated
at
a relatively high
level.
56. Nasal
bins, 1948).
some
15
sumably
concha
is lost
some
a long vesti-
is
over the nasal chamber and enters the nasal chamber posterodorsally (Parsons, 1970). The nasal
is
intermediate
(0)
A-condi-
KU
Fig. 6.
ized.
(a)
Sceloporus torquatus,
91414,
umbra,
KU
MISCELLANEOUS PUBLICATIONS
16
tially
mally present but small in other lizards, is enlarged to the point that in superficial examination
it seems to have a
Because Sphenodon lacks
transformation must be re-
median
lobe.
a hemipenis, this
garded as unpolarized.
61. Hemipenis, capitation
6).
and
sulci (Fig.
distinctly divided sulci; (1) bilobate with distinctly divided sulci; (2) strongly bicapitate.
enough
to
polarity
0>1>2. However,
The
out-group similar
first
be comparable,
teiids,
support the
because Sphenodon
must be
Hemipenis, m. retractor
(0) not
lateralis
completely
septa in
Hydrosaurus.
66. Paired ventrolateral belly patches in
males (Etheridge and de Queiroz, 1988). (0)
absent; (1) present. "Stenocercus" is coded as
(e.g.,
large-scaled species
patches.
67. Reticular papillae
1988).
on tongue (Schwenk,
Because we
depended
regarded as unpolarized.
62.
entirely
on the
coded as "unknown."
RESULTS
A total of 225 alternative supported (as opposed to unrejected) tree topologies were discovered (208 steps; C.I.=0.385). Twelve networks
cluded
7).
gardless of the impression given by the consensus tree, not all of the phy logenetic trees logically
fact,
If
(e.g.,
Many
is
the only
measure
tionships,
gies
two
in the acrodonts,
is
is
presented in Figure
not a parsimonious
the
we have
failed egregiously.
in-group taxa
is
4.89 x 10
**'
made
(i.e., all
we
However,
trees for
35
(Felsenstein, 1978).
all
^^% of
we have
but 1.12 x 10
consider that
It is
The
trees.
MO
BA
AC
AG
CR
SC
CR
,OP
AN-(!)
AC
TR
,OP
BA
,AC
CR
AN
SC
I
MO
TR
I
OP
AN
MO
IG
UUh!)
CR
I
SC
I
TR
I
,OP
MO
,AC
IG
BA
IG
CR
SC
TR
U>
AN
MO
BA AC
OP
AN
BA
CR
SC
TR
IG
,OP
AN
CR
SC
TR
,OP
AC
IG
AN
SC
(!>
BA
IG
B
IG
TR
AN
MO
MO
BA
IG
17
"
MISCELLANEOUS PUBLICATIONS
18
fPriscagama
Agamas
rE Physignathus
EUromastyx
ACRODONTS
Leiolepis
Chameleons
Polychrus
HI Anoles
Para-anoles
Enyalius
"Pristidactylus"
"Enyalioides"
Basiliscus
Corytophanes
Laemanctus
'
Petrosaurus
all
10).
Topology 1 was discovered in intergroup Networks A-D and F-K, but Topology 2 was discovered only in Networks B, E, and L (Fig. 7).
Topology 1. The monophyly of the agama
operational taxonomic unit (=agamids,
excluding Physignathus, Uromastyx, and Leiolepis) may be supported by 32. 1 (well-developed
anterior process of interclavicle), however, the
interclavicle is lost in chameleons and is un-
known
tive
must be entertained
Uta
Urosaurus
Phrynosoma
Sand
lizards
'
wide lateral
ambiguous placement because the clavicle is absent in chameleons and unknown in fPriscagama*. Stem 1
(agamas + Physignathus) is corroborated by 15.1
(epiotic foramen) and is in agreement with the
Sceloporus
Phymaturus
results of
Moody
(1
is
also an
on scapula) may belong here, but placement is made ambiguous by the lack of a clavicle
in chameleons and being unknown in fPriscagama*. Chameleon monophyly is supported
by a number of characters that are apomorphies
clavicle
Ctenoblepharys
Liolaemus
'
Leiocephalus
"Stenocercus"
"Tropidurus
HZ Uranoscodon
C
C
num-
Opiums
Chalarodon
and 40.1 (midventrally confluent postxiphisternal inscriptional ribs) also are apomorphic.
