~oologicaljoumalofthe Linnean So&$ (2000), 128: 189 233.

With 14 figures
to
doi: 10.1006/zjls. 1999.0203, available online at [Link] on 10E bl
*

A phylogenetic analysis of the tropidurine

lizards (Squamata: Tropiduridae), including new
characters of squamation and epidermal
microstructure

MICHAEL B. HARVEY' AND RONALD L. GUTBERLET JR'

Department of Biology, Box 19498, ?he UniversiQ o f Texas at Arlington, Arlington,
T X 76019-0498, U.S.A.

Receiued March 1997; accepted f o r publication February 1999

Characters of scale surface microstructure are combined with 'traditional' morphological

characters in a phylogenetic analysis of the Tropidurini. Tropidurid lizards show variation
in types of coarse and fine scale surface microstructure, in the anatomical distribution of
different scale surface features, and in scale organ morphology and distribution. The
morphology of the inner surface zone of scales is here described for the first time using
scanning electron microscopy. Our phylogeny differs considerably from those proposed in
earlier studies. New characters and frequency coding of polymorphic characters help resolve
the problematic relationships of several species. Statistical confidence supports recognition
of one large cis-Andean and one large trans-Andean clade of species. Based on our
results, we synonymize Plesiomicrolophzls with Microlophus and Uranoscodon with Tmpidurus. The
phylogenetic relationships of newly discovered Tropidurus are resolved: I: callathehs is the
sister species of 1: melanopleurus; I: xanthochilus is the sister species of I: spinulosus. Tropidurus
spinulosus is found to be more closely related to 1:stmbilurus and a clade of Amazonian species
than to I: melanopleurus. The species previously placed in Uracentron are more closely related
to species previously placed in Plica than to 1: stmbilurus as previously thought.
0 2000 The Linnran Society of London

ADDITIONAL KEY WORDS:-Tmpidurus -- Uranoscodon Micro1ophu.r ~ ~ Plesiomicmlophus

microdermatoglyphics epidermis - frequency coding systematics.
~ ~

CONIENIS

Introduction . . . . . . . . . . . . . . . . . . . . . . . 190
Methods . . . . . . . . . . . . . . . . . . . . . . . . 191
Source of characters . . . . . . . . . . . . . . . . . . 191
Character coding and polarity . . . . . . . . . . . . . . . 192
Phylogenetic analysis . . . . . . . . . . . . . . . . . . 196
Evaluation of confidence and phylogenetic signal . . . . . . . . . 196

' Corresponding author. Present address: Dept Biological Sciences, Box 70703, East Tennessee State
University, Johnson City, T N 376 14-0703, U.S.A. E-mail: harveym@[Link]
Present address: Dept Biology, University of Texas at Tyler, 3900 University Blvd, Tyler, T X 75799,
U.S.A.
189
0024-4082/00/020189+45 $35.00/0 0 2000 The Linnean Society of London
190 hl. B. HARVEY AND R. I,. GUTRERLET JR

Scale surface morphology of tropidurids . . . . . . . . . . . . . . 197

Coarse scale surface microstructure . . . . . . . . . . . . . . 198
Fine scale surface microstructure . . . . . . . . . . . . . . 202
Scale organs . . . . . . . . . . . . . . . . . . . . . 203
Transformation series of new characters . . . . . . . . . . . . . . 208
Results of the phylogenetic analysis . . . . . . . . . . . . . . . 212
Discussion . . . . . . . . . . . . . . . . . . . . . . . 216
Proposed taxonomic changes and some persistent problems . . . . . . 216
Scale surface microstructure and phylogenetics . . . . . . . . . . 217
Acknowledgements . . . . . . . . . . . . . . . . . . . . 218
Referenrcs . . . . . . . . . . . . . . . . . . . . . . . 218
Appendix 1 . . . . . . . . . . . . . . . . . . . . . . . 221
Appendix 2 . . . . . . . . . . . . . . . . . . . . . . . 224
Appendix 3 . . . . . . . . . . . . . . . . . . . . . . . 231
Appendix 4 . . . . . . . . . . . . . . . . . . . . . . . 232

INTRODUCTION

Iguanian lizards of the tribe Tropidurini occur in the Galapagos Islands and
across much of South America. The group is both ecologically and morphologically
diverse containing members as different as the dorsoventrally flattened crevice
specialist Tropidurus semitaeniatus, arenicolous species like Microlophus thoracicus and T.
hygomi, tree-hole specialists with curiously blunt and armored tails such as [Link],
and the tree-trunk specialist T. plica with its long, spindly arms and flattened body.
Not surprisingly, a complex taxonomic history (see Frost, 1992, for a thorough
review) characterizes this group: until recently, morphologically specialized species
were placed in small genera such as Plica Gray, Strobilurus Wiegmann, and Uracentron
Kaup, and more generalized species were lumped into Tropidurus Wied-Neuwied.
Except for descriptions of many new species, primarily through revisions of the ?:
torquatus and I: hispidus complexes (Cei, 1982; Rodrigues, 1987, 1988; Vanzolini &
Gomes, 1979), relative nomenclatural stability followed publication of Catalogue of
theNeotropica1 Squamata (Peters & Donoso-Barros, 1970).Then, in 1992, Frost proposed
a taxonomy of the Tropidurini consistent with its recovered evolutionary history.
Convincing phylogenetic evidence (Frost, 1992) allowed resurrection of Microlophus
DumCril & Bibron (1 837) to accommodate all but one of the species occurring along
the Pacific coast and Galapagos Islands and required the transfer of species previously
assigned to Plica, Strobilurus, Uracentron, and Tipinurus to Tropidurus. Although originally
described as a subspecies of Microlophus occipitalis, T. 0. koepckeorum Mertens was
plesiomorphic for nearly all characters examined by Frost and was necessarily placed
in a monotypic genus, Plesiomicrolophus Frost. Following these taxonomic changes,
four monophyletic lineages were recognized within the Tropidurini: the cis-Andean
(i.e. east of the Andes) Tropidurus and Uranoscodon with 27 and 1 species respectively,
and the trans-Andean (i.e. west of the Andes) Plesiomicrolophus and Microlophus with
1 and 19 species.
Several factors prompted us to reconsider phylogenetic relationships of the
Tropidurini. Ada-Pires (1995: 142) called for confirmation of Frost’s (1992) results
and preferred to recognize Plica and Uracentron. Frost (1992) suspected that Ple-
siomicrolophus might be the sister taxon of Microlophus, but at the time lacked
synapomorphic evidence to support his conjecture. Finally, we recently discovered
three new species of Tropidurus (Harvey & Gutberlet, 1998) inhabiting a remote
 PHYLOGENY OF TROPIDUKINI 191

mountain range on the Bolivia-Brazilian border. Our observations raised several

questions regarding these species’ biogeography, a subject best explored within a
phylogenetic framework. We noted that T. melanopleurus, T. etheridgei, and T. spinulosus
occur in adjacent habitats along the Central Andes of southern Bolivia and northern
Argentina, and that each of these species appears to be closely related to one of our
new species: respectively T. callathebs, T. chromatops, and T. xanthochilus. Also, we
observed several characteristics that, when viewed in an evolutionary context,
suggested resolution of the problematic placements of T. spinulosus and T. melanopleurus
in earlier phylogenetic hypotheses.
T o address these disparate questions, we undertake a phylogenetic analysis of the
Tropidurini, combining new phylogenetic characters with old and critically evaluating
taxonomic changes proposed in earlier studies. Although we describe three new
macroscopic characters of scalation, surface morphology of the epidermis is our
primary source of new characters. These ‘nontraditional’ characters of epidermal
microstructure have shown promise as a source of phylogenetic information (e.g.
Price, 1982; Williams, 1989; Harvey, 1993), although they have rarely been used
in an explicitly phylogenetic analysis (Peterson & Bezy, 1985; Harvey & Gutberlet,
1995).

RIETHODS

T o recover the evolutionary history of tropidurines we conducted phylogenetic

analyses under the criterion of parsimony (Kluge & Farris, 1969; Farris, 1983; Farris
& Kluge, 1985, 1986) using a total evidence approach (Kluge, 1989). Thus, we seek
a hypothesis of relationships that is the most parsimonious explanation of all available
character evidence.

Source of characters

We examined museum specimens (Appendix 4) to score 18 new characters

described below and 48 of 77 characters identified in earlier studies (e.g. Etheridge,
1970; Rodrigues, 1986; Etheridge & de Queiroz, 1988; Frost, 1992) and summarized
by Frost (1992). The remaining 29 characters used by Frost (1992) are skeletal
features. We are unaware of any new skeletal preparations for species of this group;
skeletons were not prepared for new species included in this analysis. For the
remaining taxa, we used Frost’s data for these 29 characters.
We mounted scales taken from 39 species of tropidurid lizards (Appendix 3) for
scanning electron microscopy (SEM) and compared our observations to published
descriptions of dorsal body scale surfaces of Liocephalus carinatus, L. schreibersi schreibersi,
Stenocercus guentheri, Tropidurus semitaeniatus, and T. sp. (Irish, Williams & Seling, 1988;
Peterson, 1984). T o our knowledge, descriptions of ventral body, supraocular, and
supralabial scale surfaces are nonexistent for tropidurid lizards.
Scales taken from museum specimens were dehydrated in an ethanol series,
sonicated to remove debris, and air-dried in a drying oven set at 8OOC. T o preserve
the integrity of scale organs in the alpha and beta keratin layers (Harvey & Gutberlet,
1995), intact supraocular scales of Uranoscodon superciliosus were critical point-dried
192 hl. B. HARVEY AND R. I,. CUTBERLET JR

in a Samdri PVT-3B critical point dryer. Samples were affixed to EM stubs with
double-stick tape and coated with 200-300 nm of a gold-palladium mixture in a
Hummer VI sputter coater. The scales were examined under a JEOL JSM 35C
scanning electron microscope.
Descriptions of most scale features are based on the beta keratin-containing layers,
removed by pushing on the surface of scales with a dissecting needle (Irish et al.,
1988). Although this method also removes the beta and part of the mesos layers,
descriptions primarily refer to the oberhautchen. [The German noun Oberhautchen
originally used to describe the outermost layer of the squamate epidermis (Schmidt,
1914) was considered anglicized by Irish et al. (1988) and requires neither the
capitalized first letter nor the umlaut.] Intraspecific variability was explored by
examining scales from many individuals of most species (Appendix 3). Scale surface
morphology changes during ontogeny in some lizards (Harvey, 1993; Harvey &
Gutberlet, 1995); we examined adults only for all species except Tropidurus spinulosus,
Stenocercus marmoratus, and S. quenthen’ for which we also examined juveniles. For
most specimens, the beta layers were removed from a supraocular, one or more
paravertebrals, ventrals, and the second or third supralabial. For all species, the
beta layers of some dorsals and ventrals were mounted upside down. Mounting
scales in this manner allowed us to examine the proximal surface of the beta layer
and the inner surface zone (Jackson & Reno, 1975), a crescent shaped strip of
epidermis beneath the caudal edge of scales.