Crotaphytus
Gambelia
Dipsosaurus
Brachylophus
Iguanas
unambiguously only one charmonophyly of the chameagamids, excluding Leiolepis and Uro-
Stem 2
carries
leons
-I-
men). Although
19
tPriscagama
tC
Agamas
Physignathus
Uromastyx
Leiolepis
Chameleons
"Enyalioides"
11
^-C
Enyalius
"Pristidactylus"
Para-anoles
10
14
tC
Polychrus
Anoles
Basiliscus
tc
13
32
Corytophanes
Laemanctus
Iguanas
Brachylophus
30
Dipsosaurus
28
31
Crotaphytus
Gambelia
Petrosaurus
Uta
16
29
Urosaurus
Sceloporus
18
41
-h
27
25
26
23
22
C
c
24
Phymaturus
Ctenoblepharys
Liolaemus
"Stenocercus"
20
Appendices 2 and
Chalarodon
Leiocephalus
21
Fig. 9.
Sand lizards
Phrynosoma
Opiums
among
tc
"Tropidurus"
Uranoscodon
MISCELLANEOUS PUBLICATIONS
20
Topology
Agamas
and Uro-
posterior).
Stem
mastyx)
is
3 (uniting Leiolepis
Physignathus
on
of
Chameleons
ribs),
row). 3.0
Uromastyx
Leiolepis
'^Priscagama
of
Topology 2
I
Agamas
Physignathus
Uromastyx
+ Leiolepis
I
Chameleons
^Priscagama
this stem:
toparietal suture)
is
Uromastyx
likely reversed in
(postfrontal reduced)
greater universality;
19.1
foramen ventral
some topologies
in
is
of
(anterior surangular
to posterior extremity
of den-
groups.
because
saurs and extremely enlarged in contrast to the
condition found in
sternal ribs
gies this
also
in
Hydrosaurus and
it is
unknown
in
ambiguous
in
cagama*.
Topology
2.
unknown
Agamas,
in fPris-
Physignathus,
Uromastyx)
in this
Stem
Topology
3 as in
1,
is
nearly the
same as
ambiguous
because of strong modification of chameleon
sterna and being unknown in -fPriscagama*, and
sternal fontanelles), although this
is
21
Topology
was sup-
Polychrus
Anoles
Para-anoles
Enyalius
"Pristidactylus"
are
signed 19.1
tral
(clavicular flange
flat,
Topology 2
Anoloids
do
is
Enyalius
"Pristidactylus"
Topology 3
its
Anoles
Para-anoles
ously
Polychrus
i-
Polychrus
Anoles
Para-anoles
Enyalius
+
5L_
"Pristidactylus"
we consider it to
MISCELLANEOUS PUBLICATIONS
22
ported in
all trees
Topology
Enyalius
is
by an
and 2,
its
placement
is
ambiguous.
(little
anterior
ele-
-i-
cor-
papillae).
Stem
(anoles
+ Polychrus)
is
Some
icle),
bra
placement
may have
a rela-
and 2
(Fig.
tail
autotomy
reversed
in anoles),
and ambigu-
which
is
shared with chameleons, morunasaurs, Brachylophus, and [in modified form] oplurines). In
Topologies
is
and
organs)
Topology
3, this is
of anoloids
-i-
regarded as a synapomorphy
oplurines.
Basiliscines
is
relationships
was
Apomorphies of
10.2 (Y-
anoloids (Fig. 11
weak
Stem
may
support a
vertebrae). In
(m. retractor
hemipenial
has
+ "Prisddactylus."
Enyalius
is
it
IGUANIAN LIZARD PHYLOGENY
Iguanines
donts, anoloids, or
perciliary
return to
su-
23
is
monophyly of
all
evidence supporting
Corytophanes
is
this relationship.
well-corroborated by 13.1
is
also a
Brachylophusnone,
1 1
three terminal
foramen in
and iguanas
(parietal
cion, however, is
character
frontal),
(V or Y-shaped
10.1
i\i2^.
parietal table).
Our
Dipsosaurus, with
suspi-
its
sce-
of Brachylophus
-i-
other
and modified
in-
scriptional ribs.
embryonically).
MORUNASAURS
Crotaphytines
The crotaphytines
phic in
is demonstrable only against the background of their out-groups, which possibly are
either a group composed of sceloporines,
oplurines, and tropidurines (and possibly including anoloids), or a group composed of acrodonts
and basiliscines. With respect to these, the crotaphytines have arguably made three reversals:
on the
More
ribs
interest-
the crotaphytines
is
extensive
skull rugosity in
the
may
larged),
It
8. 0(coronoid labial
MISCELLANEOUS PUBLICATIONS
24
(anterior surangular
Topology
uta
Petrosaurus
Oplurines
In all topologies, the oplurines
as monophyletic by:
were supported
Sceloporus
Urosaurus
Phrynosoma
c Sand
Lizards
forming paired
splints)
Topology 2
when
| uta
Networks
features are
Petrosaurus
Urosaurus
Sceloporus
Phrynosoma
c Sand
Lizards
SCELOPORINES
Three, equally parsimonious sceloporine topologies were discovered that were independent
lost),
Topology 3
monophyly of
was supported by: 28.1
all three,
uta
r^C
8 ^
Also
common
r Phrynosoma
to all
Petrosaurus
Sand
Lizards
Sceloporus
+
Fig.
12. Alternative
sceloporines.
Urosaurus
25
(e.g.,
subtended by this stem have relatively welldeveloped frontal scales that, if independent of
the interparietal scale development in this clade,
would lend support to either this arrangement or
Petrosaurus
was supported by a
saurus)
Topology
2.
may
is
is
to be
apomorphic when compared with the 3-sternalribbed tropidurines. Additionally, shimmy-burial, noted by Paull et al. (1976) and Etheridge and
de Queiroz (1988) as a possible synapomorphy
of the sceloporines, has been described in one
species of "Tropidurus" (Dixon and Wright,
1975) and may be found in at least one species of
''Agama" (Patterson, 1987). We suspect that this
behavior could be simply plesiomorphic within
Iguania.
and
Urosaurus was supported by no apomorphies in
Tropidurines
this analysis.
One
sal,
tion)
Queiroz (1988). In
all
networks
Stem
supported by
two unambiguously placed features: 23.1 (reduced angular) and 47.1 (gular fold incomplete
medially). In Network J, these features are joined
by 22.1 (posterior extension of splenial) and in
Network K by 37.1 (enlarged, median sternal
fontanelle). Although this character list is not
pidurines (Fig. 13
1) are
Topology
lizards,
reversal
if
3.
is
Liolaemus group
by 35.1
Leiocephalus
this to-
"Stenocercus"
pology.
r-
"Tropidurus"
Uranoscodon
sister-
we
MISCELLANEOUS PUBLICATIONS
26
first
out-
is
supported in
all
all
vari-
of the cladogram
cephalus
is
which
is
in this
likely
"neighborhood"
work
(Fig. 14). In
carries
3), and 39.0 (sternal ribs 4 [although Liolaemus was coded "unknown" because it has 3 or
4]), plus 30.1 (clavicular flange reduced); and
bra
Liolaemus has a reversal (36.0 posterior coracoid fenestra present). In Topology 1, Phymaturus is the sister-taxon of Ctenoblepharys +
parietal roof;
and 32.1
Topology
Stem
4, supporting the
by
shape of
apomorphic. Leio-
Taxa" (10.1
No
monophyly of
is
Stem
monophyly of
the
Liolaemus
Ctenoblepharys
(extreme posterior position of the posterior mylohyoid foramen), 36.0 (regaining of a posterior
Phymaturus
Topology 2
Liolaemus
by sternum
Phymaturus
Ctenoblepharys
Fig.
14.
Liolaemus group.
27
groove on the supratemporal process of the parietal (12.2). In this topology Ctenoblepharys is
apomorphic
in
suboculars.
may
-i-
tropidurines
noted by Arnold (1984) as possibly synapomorphic for the Liolaemus group, m. retractor later-
fossa)
more well-developed
feature
Liolaemus
and Ctenoblepharys than in Phymaturus. Degree
of development of this muscle supports Topoltion, is also
in
ogy 1.