Character coding and polarig

Our coding of a few of Frost’s (1992) characters differs from his and requires
explanation. Females of the highly dimorphic species Tropidurus melanopleurus have
granular paravertebral scales that lack keels, whereas males have strongly keeled,
imbricate paravertebrals. In contrast, paravertebrals of both sexes lack keels in
arenicolous Microlophus. Granularization of the dorsals in female I: melanopleurus is
an expression of sexual dimorphism, and the process of ‘granularization’ appears to
have simultaneously obscured keels. Reduction of these scales is a trait shared with
other Tropidurus, many or all of which have smaller paravertebrals in females than
in males. In our opinion, absence of keels in both sexes is not equivalent to absence
of keels in females only. For this reason, we restrict the definition of character 73
to ‘paravertebral scales of both sexes’ and change the coding of T. melanopleurus from
1 (i.e. in Frost’s, 1992, data matrix) to 0 for this character.
Tropidurus oreadicus lacks an axillary mite pocket (Rodrigues, 1987; Ada-Pires,
1995; and personal observation), although Frost (1992) coded it as having this
derived feature. We have changed its coding from 1 to 0 for character 54 in our
analysis.
Tropidurus spinulosus, T. xanthochilus and several related species possess two rows of
circumorbitals (Fig. 1) identified by their position between the enlarged supraoculars
and median head scales. However, this is not the case in most ?: melanopleurus. One
of 14 T. m. pictus (UTA R-37955) and one out of four 7: m. melanopleurus (USNM
281589) had two complete circumorbital rows. We did not examine any of the
seven specimens of I: melanopleurus listed in Frost’s (1992) ‘Specimens Examined’,
although this character can be assessed for one of these, the specimen pictured in
his figure 42. KU 182998 has one row of circumorbitals on its left eyelid.
 PHYLOGENY OF ‘TROPIUUKINI

10 mm

10 mm

Figure 1. Dorsal head scales of Troopidurus callathebs (Top, UTA 3961 1) and 7: xanthochilu (Bottom,
UTA 38052). Circumorbitals are stippled.

In addition to the number of circumorbital rows, other characters such as 55

(according to Frost, 1992) and 65 (Harvey & Gutberlet, 1998), as well as the new
characters 78 and 79 exhibit polymorphism in some species. Although he did not
explicitly say so, Frost’s (1992) coding of polymorphic characters suggests that he
may have used a criterion of majority (Johnson, Zink & Marten, 1988) or any-
instance (Campbell & Frost, 1993) when coding. The latter appears to be his reason
for coding fiopidurus hispidus as having multiple axillary mite pockets: “some T.
‘hispidus’ from the Guyana Region have doubled ones” (Frost, 1992: 30). The reason
for his use of quotation marks around the specific epithet suggests to us that he
shares our opinion that these specimens may represent a cryptic species similar to
T. hispidus. Avila-Pires (1995) also raises the possibility that T. hispidus may be a
composite of two or more distinct species. Whether character 55 is polymorphic
but state one rare within a single species or whether this variation indicates the
194 M. B. HARVEY AND R. 1,. GUTBERLET JR

presence of a cryptic species, T. hispidus should not be coded as having state one for
this character. The low frequency of two axillary mite pockets in T. hispidus does
not meet the majority criterion, and we code T. hispidus as having one mite pocket
(state zero). After the frequency coding method, the criterion of majority is preferred
to any instance (Wiens, 1995).
Tropidurus azureus, Tjuviceps, and T. itambere are the only species of Tropidurus to
have an interparietal substantially longer than wide (Frost, 1992). However, this
characteristic is polymorphic in T. itambere (see also Harvey & Gutberlet, 1998). In
USNM 148773, the interparietal was slightly wider than long (width 101% of
length), whereas in USNM 148775 it was slightly longer than wide (width 82% of
length). Rodrigues (1987) describes this species as having an “occipital de forma
irregular gerulmente mais longa que larga” (emphasis ours). We have no idea how
common narrow interparietals are in this species (few specimens were available for
study). However, assigning either a state of 1 or 0 to character 68 does not affect
the placement of T. itambere in the phylogeny.
T o deal with polymorphism in characters 65, 78, and 79 we used the frequency
coding method developed by Wiens (1993, 1995). Our choice of this method is
based on the results of simulation and congruence studies of its accuracy (e.g. W e n s
& Servedio, 1997) which suggest that it recovers the correct phylogeny more often
than other parsimony coding methods. Characters 65, 78, and 79 were assessed for
all of the specimens included in this study, and the observed frequencies of their
states are presented in the data matrix. Following the methodology of Wiens (1995),
frequencies were divided into lettered bins, each spanning 4% of the total range of
possible frequencies (e.g. a = 0-3.99%, b = 4-7.99%, etc.). Each species was assigned
a letter ‘a’ to ‘y’ based on the observed frequency of the derived trait, and traits
were ordered from trait absence (a) to fixation (y). T o compensate for the addition
of a large number of steps added to trees constructed using the frequency method
(24 when going from state a to state y), researchers have used weighting techniques
(e.g. Wiens, 1993, 1995; McGuire, 1996; Gutberlet, 1998). In our analysis, we
assigned a weight of 24 to nonpolymorphic characters and one to polymorphic
characters (Wiens, 1995). Thus, fixation of a derived state for polymorphic characters
adds the same number of steps as transition from a primitive to derived condition
of a nonpolymorphic character. The frequency approach potentially reduces noise
and the effects of sampling error by giving minimal weight to conditions occurring
at low frequencies (Swofford & Berlocher, 1987; Wiens, 1995).
Missing data entries in our matrix mean that (1) the data are unavailable, such
as skeletal characters from rare species, or (2) the characters are not applicable,
such as the condition of mite pockets in species that do not have them. Coding taxa
as having missing characters when the characters are not applicable can result in
assignment of impossible ancestral states (Platnick, Griswold & Coddington, 1991;
Maddison, 1993). However, such coding may only present a problem when taxa
with the nonapplicable states are in two or more regions of the cladogram (Maddison,
1993). One solution to this problem, assigning a separate state for ‘not applicable’
(Platnick et al., 1991), could potentially result in clades being diagnosed by ‘not
applicable’, and thus might cause more problems than it would solve. In our analysis
we were able to eliminate some nonapplicable entries by combining Frost’s character
48 with 49 and character 54 with 55. The new transformations for these characters
are then:
 PHYLOGENY OF TROPIDURINI 195

+
48 49. Antegular-oblique neck fold mite pockets, condition: (0 = 48{ 0)) weak single
mite pocket; (1 =48{ 1)) no mite pocket, complex neck folding; (2 =49{0}) two mite
pockets in the oblique neck fold; (3 = 49{ 1}) a single well-developed mite pocket in
the upper position of the oblique neck fold; (4=49{2)) single, very enlarged mite
pocket in the upper position of the oblique neck fold; (5 = 48{3}) ventrolateral mite
pockets; (6 =48{4}) no obvious mite pocket, aithough weak depressions are in the
ventrolateral side of the antegular fold.
+
54 55. Axillary pocket: (0 = 54{0)) axillary pocket absent; (1 = 55 { 0)) present,
single; (2 =55( 1)) present, multiple, usually 3, sometimes only 2.
The ancestral state of character 54 is 0 and that of 55 is unknown because absence
of axillary mite pockets is characteristic of Laocephalus and plesiomorphic in Stenocercus
(Frost, 1992). For this reason we coded zero as the ancestral state for our combined
character.

We were unable to combine some of our new characters in such a way that these
characters would be applicable to all species. With the exception of character 86
(see below), we decided not to assign a separate state for ‘not applicable’ in favour
of the traditional solution of simply coding these species as unknown for the character.
Whether multistate characters should be treated as ordered or unordered is
controversial (Hauser & Presch, 1991; Slowinski, 1993). Of Frost’s (1992) 77
characters, 16 are multistate. He treated characters 17, 41, 42, 48, 50, and 71 as
unordered and, treated the other multistate characters (6, 12, 29, 33, 35, 47, 49,
63, 70, 75) as ordered based on a criterion of morphological intermediacy. Such an
approach requires an ad hoc assumption that evolution of these characters proceeds
orthogenetically(Hauser & Presch, 199l),a potentially spurious model of evolutionary
change (Mabee, 1989) that assumes phyletic gradualism (see Eldredge & Gould,
1972).
In effect, such ordering potentially dilutes phylogenetic signal by increasing
homoplasy in the data set. On the other hand, by treating characters as unordered,
some legitimate synapomorphiesmay be lost (Kluge, 1991, 1993; Campbell & Frost,
1993).Campbell & Frost (1 993) provide a convincing example of this problem when
they discuss a hypothetical transformation series (0) no antlers + (1) small antlers
-+ (2) large antlers. As they explain, treating such a character as unordered would
potentially assign no cost to antlers evolving twice. That is to say, the synapomorphy
‘antlerspresent’ would be excluded from the analysis. They proposed partial ordering
as a solution to this problem.
+
Among the multistate characters in our analysis, characters 47, 48 49, 54 55, +
and 75 are composed of states similar to the antler example: state zero for each
character represents absence of a characteristic and derived states represent both
the character’s presence and condition. Each of these characters was partially
ordered (i.e. 0 + [ 1 ++ 21) in our analysis by having one transformation for presence/
absence and one unordered character set for ‘condition of presence’. To prevent
‘presence’being weighted twice, absence was coded as unknown (Campbell & Frost,
1993) in the characters for condition (Appendix 2). We do not see that the antler
analogy fits the remaining multistate characters, and they were left unordered.
However, a posteriori we mapped these characters onto our phylogenetic hypothesis
to evaluate how ordering would have affected the outcome. States 2 for characters
6 and 29 become autapomorphies of Tropidum semitaeniatus. Ordering them would
have added two synapomorphies to the otherwise well supported clade linking this
196 hI. K. HARVEY AND K. L. GUTBERLEI‘JK

species with T. bogerti. State 2 for character 63 is unique to T. plica; this species’s
placement on the phylogeny is not affected by treating the character as unordered.
Frost’s hypotheses for ordering the remaining characters, 12, 33, 35, 70, are
congruent with our hypothesis, thus ordering them would not change the results.
For each of our new characters, a hypothetical ancestral state was assigned by
assessing the characters for eleven outgroup taxa (Appendix 1). When several states
were present among the outgroups we used the argumentation of Maddison,
Donoghue & Maddison (1984) to determine the ancestral state. T o that end, states
were mapped on a phylogeny of the outgroups and ingroup consistent with earlier
studies (Etheridge, 1995; Etheridge & de Queiroz, 1988; Frost & Etheridge, 1989;
Pregill, 1992) of tropidurid relationships: ((Liolaemus archeforus, L. chiliensis, L. elongatus)
(((Leiocephaluspersonatus, L. semilineatus) L. schreibersz) ((Stenocercuscaducus, S. jimbriatus, S.
quentheri, 5’. iridescens, [Link]) (Tropidurini)))).

Phylogenetic anabsis

Our data matrix contains 29 taxa and 99 characters. We collected scale surface
data for Uranoscodon superciliosus,five species of Microlophus, Plesiomicrolophus koepckeorum,
and 2 1 species of Tropidurus. Frost (1992) reported that several species were identical
to one another for the characters he used. Although we did not examine Microlophus
atacamensis, we did examine M. theresiae, and this species is substituted for the former
species in the analysis. Tropidurus cocorobensis was not available to us but is identical
to T. etheridgei and T. tygomi except for unknown character states. The latter two
species are also identical to T. chromatops as characterized in the data matrix and
excepting osteological characters which are unknown for T. chromatops. In our analysis,
we include only T. hygomi among these four very similar species. We coded our new
characters as unknown for species not examined by us.
The data were analysed using PAUP 3.1.1 (Swofford, 1993). Frequency coding,
partial ordering, and addition of many unknown cells to the data matrix resulted
in a large increase in the number of trees and time for the analysis. For this reason,
only heuristic algorithms were used on the complete data matrix. We performed 24
random addition sequence replicates with the collapse zero-length branches and
steepest descent options disabled and tree bisection-reconnection activated.
Several species were incomplete for osteological, scale surface, or both types of
characters. In a series of subsampling experiments (Wiens & Reeder, 1995), addition
of taxa with incomplete data caused a small decrease in similarity of estimated trees
to the true phylogeny, but allowed a phylogenetic hypothesis to be postulated for
the incomplete taxa as opposed to having no hypothesis at all. In this study, we
performed two analyses of the data. Using heuristic methods we analysed the
complete set of taxa, then using the branch and bound algorithm (Hendy & Penny,
1982)we analysed a reduced set of taxa with Microlophus bivittatus, Tropidurus callathebs,
7: eythrocephalus, T. insulans, T. mucujensis, and T. xanthochilus removed.

Evaluation of confidence and phylogenetic signal

We used the g,-statistic (Hillis, 1991; Hillis & Huelsenbeck, 1992; Huelsenbeck,
199l), to see if our data matrix contained levels of covariation among characters
 PHYLOGENY OF ‘IROPIDURINI 197

(i.e. phylogenetic signal) significantly greater than expected for random data sets.
Because of the large number of taxa and characters, we calculated the g,-statistic
from a tree-length frequency distribution using a random sample of 10 000 trees
rather than all possible trees. Significance levels of the g,-statistic are dependent
on number of taxa, number of characters, and number of character states (Hillis,
1991). Hillis & Huelsenbeck (1992) pointed out that direct calculation of
confidence limits for the g,-statistic avoids assumptions about its underlying
distributions for random data sets of different dimensions. They went on to
provide tables of critical values for data sets of different sizes and numbers of
character states. We did not directly calculate the confidence limits for our test
of skewness, because our calculated g,-statistic was much more negative (= more
significant) than any of Hillis & Huelsenbeck’s (1992) tabulated critical values
and because the size of our data matrix made generation of critical values from
randomization of the data set intractable.
Our confidence about the relationships of specific clades of tropidurines is based
on the probability that a specified group is contained in the true phylogeny (Hillis
& Bull, 1993) as determined using nonparametric bootstrapping (Felsenstein, 1985).
Using simulations and a known phylogeny for the T7 bacteriophage, Hillis &
Bull (1993) convincingly demonstrated that nonparametric bootstrapping provides
biased but conservative estimates of phylogenetic accuracy. High bootstrap values
typically underestimate accuracy, whereas low values overestimate accuracy.
For the bootstrap analysis, 100 pseudoreplicated data sets based on the reduced
data set were analysed using heuristic searches, each with 100 sequence replicates.
Hillis & Bull (1993) found that bootstrap values of 70% or greater generally
correspond to 95% or greater probability of a clade being correctly resolved.
In the results, the consistency, rescaled consistency, and retention indices (Farris,
1989) are reported for our data but are not interpreted.

SCALE SURFACE MORPHOLOGY OF TROPIDURIDS

Coarse scale suface microstructure

Scale surface morphology has rarely been incorporated into phylogenetic analyses,
and many workers in iguanian systematics may not be familiar with scale surface
terminology. In particular, few descriptions of tropidurid scale surfaces have been
published (Peterson, 1984; Irish et al., 1988). Our purpose here is to summarize
characteristics of scale surface microstructure that aid in species recognition and in
establishing phylogenetic relationships among tropidurids. We do not attempt a
more in depth description of these features’ morphology but refer the reader to
published descriptions of similar features.
Two types of high relief coarse microstructure occur in tropidurids. Macro-
honeycomb (see Harvey, 1993; Harvey & Gutberlet, 1995 for detailed descriptions
of this type of morphology) consists of folded keratinized layers of the epidermis
and is the most common type of coarse microstructure among tropidurids. Often,
though not always (Harvey & Gutberlet, 1995), individual macrohoneycomb cham-
bers are overlapped by single oberhautchen cells whose margins may be visible
198 M. B. HARVEY AND R. L. GUTBERLET JR

Figure 2. A & B, paravertebral of Uranoscodon superciliosus, arrow indicates position of a lenticular scale
organ on keel, frame in (A) surrounds area enlarged in (B). C, sublamellar cell margins of the distal
oberhautchen on the inner surface zone of a dorsal taken from Tropidurus callathebs. D, modified
macrohoneycomb of T aeureus. E, supralabial of Tmpidurus melanopleums pictus. F, supralabial of T
xanthochilus.

along the apex of ridges (Figs 2D, 6E). Only among species of LeRioce$hzlus within
the Tropiduridae, macrohoneycomb is replaced by ‘hillocks’ (Fig. 4A-D, see also
Irish et al., 1988).To the exclusion of macrohoneycomb, hillocks occur on vertebrals,
paravertebrals, ventrals, labials, supraoculars, and inner surface zones in species of
this genus.
Coarse microstructure is usually less distinct on ventrals, and macrohoneycomb
on labials and supraoculars is usually non-imbricate or weakly so (Figs 6E, 7E,F) in
 PHYLOGENY OF TROPIDURINI 199

Figure 3. Macrohoneycomb from the hinge regions of (A) Trobidums stmbilums, (B) T. bgomi, and (C)
T onadicus illustrating structure of caudal wall and degree of imbrication.

contrast to the highly imbricate macrohoneycomb on other scales (Figs 2D, 3A,B,
5).
Tropidurids may be distinguished from one another by the shape of individual
macrohoneycomb chambers and by the distribution of this morphology over different
epidermal surfaces. Stenocercus and cis-Andean Tropidurini have high macro-
honeycomb similar to that of xenosaurids and cordylids, whereas the macro-
honeycomb of Liolaemus and the trans-Andean taxa Microlophus and Plesiomicrolophus
is very low (Fig. 4E,F). Both types of macrohoneycomb are modified across the
hinge region (Figs 4F, 7A), the anterior edge of the scale where the scale is overlapped
by its cephalad neighbour. In the hinge region of dorsals and ventrals, high walls
do not completely enclose tropidurid macrohoneycomb chambers (Fig. 3), although
this is often the case near the caudal edge of these scales. Rather, the caudal wall
of chambers in the hinge region is flat; the only exception being Tropidurus oreadicus
which has juxtaposed chambers across the entire surface of its dorsals and in the
hinge region of its ventrals (Fig. 3C). In some species, the oberhautchen of each
chamber in the hinge extends as a jagged projection over the next caudal chamber
(Figs 2D, 3A), whereas in the vast majority of species this is not the case and the
anterior wall appears rounded and smooth (Fig. 3B). In T. umbra and apparently
Uranoscodon superciliosus, a jagged projection of the oberhautchen is absent from most
of the hinge but appears between macrohoneycomb chamber rows 10-15 (counted
next to keel). Unique to Stenocucus marmoratus and U. superciliosus among the tropidurids
we examined, chamber walls are irregularly fragmented over most of the middle-
third of paravertebrals (Fig. 2A,B) but intact elsewhere on these scales.
Macrohoneycomb chambers are poorly defined to absent on the central and
caudal portions of dorsals in many tropidurids (Figs 4E, 5A); but in some species,
macrohoneycomb chambers are clearly defined on all areas of paravertebral scales
except the distal surface of keels. Some taxa such as Liolaemus and the arenicolous
species of Microlophus appear to take the former condition to an extreme by having
low, rectangular macrohoneycomb present only in the hinge region of all scales
(Fig. 4E,F'). Elsewhere on the scale, juxtaposed polygonal cells are visible, but visible
ridges are absent in these species.
In all taxa studied, supralabials exhibit at least some coarse microstructure.
Macrohoneycomb or hillocks are restricted to edges of the scale other than that
200 M. B. HARVEY AND R. L. GUTBERLET JR

Figure 4. A-D, Leiocephalus schreibersi. A, supraocular, arrow indicates narrow region of a keel enlarged
in (B). B, oberhautchen cells along apex of keel covered in shallow pits, cells lateral to keel covered in
spinules. Coarse (C) and fine (D) structure of a paravertebral scale. Arrows indicate fused spinules like
those present in some Stenocercus. E & F, dorsal of Liolaemus chiliensis. Frame encloses area enlarged in
(F). Arrows indicate folding of beta keratin layers to form low macrohoneycomb in hinge region of
scale.

bordering the mouth (Fig. 2E) in most tropidurids. Except for the area immediately
bordering the mouth, macrohoneycomb covers all of individual labials in highly
derived species of Tropidurus (Figs 2F, 7 E), many Stenocercus, and Uranoscodon superciliosus.
Trapidurns nanuzae exhibits an intermediate condition, having irregular patches of
macrohoneycomb extending across portions of the lower two-thirds of supralabials.
 PHYLOGENY OF TROPIIIUKINI 20 1

Figure 5. Fine microstructure on a paravertebral scale of Tr@durus callathebs. Letters in (A) refer to
pitted region enlarged in (B), pit-and-groove region enlarged in (C), and spinulate region cnlarged in
PI.