Although Topology 2 is analytically equal to
Topology 1 it is important to note that it takes
advantage of an "unknown" in Phymaturus (45.0
in Phymaturus patagonicus; 45.1 in P. palluma).
Because P. patagonicus is otherwise more plesiomorphic than P. palluma, and because of the
hemipenial musculature character of Arnold
(1984) mentioned above we support Topology 1
as the most likely.
,
and 48.1
tary)
ally, 17.1
may
the
Stem
of Topology
fontanelle)
1").
placed ambiguously on
is
Also
in
1,
this
stem
(37.0) in oplurines
of tropidurines + (oplurines
SCELOPORINES + OpLURINES +
Tropidurines i+ Anoloids)
In Networks A-H, and L (Fig, 7), sceloporines
form the sister-taxon of oplurines tropidurines
Topology 1). In Network I, this ar(Fig. 15
rangement is augmented by anoloids being
-i-
Topology
2).
Network
In
J,
oplurines
Topology
15
(anoloids
-i-
Topology
3),
and
in
is
placed on
this
5) is also supported
-i-
Assuming
Net-
+ tropidurines]) (Fig.
The "fence lizard" habitus of
homologous
[oplurines
4).
we
is
at a
more
would
and anoloids.
ity is
due
to
plasy or plesiomorphy.
Topology
group)
is
1.
Stem
3,
is
MISCELLANEOUS PUBLICATIONS
28
Topology
Topology 2
Sceloporines
Sceloporines
I Tropidurines
Tropidurines
Opiurines
Opiurines
'
Anoloids
Topology 4
Topology 3
Sceloporines
Sceloporines
Tropidurines
Anoloids
+
+
Opiurines
10
L Anoloids
11
Opiurines
'
Tropidurines
in this topology
depends on the enlarged median sternal fontanelle of sceloporines and tropidurines
being a synapomorphy of the entire group (sce-
because
ture).
it
with a reversal, rather than the independent acquisition of this feature in the sceloporine
and
tropidurine clades.
Topology 3.
ously by 21.1
Stem
8,
(in-
also supported
is
character, 52.1
Stem 6
and 59.1
supported unambigu-
is
ribs)
gate)
convergently.
-i-
the
monophyly of
Two
may belong
superciliaries)
Stem
also sup-
is
11,
acter, 22.1
though
this is
convergent in anoles.
among
oplurines
at-
less
show
that contin-
collectives,
strict
consen-
Iguania (Fig.
8).
informal equivalents)
is listed
form
in
is
Figure 16.
relation-
is
illustrated in tree
The
29
parsimoni-
ous.
Summary of Results
Our
results
its
Chamaeleonidae
Chamaeleoninae
loids
Corytophanidae
Crotaphytidae
Hoplocercidae
Iguania
is
Iguanidae
-i-
Agaminae
No clear resolution
Leiolepidinae
Opiuridae
Phrynosomatidae
paraphyletic
Chamaeleonidae,
Polychridae
it
Liolaeminae
For
is
this
would require
Tropiduridae
Fig. 16.
Taxonomic
tree.
Leiocephalinae
Tropidurinae
MISCELLANEOUS PUBLICATIONS
30
Chamaeleonidae
Rafinesque,
1815
idae)
inae)
Corytophanidae Fitzinger,
1843 ("Igua-
stituent groups.
Within Tropiduridae
we
recog-
nidae": basiliscines)
family ("Iguanidae":
de Queiroz (1987) proposed a suprageneric phylogenetic taxonomy within his Iguaninae (our
("Iguanidae": crotaphytines)
Hoplocercidae new
morunasaurs)
Iguanidae),
nines)
Opluridae Moody,
it
1983
("Iguanidae":
nidae": sceloporines)
we
tro-
pidurines)
in-
anoloids)
ings,
oplurines)
is
different
nia).
(i.e.,
new subfamily
Liolaeminae new subfamily
category as
Leiocephalinae
tural
The
we
have,
redundancy.
We
we
between this taxonomy and the traditional one (e.g.. Camp, 1923;
Estes et al, 1988) are: (1) Agamidae* and Chamaeleonidae have been combined and (2) the
informal groups within "Iguanidae" are accorded
formal, independent taxonomic status. With the
ranking
salient differences
this
is
case
is
arbitrary
some
much
and much
to
little to
do with a crude
do with evolution
sort of essentialism.