Intraspecific variation in this character was never observed even when scales from
as many as sixteen individuals were mounted for scanning electron microscopy.
The distribution of coarse microstructure on supraocular scales is also variable
among taxa. Although not exactly, this variability is nearly congruent with the
distribution of macrohoneycomb over supralabials. Coarse microstructure is restricted
to scale edges in Liolaemus, Microlophus, Plesiomicrolophus,Stenocercusguentheri, S. marmoratus,
Tropidurus hispidus, T. bgomi, T. oreadicus, T. semitaeniatus, and T. torquatus. With the
exception of smooth areas along the crests of macroscopic ridges, coarse mi-
crostructure covers the entire surface of supraoculars in Leiocephalus, S. caducus, S.
jimbriatus, S. iridescens, T. azureus, I: etheridgei, Tjauiceps, [Link], 7: strobilurus, 7:umbra,
and Uranoscodon (Fig. 1OA,F).
In many lizards, ventral scales lack macrohoneycomb over most of their surface
(Harvey, 1993; Harvey & Gutberlet, 1995),and such is the case with most tropidurids.
Usually, macrohoneycomb occurs only over the hinge region of ventrals (Fig. 7A).
However, in some Stenocercus, in Uranoscodon, and in Tropidurus previously placed in
Plica and Uracentron, macrohoneycomb extends across the entire surface of ventrals
(Fig. 7B). Notably, fiopidurus strobilurus lacks this derived condition.
The crescent shaped strip of epidermis beneath the caudal edge of imbricate
scales (hereafter referred to as the inner surface zone, following the terminology of
Jackson & Reno, 1975; Figs 2C, 9B) overlies and contacts the hinge region of the
202 M. B. HARVEY AND R. L. GUTBEFUET JR

next scale in the direction of scale imbrication. The inner surface zone has never
been described in any detail using scanning electron microscopy (see however
photos in Cole & Van Devender, 1976; Harvey & Gutberlet, 1995); however, its
microstructure differs from that on the outer surface and hinge region of scales in
several squamate lineages (pers. observ.). Among tropidurids, the inner surface zone
always bears coarse microstructure. Macrohoneycomb extends from the exposed
surface of imbricate scales to completely cover the inner surface zone of dorsals in
many Microlophus, Plesiomicrolophus, some Stenocercus, some Tropidurus, and Uranoscodon
superciliosus. At the opposite extreme, lamellae completely cover the inner surface
zones of Liolaemus, Stenocercus guentheri, T. melanopleurus, T. strobilurus, and Tmpidurus
callathebs. In the other species, a transition from low-walled macrohoneycomb to
lamellae occurs from the caudal edge of the inner surface zone to its cranial edge.
In these species exhibiting the transitional morphology, some intra- and interspecific
variation exists in the relative amounts of macrohoneycomb and lamellae. For
instance, macrohoneycomb on inner surface zones of dorsals in T. xanthochilus is
usually more extensive than in the other taxa with transitional morphology. Similar
trends are observed on the inner surface zones of ventrals although they generally
tend to have lower macrohoneycomb and a higher frequency of lamellae.

Fine scale surface microstructure

The epidermis of iguanian lizards exhibits a variety of subcellular features such

as setae, spinules, and pits. Terminology for these features varies among authors.
Our terminology is that of Peterson (1983) and Peterson & Williams (1981) who
established a nomenclature for the spinule-setae-spike morphotypic series (sensu
Olson, 1975) on the digital pads of polychrotids. Detailed morphological descriptions
of these structures have been published by several authors (see especially Peterson,
1984, and citations therein).
All tropidurids have a pitted epidermis that usually is also covered in spinules on
at least some scale surfaces. In some Tropidurus and all the Liolaemus, Microlophus, and
Stenocercus we examined, a morphotypic series of several microstructural morphologies
is present on, at least, paravertebrals and supraoculars. The edge of scales is entirely
or almost entirely pitted (Fig. 5B). Approaching the centre of the scale, the pitted
region grades into a mix of pits, pit and groove, and d f i s e spinules, followed by a
region where spinules are often very dense (Fig. 5C,D). The relative extent to which
scales are covered by these different morphologies varies among species. In many
Tropidurus, spinules appear within the cranial-most row of macrohoneycomb and
extend across the entire scale, leaving only the edges bare of spinules. At the opposite
extreme, Stenocercus iridescens has pitted and pit and groove microstructure extending
up to the caudal margin of dorsals where dense spinules appear in a relatively
narrow band along the scale’s edge. Some species, such as Tropidurus nanuzae, appear
to completely lack spinules.
This transitional morphology may characterize every scale on the body as is the
case in most Tropidurus, or it may be restricted to scales from one or more body
regions. For example, Microlophus has partially spinulate dorsals, supralabials, and
supraoculars but uniformly pitted ventrals, whereas both the ventrals and supralabials
are uniformly pitted in Plesiomicrolophus.
In Lhocephalus, densely distributed spinules occur over dorsals, dorsal head scales,
 PHYLOGENY OF TROPIDURINI 203

supralabials, ventrals, and inner surface zones. At the tops of hillocks, spinules are
usually pointed into thin spikes, and in the ‘valleys’ between hillocks, groups of
spinules tend to unite distally (Fig. 4D). The hillock morphology of L. schreibersi is
flattened on the surface of keels and in isolated patches on the caudal two-thirds of
dorsals; these areas lack spinules and are uniformly pitted (Fig. 4B). Pit and groove
microstructure appears to be absent from the epidermis of L. schreibersi.
Newly hatched Tropidums spinulosus have quite different microstructural patterns
than do adults. Scales taken from a juvenile with a snout-vent length of 37mm
(UTA R-3459 1) lack spinules (Fig. 6F). Instead, pit and groove microstructure covers
the entire scale. We did not observe such dramatic differences between scales taken
from different adult specimens. In snakes, ontogenetic shifts in scale morphology
are documented for a diverse array of species (Price & Kelly, 1989). The shift in
snakes occurs with the first shed; however, this may not be the case in some other
squamates (e.g. Shinisaurus) where some neonatal morphologies apparently persist
through several sheds (Harvey, 1993).
Both Uranoscodon superciliosus and Tropidurus plica exhibit a fine microstructure of
large, widely spaced papillae occurring on all scales and extending to scale inner
surface zones. Papillae may be rounded distally (Fig. 6B) or attenuated into spikes
(Fig. 6D) and both shapes typically occur in the same individual of either species.
Possibly, these papillae form through the fusion of spinules as occurs in Leiocephalus
schreibersi and many polychrotids (Peterson & Williams, 1981; Peterson, 1983). This
appears to be supported by irregular sizes of papillae and spinules on some scale
surfaces in TT: plica such as those pictured in Figure 6A,B. However, the distribution
of papillae is opposite that of spinules. In TT: plica where both morphologies occur,
the centres of scales are covered in diffuse (on dorsals) to dense (on supraoculars)
spinules, whereas papillae are restricted to the borders of scales. Papillae are even
densest on the cranial edge of dorsals, anterior to the first row of macrohoneycomb.
This region is almost always devoid of spinules in even the most heavily spinulate
tropidurids.

Scale organs

Multiple lenticular scale organs are present on head scales of all species examined
(Figs 2E,F, 4A, 7E,F, 8, 10). These organs consist of a dermal papilla, the sense
organ ‘column’ of Jackson & Sharawy (1980) (Fig. 10D,E), that extends up through
keratinized layers of the epidermis to the scale surface. The oberhautchen of
tropidurid scale organs is occasionally pitted or has ‘ghost images’ of the clear layer
(Stewart & Daniel, 1972);it lacks a hair-like sensilla, coarse microstructure, spinules,
and pit and groove. Scale organs in tropidurids do not project substantially above
the surrounding surface of the oberhautchen as occurs in some cordylids (Harvey
& Gutberlet, 1995) and xenosaurids (Harvey, 1993). Substantial lateral expansion
of the alpha layers like that of scale organs in some Cordylus (Harvey & Gutberlet,
1995) was not observed (Fig. 10).
Typically, these organs appear to be randomly distributed over the surface of
individual head scales (Fig. 8), but some head scales bear decidedly nonrandom
distributions of scale organs. On supraoculars of Lezocephalus, most scale organs are
positioned at the termini of parallel, macroscopic ridges (Fig. 4A). Scale organs on
supraoculars of Tropidums azureus, T javiceps, TT: itambere, ?: umbra, and Uranoscodon
204 M. B. HARVEY AND R. L. GUTBERLET JR

Figure 6. A & B, fine microstructure of a paravertebral taken from Tmjidurusplica. Frame indicates deep
macrohoneycomb chamber enlarged in (B). Arrows indicate papillae. C & D, deep macrohoneycomb and
papillae of Uranoscodon superciliosus. Frame in (C) encloses chamber enlarged in (D). E, spinulate
macrohoneycomb of adult Tropidurus spinulosus. Arrow indicates low ridge left from clear layer, not to
be confused with prominent cell margins of the heptagonal oberhautchen cells lying atop the ridges
of the macrohoneycomb. F, pit and groove pattern of hatchling T s/inulosus.