31
characterizations provided for the famiand subfamilies are not lists of apomorphies;
those data are available in Appendices 2 and 3
lies
and
in "Results."
suspended
in
synonymies
1815.
Nat.:75.
Lacert.: 12.
1802.
Philadelphia, 16:226.
Characterization.
em-
some groups)
between the
(Estes et
al.,
orbits (reversed
Chamae-
name
[Opinion 89]).
1825. Stellionidae Bell, Zool.
J.,
London,
See
dae.
Distribution.
tropical regions.
attribution
is in error;
Cuvier used
1768).
Neue
Classif.
Kaup
1825).
Type genus:
"'Calotes
Kaup" (=Calotes
Cuvier, 1817).
"Lophura Wagl[er],
Type genus:
old sense),
onidae.
now
equivalent to Chamaele-
Kaup, 1825.
MISCELLANEOUS PUBLICATIONS
32
GeoL
be bicapitate
Distribution.
ment.
to
maxillae meet
(1)
Tropical
and temperate
re-
Comment.
is
meleons and "[Priscagama*); (4) jugal and squamosal broadly juxtaposed (not ^Priscagama*)',
(5) parietal roof shape quadrangular (or domed in
chameleons); (6) parietal foramen usually ab-
sent, if present,
on frontoparietal suture;
(7)
Chamaeleonidae
tional
supported.
its
(1988).
which
al.
The
(the
the tradi-
chameleons)
is
well
Moody
(1984) are
Chamaeleonidae because they are not tied together unambiguously by apomorphies, although
lost entirely
[Priscagama*); (10) no
labial blade
foramen inferior
Chamaeleonidae.
of coronoid;
AGAMINAE
to pos-
groove broadly open; (13) splenial short anteriorly, or absent; (14) dentary and maxillary teeth
acrodont, fused to underlying bone in adults; (15)
1825.
SPIX, 1825
Agamae
Spix,
1802.
1825. Stellionidae Bell, Zool,
J.,
London,
number of
sternal
1768).
See
member of Iguania.
1826. Draconoidea Fitzinger,
Rept.:ll.
Neue
Classif.
1:15.
Kaup
ribs variable;
row
variable; (25)
1825).
some
organs of some
agamines are clearly not homologous); (28) acrodontan nasal apparatus (except Physignathus),
Cuvier, 1817).
60
120
150
33
180
60
60
30
30
Type genus:
"Lophura Wagl[er].
Kaup, 1825.
Characterization.
(1)
vomer
flat
or con-
normal feet.
Content*. Acanthosaura
phically; (9)
Until
the
controversy
surrounding
the
is
resolved,
we
refrain
from
Gray,
1831;
dosaurus.
to
Chlamy-
MISCELLANEOUS PUBLICATIONS
34
we simply
Klaver and
siini
Peters, 1863;
Xenagama Boulenger,
Distribution.
Temperate and
1895.
tropical Eura-
Bohme
Furcifer, Bradypodion,
sia,
Comment.
equivalent to
As
Agamidae of previous
is
authors, but
J.
Linn. Soc.
Zool., 10:34.
CHAMAELEONINAE RAFINESQUE,
1815
1820).
Characterization.
Nat.:75.
Chamae-
name
(Opinion 89).
1923. Brookesinae Nopsca, Fortschr. Geol.
Palaeont., 2:124. Type genus: Brookesia Gray, 1865.
Characterization.
See comment.
(1)
vomer
flat
or con-
domed and
foramen absent;
no
(1)
vomer concave;
(2)
normal
recognized previously
(e.g.,
Borsuk-BiaZynicka
is
Calumma*
1843;
Mus.
for 1899:223.
Natl.
em
and northwestern Arabia, Madagascar, Seychelles, India, and Sri Lanka, and associated
Characterization.
Type genus:
Comment.
Our purpose here is not to evaluwork on the phylogeny of chameleons, and although we have Brookesiinae Nopsca (Klaver and Bohme, 1986) in the synonymy
ate previous
of Chamaeleonidae, this
is
only in recognition of
Laemanctus);
(4) jugal
postembryonically
our Chamaeleoninae
in
is
embryonically
(6) parietal
in frontal (parietal
Laemanctus);
(7)
35
foramen absent in
sits on lateral
supratemporal
(24)
in
some
Basilis-
Content.