superciliosus are almost completely restricted to the surfaces of macroscopic ridges

(Fig. lo), whereas Tropidurus lacking rugose supraoculars usually have scale organs
distributed randomly across the scale. Scale organs of Liolaemus, hiocephalus, Micro-
lophus, and Plesiomicrolophus we examined were in a line along the caudal edge of all
enlarged supraoculars (Fig. 8D). Very rarely, scale organs in these species appeared
away from the caudal edges of these scales. Scale organs on other dorsal head scales
also tend to be positioned along the caudal edge of the scale in these same taxa.
 PHYLOGENY OF TROPIDURINI 205

Figure 7. A, ventral of Tro~idumsxanthochilus showing macrohoneycomb restricted to the hinge region,

absence of a keel, and lenticular scale organ (arrow). B, midventral scale of I: umbra. Arrow indicates
one of two scale organs flanking caudal end of large keel. C, paravertebral of 7: umbra showing location
of large scale organ atop keel. D, dorsals of ?: chromatops showing position of scale organs. Scale to
right was taken from specimen’s right, scale on left from specimen’s left. E & F, lenticular scale organs
and macrohoneycomb on a supralabial taken from I: umbra. Arrow indicates scale organ enlarged in
(0

Number of scale organs on individual head scales can vary considerably within
a species or even the same specimen (Harvey & Gutberlet, 1995; Matveyeva &
Ananjeva, 1995). Some Liolaemus, arenicolous species of Microlophus, and some
Stenocercus have obvious reduction in the number of scale organs on supraoculars.
206 M. B. HARVEY AND R. L. GUTBERLET JR

A 3 mm C 3 mm

B 3 mm D 3 mm

Figure 8. Snout scales of (A) Tropidums culluthebs (UTA 39612) and (B) T rnelunopleurus (UTA 37951)
illustrating distribution of lenticular scale organs and canthal-preocular contact (stipple). Supraocular
region of (C) Tropidurus culluthebs (UTA 39622) and (D) Microlophus occzpitulk (TCWC 28388) showing
distribution of lenticular scale organs and enlarged supraoculars (stipple).

At most, supraoculars in these species have five scale organs with most scales having
one or none.
Among tropidurids, scale organs are also located at the caudal edge of para-
vertebrals. Scale organs are usually absent from vertebral scales and were observed
only on some smaller vertebrals on the caudal half of the body in Tropidurus umbra.
Species of Leiocephalus and Stenocercus may have three to five scale organs on a single
paravertebral; but, in tropidurines, scale organs occur in one of four places: (1) at
 PHYLOGENY OF TROPIDURINI 207

Figure 9. Caudal notch in ventrals of Mimlophus occipitalis viewed on the outer (A) and inner (B) surface
zones. Arrow in (B) indicates position of the terminal lenticular scale organ directed into the notch.

the caudal tip of the scale, (2) to the left of the keel, (3) to the right of the keel, or
(4) on both sides of the keel (Fig. 7B,D). Scale organs may also be absent from
individual paravertebral scales. The position of scale organs on paravertebrals is not
random. Organs at the caudal tip of scales occur on paravertebrals near the dorsal
midline or on scales that are not mucronate or only weakly so, such as those of
Stenocercus mannoratus, Tropidurus azureus, and females of I: melanopleurus. Scale organs
to the left of the keel occur on paravertebrals on the left side of the body. Scale
organs located to the right of the keel occur on paravertebrals on the right side of
the body. Paravertebrals with paired scale organs or no scale organs may be
interspersed among scales that follow the above pattern. Zopidurus plica, I: umbra,
and Uranoscodon superciliosus possess large, flat scale organs positioned medially on
the keels of paravertebrals (Figs 2A, 7C). These organs are circular or have an
irregularly rounded contour (Fig. 7C). On one specimen of I: umbra, two of these
organs were observed on the same scale-one in the typical position and another
cranial to it on the same keel.
Scale organs are present on some ventral scales in all taxa studied. Tiny scale
organs (Fig. 7A) are present at the caudal tip of ventral scales in most species.
Tropidurus umbra and Uranoscodon have well-developed scale organs located to one
side or on both sides of a central keel (Fig. 7B) present on each of their ventrals. As
with dorsal scales, there is a consistent pattern: ventrals on the right side of the
midventral line have scale organs to the right of the keel; ventrals on the left side
of the midventral line have scale organs to the left of the keel. Ventrals with paired
scale organs are interspersed among the scales that follow this pattern.
Species of Microlophus and Plesiomicrolophus koepckeorum were unique among the
species we examined in possessing a caudal notch (Fig. 9) in some ventral body and
femoral scales. Apparently, the notch is associated with the terminal scale organs
on these scales. When present, the notch extends to the usual position for the scale
organ which may be absent or directed caudally into the notch (Fig. 9B). Notches
are most clearly visible in the beta-keratin containing layers but also extend for a
short distance into the alpha layer. In some species such as M. occipitalis, M. ti@,
M. thoracicus, and R koepckeorum, most ventrals bear a notch, whereas in others, M.
stolzmanni, M. peruvianus, and M. theresiue, very few ventrals bear notches. Two of the
latter species are arenicolous and have mostly rounded ventrals. In these species,
208 M. B. HARVEY AND R. L. GUTBERLET JR

Figure 10. A-E, distal oberhautchen (A), proximal beta layer (B,C), and distal alpha layer (D,E) of a
strongly rugose supraocular taken from Uranoscodon superciliosus. Arrow in (B) indicates hole in beta
indicates structure enlarged in (E), the alpha layer covering a long,
layer enlarged in (C). Arrow in (D)
tubular dermal papilla. Dermal papillae fit into holes in the beta layer. F, weakly rugose supraocular
of Tropidurus umbra.

the notch is most frequent in more pointed scales such as the patch of scales just
caudal to the sternum in M. theresiae.

TRANSFORMATION SERIES OF NEW CHARACTERS

The variation described in the preceding section on scale surface morphology

provides the basis for 15 new phylogenetic characters. T o those 15 we add three
 PHYLOGENY OF TROPIDURINI 209

A 5 mm B 5 mm

Figure 1 1. Contact between postmentals and infralabials (stipple) in (A) Tmbidurus callathe&s (UTA
39612) and (B) 7: chrornatoju (UTA 39603).

macroscopic characters: two characters of squamation found to be useful in diagnosing

new species in two earlier studies (Manzani & Abe, 1990; Harvey & Gutberlet,
1998) and variation in palpebral scale morphology discovered while we examined
the distribution of scale organs. In the following transformation series, states 0 and
A are primitive unless the character is unpolarized. Complete descriptions of
characters 1-77 were given by Frost (1992), and his characters are summarized in
Appendix 1.
78. Preocular-canthal contact (Fig. 8): (A) posterior canthal contacting preocular;
(B-X) polymorphic frequencies; (Y,! canthal and preocular separated by small
rectangular scale that is part of series separating superciliaries from upper eyelid; (?)
character not applicable or unknown (=species not examined by us).
Preocular-canthal contact has been used to distinguish between species of Tropidurus
(Manzani & Abe, 1990; Harvey & Gutberlet, 1998). Among some of the more
derived species of Tropidurus, the canthals and preocular are separated by one or
more smaller rectangular scales lying between the eye and superciliaries. Whereas,
among all the other tropidurids we examined contact between these scales rarely
occurs. This character is difficult to assess in species having a fragmented preocular.
In Tropidurus azureus, I:javiceps, and I:plica scales between the preocular series and
the canthals are shaped like the loreals or scales on the eyelids, and we coded these
species as having state Y. In Uranoscodon superciliosus and T. umbra all the scales in
this region are so fragmented that the character is not applicable or at least impossible
to assess.
79. Postmental-infralabial contact (Fig. 1 1): (A) one or no postmentals contacting
infralabials; (B-X) polymorphic frequencies; (Y,! two or more postmentals contacting
infralabials; (?) not applicable or unknown.
Among most tropidurids, a single postmental contacts the infralabial series, but
in Eopidurus melanopleurus and I: callathebs the postmentals are angled more obtusely
than in other species, and the sublabials do not extend as far anteriorly so that 2-3
(rarely 1 or 4) postmentals contact the infralabial series. Tropidurus bogerti, I: hispidus,
and I: semitaeniatus are polymorphic for this character, with about half of the
specimens we examined having the derived state. In most other tropidurids, a second
postmental rarely contacts the infralabials, but the first postmental almost always
does. Some specimens of Tropidurus torguatus exhibit an extreme condition of state A
where the sublabials completely separate the postmental and infralabial series.
The postmental series is so fragmented in Tropidurus plica and I: umbra that this
210 M. B. HARVEY AND R. L. GUTBERLET JR

character is not applicable. Tmpidurus atureus and [Link] usually have more than
one postmental contacting the infralabials; however, the postmentals are fragmented,
have an unusual shape, and are not angled obtusely as in T. melanopleurus and ?:
callathelys. We also chose to treat this character as inapplicable to these species.
Occasional specimens (e.g. four of 26 Plesiomicrolophus koepckeorum) exhibited very
narrow contact between the second postmental and infralabial. We did not consider
this to be equivalent to the derived condition of this character.
80. Type of coarse microstructure (Figs 4E,F, 5): (0) very low, rectangular
macrohoneycomb; (1) well developed, imbricate macrohoneycomb.
Two well defined states exist with the former characterizing Plesiomicrolophus and
all Microlophus we examined and the latter characterizing Uranoscodon and all Tropidurus
we examined. Applying the argumentation of Maddison et al. (1984), this character’s
polarity is found to be equivocal, because each outgroup has a different type of
coarse microstructure: low macrohoneycomb in Liolaemus, high macrohoneycomb
in Stenocercus, and hillocks in Leiocephalus.
81. Caudal wall of macrohoneycomb on dorsals (Fig. 3): (0) caudal wall lacking
a jagged projection of the oberhautchen; (1) caudal wall having a jagged projection
of the oberhautchen on at least some portions of the scale.
In most species having the derived condition (Tropidurus azureus, ‘7: jlaviceps, 7:
nanutae, T. plica, and I: strobilurus),jagged areas are present in the hinge but lost
posteriorly. In Tropidurus umbra jagged walls are absent from the hinge but appear
on several rows of macrohoneycomb beginning between rows 10 and 15. The same
appears to occur in Uranoscodon; however, the jagged walls are not clearly visible in
this species because this area was dirty in all our samples. Also, this area lies in the
heavily fractured region unique to this species and Stenocercus marmoratus, further
complicating assessment of this character.
82. Distribution of macrohoneycomb on dorsals: (0) poorly developed to absent
on caudal two-thirds of scale; (1) well developed over entire surface of dorsals except
for distal surface of keel.
83. Distribution of coarse microstructure on supralabials (Figs 2E,F, 7E): (0) coarse
microstructure distributed in a narrow band along dorsal edge of the scale; (1) coarse
microstructure distributed across most of the scale surface, edge bordering the buccal
commissure smooth.
The labial scale surface is smooth except for a narrow band of coarse microstructure
along the dorsal, cranial, and caudal edges of the scale in all Liolaemus and Leiocephalus
we examined, but both character states occur in Stenocercus. Applying the polarity
argumentation of Maddison et al. (1984), extensive covering of supralabials by
macrohoneycomb is derived independently within the Tropidurini and Stenocercini.
84. Distribution of coarse microstructure on supraoculars: (0)coarse microstructure
restricted to the edge of supraocular scales, dorsal surface lacking coarse micro-
structure; (1) coarse microstructure extending across entire surface of supraocular
scales except for the surfaces of macroscopic ridges and keels.
This character may not be independent from character 83; however, the occurrence
of smooth supralabials and smooth supraoculars is not entirely congruent in tro-
pidurids. All species having macrohoneycomb across most of their supralabials also
have macrohoneycomb across their supraoculars. However, the reverse is not true.
Polarity of this character is equivocal because coarse microstructure is low (i.e.
state A) on the supraoculars of Liolaemus and high on those of Lhocephalus. Both
character states occur among species of Stenocercus.
 PHYLOGENY OF TROPIDURINI 21 1

85. Distribution of coarse microstructure on ventrals (Fig. 7A,B): (0) coarse

microstructure confined to hinge region of scale; (1) coarse microstructure distributed
over most or all of the scale.
86. Type of coarse microstructure on inner surface zone of dorsals (Figs 2C, 9B):
(0) macrohoneycomb or macrohoneycomb and lamellae; (1) lamellae only; (2) not
applicable, inner surface zone reduced to a narrow strip; (?) unknown, species not
examined by us.
Morphological extremes on inner surface zones are easily differentiated (e.g.
lamellae of Tropidurus melanopleurus vs. macrohoneycomb of Uranoscodon superciliosus),
but because macrohoneycomb shifts so gradually to lamellae in taxa with the
transitional morphology, no clear 'cut-off distinguishes very low macrohoneycomb
from lamellae. Thus, a different researcher may be inclined to code more taxa as
having all lamellae. Despite our a priom' acknowledgement that the coding of this
character is problematic, we have included it in the phylogenetic analysis because
we feel that the extreme condition is potentially informative.
Dorsals of arenicolous Microlophus, Tropidurus azureus, T. bogerti, [Link], and T.
semitaeniatus are predominantly nonimbricate and possess very narrow inner surface
zones; the character cannot be assessed in these species. Assigning a separate state
(2 = not applicable) to these species is equivalent to adding a character 'Reduction
of inner surface zone', and this state appears as a convergent synapomorphy at
three places in the phylogeny: the nodes joining (1) T. bogerti +T. semitaeniatus, (2)
[Link] + + +
T. azureus, and (3) M. peruvianus M. theresiae M. tkris.
87. Regional expression of spinules: (0) present on all scales; (1) absent from
ventrals; (?) not applicable, absent from all scales.
State 1 occurs in Liolaemus, state 0 in Stenocercus, and Leiocephalus has spines and
pits rather than the pit, pit and groove, and spinule morphotypic series. Therefore,
this character is left unpolarized.
As we discuss above, we are not certain whether the papillae of Uranoscodon
represent fused spinules or not. Papillae are on all scales in this species, and we
coded it as having state 0.
88. Presence of a morphotypic series of pits, pit and groove, and spinules on
dorsals and supraoculars (Fig. 5): (0) series present and occupying more than two
rows of macrohoneycomb chambers; (1) series truncated or absent, scales uniformly
spinulate or lacking spinules only on the anteriormost edge of the scale; (2) series
absent, scales uniformly pitted.
All three states are present in Stenocercus, but only state 0 occurs in the Liolaemus
we examined. We treat this character as unordered and states 1 and 2 as derived.
89. Papillae (Fig. 6A-D): (0) absent; (1) present.
90. Position of scale organs on paravertebrals (Figs 2A, 7C,D): (0) behind or
beside keel; (1) on surface of keel, near caudal tip.
91. Scale organs on ventral scales (Fig. 7A,B): (0) single, tiny, near caudal tip of
scale; (1) sometimes paired, lateral to keel, larger and more developed.
Tropidurus umbra and Uranoscodon superciliosus are the only tropidurids known to
have state 1. However, these are also the only ingroup taxa with keeled ventrals
(character 76 of Frost, 1992), and the two characters may be nonindependent.
Among our out<groups,Stenocercusjimbriatus also has keeled ventrals but lacks enlarged
ventral scale organs. Incongruence of these two features in S. jimbriatus suggests that
characters 76 and 91 may be independent.
212 hl B HARVEY AND R I, GUlBbRLbT JK

2 mm

Fi<pre 12. Second rows of palpebrals (white) illustrating the palpebral ‘comb’ of Microlophus thorucicus
(TCWC 28597).

92. Caudal notch on ventrals (Fig. 9): (0) absent, (1) present on some ventral
scales.
93. Position of scale organs on enlarged supraoculars (Fig. 8): (0) scale organs
arranged in a row along caudal edge of scale; (1) scale organs distributed across
scale surface in a seemingly random arrangement.
94. Number of scale organs on enlarged supraoculars: (0) four or more scale
organs on most enlarged supraoculars; (1) no enlarged supraocular with more than
five scale organs; zero to two scale organs on most supraoculars.
95. Palpebral comb (Fig. 12): (0) absent, palpebrals granular; (1) present, second
row of palpebrals from eye slit greatly attenuated into spines.
We use the term ‘palpebrals’ sensu Smith (1946) to refer to all the scales covering
the eyelids except for the ciliaries at the edge of the eyelids. Palpebrals are modified
to varying degrees among tropidurids; those on the second row from the eye form
a ‘palpebral comb’ (Fig. 12) among arenicolous Microlophus examined in this study:
M. peruvianus, M. theresiae, M. tkris, M. thoracicus. In other Microlophus and Plesiomicrolophus
koepckeorum, the same scales are slightly pointed, and this condition is barely dis-
tinguishable (these species are coded as lacking a palpebral comb, state 0) from the
decidedly granular condition of the same scales in Tropidurus.

RESULTS OF THE PHYLOGENETIC ANALYSIS

The distribution of 10 000 randomly generated trees was strongly left-skewed (g =

- 0.65 1, mean tree length 1 1 548 weighted steps) indicating that the data contained
phylogenetic signal or was correlated for reasons other than common history such
as character nonindependence. PAUP saved 270 trees during the 24 replicates of
the random search of the complete set of taxa. When zero-length branches and
 213

Ancestor
- Plesiornicrolophus
Microlophus occipitalis -Plesiomicrolophus
23 - Microlophus stolzmanni -Microlophus occipitalk
$- Microlophus bivittatus
80
-Microlophus stolzmanni
t
1 5 Microlophus theresiae
Microlophus peruvianus
Dopidurus nanuzae
Microlophus theresine
Microlophus peruvianus
Dopidurus hygomi
Dopidurus oreadicus
- Uranoscodon
-
Dopidurus nanuzae
3 Dopidurus hispidus -
Dop?urus hygomi
4

5 I
c Dopidurus insulans
Dopidurus erythrocephnlus
Dopidurus itambere
93
-
Dopidurus hispidus
Dopidurus itambere
+
1 -
19
Dopidurus nwntanus - Dopidurus nwntanus
6
t
Dopidurus mucujensis - Dopidurus oreadicus
I
Dopidurus torquatus - Dopidurus torquatus
+ --
Dopidurus bogerti
t
8 m
I
20 nopidurus semitaeniatus
50 Dopidurus melanopleurus

+ -
9
t
x, . Dopidurus melampleurus
Dopidurus callathelys
Dopidurus spinulosus
Dopidurus spinulosus
-
Dopidurus plica
-
Dopidurus umbra
10 & Dopidurus xanthochilus
-
Dopidurus strobilurus
I Dopidurus strobilurus
11 Dopidurus bogerti
Dopidurus sernitaeniatus
Dopidurus azureus
Dopidurus flaviceps

Figure 13. A, phylogeny of tropidurines based on scale surface morphology and traditional characters:
a majority consensus of 270 equally parsimonious trees each with 5 1 17 weighted steps, consistency
indices of 0.544, and rescaled consistency indices of 0.425. Polytomies were resolved in less than 50%
of the trees. Branch 17 was resolved in 87% of the trees; all remaining branches were recovered in
100% of the equally parsimonious trees. B, phylogeny of reduced set of taxa showing all branches
present in 50% or more of the bootstrap trees. Numbers indicate percentage of bootstrap trees
containing the associated branch.