Basiliscus
1834.
Comment.
to the
(1989).
Fig.
1 8.
Distribution of Corytophanidae.
al.
(1976),
MISCELLANEOUS PUBLICATIONS
36
HOPLOCERCIDAE
Characterization.
except in old
foramen
supratemporal
in frontoparietal suture;
sits
on
lateral side
(7)
of supratem-
NeW
Type genus.
lateral side of
expanded onto
to
on
(5) parietal
roof trapezoidal; (6) parietal foramen in frontoparietal suture (absent in some "Morunasau-
FaMILY
labial
foramen inferior
groove not fused; (13) splenial very large, penetrating far anteriorly; (14) dentary and maxillary
teeth pleurodont, not fused to underlying
bone
in
icle
to
icle
fracture planes;
"Morunasaurus" an-
number
phistemal inscriptional ribs short; (23) interparietal scale not enlarged; (24) mid-dorsal scale row
in
ent;
straight;
Comment.
Crotaphytidae
(1988).
4;
(22)
concha present,
free; (29)
hemipenes
Eastern Panama
to the Pacific
corresponds to
and de Queiroz
Brazil,
em
37
Comment.
Hoplocercidae
corresponds to
al.
We
(1973),
Fam. Gatt.
Type genus: "Iguana Linne"
38
MISCELLANEOUS PUBLICATIONS
39
comment.
Tonga Islands
Characterization.
meet-
Comment.
Fiji
and
(Fig. 21).
Iguanidae
corresponds to the
"Results").
parietal
rugose (except
Am-
(in fron-
Characterization.
(8)
Cuvier, 1829.
meet-
and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal; (6) parietal foramen in frontoparietal suture; (7) supratemporal sits on medial
nuchal endolymphatic sacs not penetrating nuchal musculature; (9) dentary not expanded onto
of cor-
in
(14) dentary
and maxillary
tate, unisulcate;
Content.
Amblyrhynchus
Bell, 1825;
Bra-
1856.
1843; Ctenosaura
splints, isolated
ribs
and not
row absent
40
MISCELLANEOUS PUBLICATIONS
41
on
lateral
free;
concha present,
Content.
1837;
Opiums
Cuvier, 1829.
Distribution.
Western
Comoro Islands.
Comment. Opluridae corresponds
"oplurines" of Smith et
al.
to the
(1973), Etheridge in
Paull et
al.
(1988).
The nomenclatural
dae
is
gascar;
coronoid poorly developed or absent; (11) anterior surangular foramen above posteriormost
status of Doryphori-
(14) dentary
by
sternum far anteriorly; (18) caudal autotomy
fracture planes present (except in Phrynosoma),
with transverse processes anterior to fracture
number
3 or
4 (Petrosaurus); (22)
Phrynosoma
imiliani
1919);
Uma
USA
to the "sceloporines" of
de Queiroz (1988).
1971. Sceloporinae Kastle, Grzimek's Tierleben, 6:181-182. Type genus: Sceloporus V^icgmann, 1828.
Characterization.
meet-
Rept.: 11.
comment.
and
(5) parietal
Neue
Classif.
stated.
See
MISCELLANEOUS PUBLICATIONS
42
Daudin, 1802).
1864. Anolidae Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:227.
lis
Daudin, 1802.
Characterization.
meet-
roof trapezoidal or
foramen normally
parietal in
sits
on
in
some
lateral side
IGUANIAN LIZARD PHYLOGENY
43
tional
bone
in
some Poly-
sor")
and Recommendation
24A
of the Interna-
Pneustes prehensilis
1957), considered to
tus vautieri
Merrem, 1820).
synonym oi Polychrus
number
2, 3, or 4; (22)
postxiphistemal inscrip-
row
lis);
plesiomorphically
some Ano-
litt.).
is
argu-
Content.
Anisolepis Boulenger,
1885
Type
genus:
Tropidurus WiedNeuwied, 1825. See comment under
Tropidurinae.
1843. Ptychosauri Fitzinger, Syst. Rept., 1:16.
to
(Fig. 23),
in
1825).