duplicate trees were eliminated, this number was reduced to 135 equally parsimonious
trees with consistency indices of 0.544, rescaled consistency indices of 0.42 1, retention
indices of 0.774, and lengths of 5 117 weighted steps. A 50% majority-rule consensus
of these trees is presented in Figure 13; 87% of the trees showed resolution of
branch 17. All remaining branches were recovered in 10Oo/o of the trees. Summaries
for character transformations along numbered branches in Figure 13 are provided
in Appendix 2.
Analysis of the reduced set of taxa produced a consensus tree with a topology
identical to that of the tree containing all taxa. Our phylogeny of all taxa is
interpreted as the best hypothesis of ingroup relationships; however, statistical
confidence is limited to relatively few branches. Figure 13B shows all branches with
bootstrap proportions greater than or equal to 50. With statistical confidence, five
clades were identified. Although placement of Urunoscodon in the bootstrap tree is at
odds with its deeply nested position in Figure 13, confidence in this alternative
hypothesis is lacking: the branch leading to a clade of all included liopidurus was
recovered in 50% of the bootstrap trees and may not be accurate.
Not surprisingly, the arenicolous species of Microlophus form a well supported clade
214 M. B. HARVEY AND R. L. GUTBERLET JR

as in Frost’s study. Six derived features of this group were identified earlier (Frost,
1992; characters 48.1, 53.1, 60.1, 7 1 . l , 73.1, and 74. l), and we have added three
additional features of scale surface morphology (86.2, 94.1) and scutellation (95.1)
that support a close relationship of at least three of these species M. peruvianus, M.
theresiae, and M. tigris. Scale surfaces of Microlophus thoracicus were not examined using
electron microscopy; however, it shares presence of a palpebral comb (character
95.1) with the other members of this group we examined.
Few characters support resolution of branches within the paraphyletic Tropidurus
torquatus group of former authors; branches 4-8 are each supported by single
characters, Our placement of these species is congruent with that of Frost (1992)
except with respect to T. oreadicus, T. hispidus, and T. insulans. Movement of these
three species to the more primitive position below the T. e@hrocephalus-T. itambere
clade may have resulted from recoding (see Methods) the axillary mite pocket
character (54) for T. oreadicus. Moreover, T. oreadicus, T. hispidus, and 7: insulans lack
an inguinal mite pocket (56. l), and this character evolved in an ancestor not shared
with these species (i.e. along branch 6). Our placement of the T. torquatus group
members requires convergent evolution of number of oblique neck fold mite pockets
+
(character 48 49). However, as other researchers have pointed out (Rodrigues,
1987; Frost, 1992), characterizing these pockets as ‘one’ or ‘two’ oversimplifies their
variation.
The relationship of Tropidurus melanopleurus to ‘I:spinulosus and species formerly
placed in other genera was unresolved in Frost’s (1992) shortest tree, and T.
melanopleurus was placed as the sister species to ‘I: spinulosus in his slightly longer
alternative topology. These species’ relationships are resolved by recognizing that
T. melanopleurus has a polymorphic condition for number of rows of circumorbitals
(Fig. 14). As in Frost’s analysis, evolution of two rows of circumorbitals unites 7:
melanopleurus and T. spinulosus with several other derived species. However, fixation
of this trait becomes a new synapomorphy not considered by Frost (1992) and forces
T. melanopleurus to a position outside the terminal clade grouping 7: spinulosus, 7:
xanthochilus, and the more derived species. Moreover, to the exclusion of I: me-
lanopleurus, 7: spinulosus and the other derived species are united by macrohoneycomb
well developed over the entire surface of dorsals (character 82) and high macro-
honeycomb extending across the supralabials (character 83).
A close relationship between T. melanopleurus and 7: callathebs is supported by
lamellae covering the inner surface zones of dorsals (character 86.1) and fixation of
character 79, two postmentals contacting infralabials. Tropidurus melanopleurus and T.
callathebs share considerable sexual dimorphism in squamation and body size (Perez-
Mellado & De la Riva, 1993; Harvey & Gutberlet, 1998). Unlike any other
tropidurids, these two species exhibit an unusual instance of ‘reverse’ dichromatism
where the females are more strikingly coloured than the males. Homoplastic in our
hypothesis is loss of preocular-canthal contact in I: callathebs (fixation of character
78), a trait otherwise uniting T. spinulosus, 7: xanthochilus, and the more derived
Tropidurini from Amazonia and the Atlantic forests.
A close relationship between Tropidurus spinulosus and T. xanthochilus is supported
by mental scale reduction (character 58), a character also convergent in T. lumaria,
T. plica, and T. umbra. Reversal of characters 17 and 28 characterize 7: spinulosus;
neither of these osteological traits have been examined in T. xanthochilus. If present
in the latter species, they would provide additional support to branch 22.
Frost (1992) found that Tropidurus strobilurus, an inhabitant of the forests along the
 PHYLOGENY OF TROPIDURINI 215

Plesiomicrolophus

I Microlophus
Caudal notch

Potential synapomorphies:
Polarity equivocal
hypotheticalancestor
B

Inner surface zone Fixation: Postmental-

lamellae infralabial contact
I: melanopleurus I: melanopleurus

G
I: callathelys I: callathelys
Z spinulosus I: spinulosus
I: xanthochilus I: urnthochilus
High macrohoneycomb 2 rows of
Branch to more circumorbitds Branch to more
across labiala
High
acrossmacrohoneycomb
caudal 2/3rds derived species Firation: derived species
preocular-canthal
of dorsals contact
C D
Figure 14. The basal polytomy in Frost’s (1992) hypothesis (A) resolved using new characters of scale
surface morphology (B). Most of these new characters cannot be unambiguously polarized without
resolution of the relationships within Stenocercus. The relationship of Tropidurus melanopleurus and ?:
spinulosus resolved using new characters from scale surface morphology (C) and frequency coding (D).
Stippled boxes represent convergences or parallelisms; black boxes are unique apomorphies.

Atlantic coast of Brazil, was most closely related to the two Amazonian species
formerly placed in Uracentron and that the three species shared a flat anterior margin
of the pubis and armed caudal scales. However, these two derived traits fail to
overcome characters contradicting a close relationship between these taxa. As Frost
(1 992) points out, independent evolution of armed caudal scales among Tropidurus
would not be surprising and has occurred in other iguanian lineages such as the
closely related genus Stenocercus. Branch 13 excludes ir: strobilurus and unites all of
the Amazonian species previously (as of 1970) placed in Plica, Uracentron, and
Uranoscodon. This Amazonian clade is unambiguously supported by characters 22.1
(posterior maxillary teeth brachydont), 46.1 (gular fold complete medially), 57.1
(rostra1 scale height reduced), and 85.1 (coarse microstructure high across ventral
scales).
Our new data together with many characters discovered in earlier studies support
placement of Uranoscodon within Tropidurus and its nesting within the clade of highly
derived species. A close relationship between Uranoscodon, T. plica, and T. umbra is
supported by three characters of scale surface morphology unique to these taxa
among the tropidurids we examined: scale organs atop keels of paravertebrals in all
three (character 90. l), papillae (89.1) in Uranoscodon and [Link], and enlarged ventral
scale organs (91.1) in Uranoscodon and T. umbra. Additional support for placement of
Uranoscodon within the clade of highly derived species comes from a variety of
‘traditional’ characters (Appendix 2). However, Uranoscodon exhibits more reversals
than any other species, and most of these contradict its placement within Tropidurus.
216 M. B. HARVEY AND R. L. GUTRERLET JR

Similarities between Eopidurus nanuzae and the derived species previously placed
in Plica, Strobilurus, Uracentron, and Uranoscodon were found by Frost (1992)) and many
more were identified in this study. Tropidurus nanuzae and these more derived species
share jagged projections of the caudal wall of macrohoneycomb (character 8 1.1))a
feature unknown outside Tropidurus and probably present in Uranoscodon. Tropidurus
nanuzae and some or all of these derived species also possess well developed
macrohoneycomb over the caudal two-thirds of their dorsals (82.1) and across their
supraoculars (83.1, both characters also in some Stenocercus).

DISCUSSION

Proposed taxonomic changes and some persistent problems

Frost (1992) established the new genus Plesiomicrolophus to accommodate Tropidurus

occipitalis koepckeorum Mertens, because this taxon lacked derived conditions for the
characters he considered; most importantly, it reportedly lacks the terminal disks of
the hemipenes diagnostic of Microlophus. He went on to say “my conjecture is that
future work will show it (Plesiomicrolophus) to be the sister taxon of Microlophus, in
which case it could justifiably be considered a junior synonym of Microlophus)) (Frost,
1992: 45-46). Our new data fulfill his prediction (Fig. 14). Plesiomicrolophus koepckeorum
and all Microlophus we examined share a unique synapomorphy: the caudal notch
(character 92). They also share very low macrohoneycomb (character 80, also in
Liolaemus), loss of spinules on ventrals (87, also in Liolaemus), a broad morphotypic
transition of fine microstructure along the anterior edge of dorsals (88, in some
outgroups and some Tropidurus), and scale organs arranged along the caudal edge
of supraoculars (93, also in Liolaemus and Leiocephalus). Except for the caudal notch,
polarity of these characters is equivocal at present, although more complete resolution
of outgroup relationships, especially within Stenocercus, may well establish some or
all of these characters as synapomorphies. Some evidence such as absence of terminal
discs in F koepckeorum support its placement as the sister group to all the other trans-
Andean species; however, this placement might be equivocal because of shared
similarities (e.g. most ventrals of I? koepckeorum and M. occipitalis are notched, whereas
very few ventrals are notched in M. stolzmannz) with some Microlophus but not others.
We propose that Tropidurus occipitalis koepckeorum Mertens be transferred to Microlophus
as Microlophus koepckeorum comb. nov., and that Plesiomicrolophus Frost be considered
a junior synonym of Microlophus Dumkril and Bibron.
One problematic result of this analysis regards the placement of Uranoscodon
superciliosus. Earlier, Frost (1992) noted the bewildering mix of primitive and derived
characters in Uranoscodon. Based on our analysis of all available characters and the
results of the bootstrap analysis, we conclude with statistical confidence (Fig. 13B)
that Uranoscodon is part of a monophyletic clade of cis-Andean Tropidurini. We lack
confidence that U. superciliosus is the sister species of T. umbra; however, this deeply
nested placement within Tropidurus is the most parsimonious explanation of all
available data. The name Uranoscodon could be maintained (i.e. as sedis mutabilis
under Wiley’s, 1981: 2 11, convention 4) until its placement (ix. as the sister species
of ir: umbra or, alternatively, of all Tropidurus) in the cis-Andean clade is determined
with confidence. However, we see no reason for maintaining Uranoscodon as a
 PHYLOGENY OF TROPIDURINI 217