1:17.
ger,
Fitzin-
(1988).
We
Fitzinger,
Characterization.
meet-
1843, and
44
MISCELLANEOUS PUBLICATIONS
IGUANIAN LIZARD PHYLOGENY
(3) skull roof not strongly rugose; (4) jugal
and
45
to
Comment.
Tropiduridae corresponds
to the
LEIOCEPHALINAE
Type genus.
NEW SUBFAMILY
Characterization.
overlapped by nasals; (2) parietal table Y or Vshaped (shared with some iguanids and anoles);
(except
some Liolaemus)', (13) splenial very short antesome liolaemines); (14) dentary
more than
Ve length
no preanal
riorly (except in
some
Leiocephalus, some "Stenocercus"); (17) posterior process of interclavicle not invested by ster-
num
"Tropidurus"
hemipenes.
Content.
Distribution.
Bahama
Islands,
Cuba and
guilla, Martinique,
al, 1988]);
and Guadeloupe
Cayman
[Pregill et
number
some "Tropidurus");
(21)
3 or 4; (22) postxiphistemal
soides" and
LIOLAEMINAE
NEW SUBFAMILY
or V-shaped: (3)
precoronoid length of mandible; (5) poorly developed anterior process of interclavicle; (6)
inae,
some
(except in
Content.
Leiocephalinae
new subfamily;
Distribution.
The
Bahama
Islands,
Cuba
hemipenes.
MISCELLANEOUS PUBLICATIONS
46
60
90
Leiocephalinae
20
Tropidurinae
20
20-
1000
Kilometers
40
Liolaeminae
40
60
90
Fig. 24. Distribution of Tropiduridae, including subfamilies.
The
Holocene distribution of
30
that taxon.
was due
to
it
is
Commission of Zoological
Nomenclature be petitioned to resolve the problem (Art. 70b; International Code of Zoological
Nomenclature, 1985).
Characterization.
1843
genus:
Tropidurus
1.
Wied-
Fitzin-
1825).
no more than
Vfe
"Ophryoessoides"
Dumeril,
ger,
Type
Content.
TROPIDURINAE BELL,
47
Fitzin-
SUMMARY
A
formed using 67 transformation series containing 147 characters of osteology, dentition, squa-
duce 18
all
units,
trees of nine
+ Chamaeleonidae],
(acrodonts [= Agamidae*
tro-
analysis
topology.
on unrooted network
name
Therefore,
publication.
MISCELLANEOUS PUBLICATIONS
48
two
in the acrodonts,
and three
new
tionships, "Iguanidae"
taphytines); Hoplocercidae
iguanines); Opluridae
Chamae-
was because of
trees
much
of
between discovered
this lack
of resolution.
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An
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APPENDIX
New
Wiley
\JR ANO=Uranoscodon;
glossa;
yalius; ?RlST='Pristidactylus"\
?ARAA=Urostro-
ENYLD
CORYT=Cory-
tophanes;
rus;
CROTA=Crotaphytus;
MISCELLANEOUS PUBLICATIONS
54
Appendix
continued.
55
continued.
Character
SPHEN
SCLER
2222222333333333
3456789012345678
?00?0000100
?000100?010
ANCES
Taxon
PRISC
AGAMI
UROMA
LEIOL
PHYSI
12
0?
CHAME
1
POLYC
ENYAL
PRIST
PARAA
ANGLE
ENYLD
?
1
BASIL
CORYT
LAEMA
PETRO
SCELO
UROSA
UTA
PHRYN
SANDL
2
7
1
1
1
1
PHYMA
CTENO
LIOLA
LEIOC
?
?
STENO
TROPI
URANO
CROTA
GAMBE
OPLUR
CHALA
DIPSO
?
1
BRACK
IGUAN
10
MISCELLANEOUS PUBLICATIONS
56
Appendix
continued.
APPENDIX
Apomorphy
57
Figure 9. See legend of Appendix 1 for abbreviations used. Asterisks note character
of ambiguous placement. Characters from unpolarized or unordered transformations are noted by a
"U." Daggers note other characters (not an exhaustive list) not used in the analysis.
lists for tree in
Transformation
Stem
PRISC
Series
Ancestral
Character
Derived
Character
MISCELLANEOUS PUBLICATIONS
58
Appendix 2 continued.
Transformation
Stem
Series
PETRO
Ancestral
Character
38
39
Derived
Character
1
1
SCELO
UROSA
UTA
PHRYN
tsecondary cusps of posterior marginal teeth reduced (Etheridge and de Queiroz, 1988).
tdark axillary spot (Mittleman, 1942) (homologous with those in sand lizards?).
SANDL
35
47
fskuU and head scales strongly modified (Reeve, 1952; Presch, 1969).
tphalanges lost from digits 4 and 5 of manus (Etheridge and de Queiroz, 1988).
1
66
tlabial scales elongated, obliquely oriented; lower jaw countersunk (Smith, 1946; Axtell,
1958).
PHYMA
21
30
38
39
CTENO
45
LIOLA
17
1
1
1
36
LEIOC
10
32
tlong, free xiphisternal rods curve forward to underly xiphisternal ribs (Etheridge, 1966).
tpostrostral scales lost (Etheridge, 1966).
STENO
TROPI
23
tarticular surface of
URANO
U
U
(Frost, 1987).
44
45
47
59
CROTA
GAMBE
7
41
tclavicular fenestrae (Etheridge and de Queiroz, 1988).
many
Crotaphytus).
OPLUR
CHALA
30
46
U
u
u
44
45
40
1
1
11
1
1
1
41
10
IGUAN
36
15
31
U
U
DIPSO
BRACK
38
39
1
1
1
1
1
59
Appendix 2 continued.
Transformation
Stem
Series
3
9
10
Ancestral
Character
Derived
Character
61
Appendix 2 continued.
Transformation
Stem
Series
24
*
*
Ancestral
Character
Derived
Character
1
1
1
25
12
2
1
1
1
26
U* 17
MISCELLANEOUS PUBLICATIONS
62
APPENDIX
Transformation
Series
Changed
From To
*
10
11
12
13
14
15
U16
U17
Along
Stem
Consistency
63
Appendix 3 continued.
Transformation
Series
U18
U19
U20
U21
U22
U23
24
25
26
U27
28
29
U30
U31
U32
33
U34
Changed
Erom To
*
Along
Stem
Consistency
MISCELLANEOUS PUBLICATIONS
64
Appendix 3 continued.
Transformation
Series
Changed
From To
U35
Along
Stem
Consistency
31
23
16
SCELO
U36
0.250
10
28
20
7
IGUAN
LIOLA
UROMA
U37
27
3
25
5
U38
0.286
0.667
30
28
7
OPLUR
PHYMA
PETRO
39
0.286
27
*
5
8
7
2
OPLUR
PHYMA
PETRO
CHAME
U40
1
3
25
BRACH
41
0.222
11
0.500
12
8
BRACH
CROTA
42
19
16
43
25
27
U44
1
1
0.200
0.500
1.000
DIPSO
URANO
U45
*
BASIL
27
DIPSO
0.250
URANO
CTENO
1
BASIL
0.167
65
Appendix 3 continued.
Transformation
Series
Changed
From To
U46
Along
Stem
Consistency
29
1
21
8
2
1
CHALA
0.200
24
47
7
1
URANO
SCELO
CHAME
U48
0.200
31
26
*
POLYC
CHAME
23
9
10
49
50
51
*
*
52
POLYC
18
0.333
1.000
31
27
23
20
0.333
1.000
55
56
57
0.500
10
25
1
53
54
POLYC
0.200
1.000
1.000
U58
PHYSI
1.000
0.500
25
CHAME
U59
18
30
21
15
URANO
U60
18
U61
0.500
0.200
1.000
10
20
1
U62
U63
19
18
0.500
LEIOL
0.500
13
U64
ENYAL
0.500
30
65
UROMA
66
0.500
15
SANDL
67
0.500
19
0.500
ANOL
0.500
Her,,
its"
QL666.i F7 1989
Date Due
RECENT
67.
An
MIJ
ecogeographic analysis of
th
fi
<^^T*^
B (\\f^
Lee.
/)
,^-^
4
I "\
3 4Q flft
c,
evolutionary
June 24,
1980.
AP^30i995
^ - ^***^
Pp
C^4i
<y^S>
i:
Leptodactyli-
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