monotypic genus (we reject the notion that the generic name should be conserved
to emphasize the unique ecological specialization of the species) now that we are
confident it is more closely related to Tropidurus than Microlophus. We here propose
that Lucerta superciliosa Linnaeus be transferred to Tropidurus as Tropidurus superciliosus
comb. nov. and that Uranoscodon Kaup be considered a junior synonym of Tropidurus
Wied-Neuwied.
Troubling are (1) the similarities among Microlophus, Liolaemus, and Liocephalus but
not Stenocercus for several scale surface features (characters 80, 87, 93); (2) similarities
between Tropidurus nanuzae and more derived species, especially 7: strobilurus (e.g.
character 81); (3) the number of reversals among species in the clade of Amazonian
species; and (4) the frequency with which both primitive and derived states in our
own and Frost’s (1992) analyses were observed among species of Stenocercus. T o us,
these observations suggest that simultaneous resolution of the ingroup and outgroups
may be a superior method for determining character polarity than an a priori form
of argumentation (i.e. Maddison et al., 1984) based on a constrained outgroup
topology. As Nixon & Carpenter (1993) point out, the latter method is not maximum
parsimony, although we would not go so far as to agree with them in calling it
‘irrelevant’. Potentially, such a reassessment of polarity could invert the cis-Andean
clade and allow revalidation of at least some of the genera synonymized by Frost
(1992) (see, however, Frost’s alternative topology, his figure 35). We do not believe
that this is wild speculation, especially because of the lack of character support for
clades between branches 3 and 10.
A final comment regards proposition of taxonomic changes in the absence of
statistical confidence. The properties of methods for assessing confidence in phylo-
genetic analyses have only recently been studied. Synonymization of Plica and
Uracentron has been questioned (Avila-Pires, 1995), and her position is supported by
the absence of statistical confidence in a more resolved phylogeny coupled with the
other problems voiced in the preceding paragraph. We accept our shortest trees
based on all the data as the best estimations of tropidurid phylogeny. In the absence
of contradictory data or until simultaneous resolution of the ingroup and outgroup
can be accomplished, revalidation of Plica and Uracentron is unjustified. We argue
that Frost (1992) was correct to synonymize these and other genera with Tropidurus
based on the available data. T o do otherwise would accord generic status to
paraphyletic taxa and would directly contradict the recovered evolutionary history
of the group.

Scale surface microstructure and phylogenetics

Osteology and squamation will no doubt continue to be the richest and most
frequently utilized sources of morphological characters in phylogenetic studies of
lizards. New analytical methods such as frequency coding have increased the number
of codeable characters (e.g. Wiens & Reeder, 1997). Nonetheless, any one group of
characters will most likely fail to resolve relationships among all the species within
large taxonomic groups such as the Tropidurini.
Several authors (especially Peterson & Bezy, 1985; Williams, 1988; Harvey &
Gutberlet, 1995) have demonstrated the value of scale surface microstructure in
systematics. This point is driven home by the present study. Though fewer in
number (15 scale surface characters versus 2 1 characters of squamation and 40 of
218 M. B. HARVEY AND R. L. GUTBERLET JR

osteology), the scale characters used in this study proved particularly important.
Whereas some features varied intraspecifically, others were found to be diagnostic
at the generic level or to suggest shared ancestry of large groups of species.
We hope that this study will encourage more researchers to include epidermal
microstructure in phylogenetic analyses. Among squamates, we suspect that iguanians
possess the greatest diversity of scale surface microornamentation, and researchers
can reasonably expect to find many useful characters among species in this group
(e.g. Cole & Van Devender, 1976; Schleich and Kastle, 1979; Peterson, 1984; Lang,
1989). Our recent study of 41 species of cordylids and gerrhosaurids (Harvey &
Gutberlet, 1995)found fewer characters than this study, because those scincomorphan
lizards lack an important source of variation present in iguanians: the pit, pit-and-
groove, and spinule morphotypic series. This added dimension to the scale surfaces
of iguanians accounted for 20% (3 of 15) of our new epidermal characters. We did
not report all additional variation in this morphotypic series that we observed among
the outgroups. Again, we emphasize the importance of examining scale surfaces
from different body regions and, notably, the inner surface zone of individual scales.
Only 6 of our 15 characters would have been found had our survey been restricted
to dorsal scales as were most earlier studies of scale microstructure. Finally, we note
that researchers lacking access to a scanning electron microscope can quite easily
assess characters of scale organ distribution and morphology (see also Matveyeva &
Ananjeva, 1995). All of our characters based on scale organs (5 of 15) could be
assessed with a dissecting microscope.

ACKNOWLEDGEMENTS

This project grew out of research on Bolivian Tmpidurus and field studies in Bolivia
funded by Conservation International. The senior author thanks A. Forsyth and T .
Schulenberg for inviting him to participate in Rapid Assessment Program expeditions
to Bolivia. The second author’s study of scale surface morphology is supported by
a scholarship from the North Texas Herpetological Society and a grant from the
Phi Sigma Biological Society. For loan of specimens under their care, we thank
J.E. Cadle (MCZ), EJ. Censky (CM), J.R. Dixon (TCWC), H.S. Greene (MVZ),
A.G. Kluge (UMMZ), and R.P. Reynolds (USNM). At the Texas Cooperative
Wildlife Collection, J.R. Dixon and at the Smithsonian, R.P. Reynolds kindly
provided us with working space. We thank HJ. Arnott for granting us access to the
electron microscopy lab at the University of Texas at Arlington. L. Lopez provided
technical assistance regarding use of the electron microscope. D. Frost, J. Wiens,
and G. Scrocchi critically reviewed an earlier draft of this manuscript and provided
many useful suggestions.

REFERENCES

Ada-Pires TCS. 1995. Lizards of Brazilian Amazonia (Reptilia: Squamata). <oologische Erhandelingen
Leiden 299: 1-706.
Campbell JA, Frost DR. 1993. Anguid lizards of the genus Abmnia: revisionary notes, descriptions
of four new species, a phylogenetic analysis, and key. Bulleiin ofthe American Museum ofNatural History
216: 1-121.
 PHYLOGENY OF TROPIDURINI 219

Cei JM. 1982. A new species of Tmpidurus (Sauria, Iguanidaej from the arid Chacoan and western
regions of Argentina. Occasional Papers ofthe Museum ofNatural Histoy the Universip ofKansas 97: 1-10.
Cole CJ, Van Devender TR. 1976. Surface structure of fossil and recent epidermal scales from
North American lizards of the genus Sceloporu (Reptilia, Iguanidaej. Bulletin ofthe American Museum
ofNatura1 Histoty 156: 451-5 14.
DumCril AMC, Bibron G. 1837. Erpltologie Ginhale ou Histoire Naturelle Compkk des Reptiles. Vol. 4.
Paris: Roret.
Eldredge N, Gould SJ. 1972. Punctuated equilibria: an alternative to phyletic gradualism. In: Schopf
TJ, ed. Models in Paleobiology. San Francisco: W. H. Freeman, 82-1 15.
Etheridge R. 1970. A review of the South American iguanid lizard genus Plica. Bulletin of the British
Museum ofNatural Histov, ~ o o l o g y19: 235-256.
Etheridge R. 1995. Redescription of Ctenoblephay adspersa Tschudi, 1845, and the taxonomy of
Liolaeminae (Reptilia: Squamata: Tropiduridaej. American Museum Novitates 3142: 1-34.
Etheridge R, de Queiroz K. 1988. A phylogeny of Iguanidae. In: Estes R, Pregdl G, eds. Phylogenetic
relationships of the lizard families: essays commmoratiy Charles L. Camp. Stanford: Stanford University
Press, 283-367.
Farris JS. 1983. The logical basis of phylogenetic systematics. In: Platnick NI, Funk VA, eds. Advances
in cladistics, Vol. 2. New York Columbia University Press, 7-36.
Farris JS. 1989. The retention index and the rescaled consistency index. Cladistics 2: 14-27.
Farris JS, Kluge AG. 1985. Parsimony, synapomorphy, and explanatory power: a reply to Duncan.
Tmon 3 4 13Ck135.
Farris JS, Kluge AG. 1986. Synapomorphy, parsimony, and evidence. Tmon 35: 298-306.
Felsenstein J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution
39: 783-791.
Frost DR. 1992. Phylogenetic analysis and taxonomy of the Tmpidums group of lizards (Iguania:
Tropiduridaej. American Museum NoUitates 3033: 1-64.
Frost DRYEtheridge R. 1989. A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia:
Squamataj. Miscellaneous Publications the Museum of Natural Histoty Uniuersip of Kansas 81: 1-65.
Gutberlet RL Jr. 1998. The phylogenetic position of the Mexican black-tailed pitviper (Squamata:
Viperidae: Crotalinae). Herpetologica 5 4 184-206.
Harvey MB. 1993. Microstructure, ontogeny, and evolution of scale surfaces in xenosaurid lizards.
Journal ofMorphology 216: 161-177.
Harvey MB, Gutberlet RL Jr. 1995. Microstructure, evolution, and ontogeny of scale surfaces in
cordylid and gerrhosaurid lizards. Journal OfMorphology 226: 12 1-1 39.
Harvey MB, Gutberlet RL Jr. 1998. Lizards of the genus Tm~idum(Iguania: Tropiduridaej from
the Serrania de Huanchaca, Bolivia: new species, natural history, and a key to the genus. Herpetologica
54: 493-520.
Hauser DL, Presch W. 1991. The effect of ordered characters on phylogenetic reconstruction.
Cladisti~s7: 243-265.
Hendy MD, Penny D. 1982. Branch and bound algorithms to determine minimal evolutionary trees.
Mathematical Biosciences 59: 277-290.
Hillis DM. 1991. Discriminating between phylogenetic signal and random noise in DNA sequences.
In: Miyamoto MM, Cracraft J, eds. Phylogenetic ana$sis ofDNA sequences. New York Oxford University
Press, 278-294.
Hillis DM, B d J . 1993. An empirical test of bootstrapping as a method for assessing confidence
in phylogenetic analysis. systematic Biology 42: 182-192.
Hillis DM, Huelsenbeck JP. 1992. Signal, noise and reliability in molecular phylogenetic analyses.
Journal of Heredip 83: 189-1 95.
Huelsenbeck JP. 1991. Tree-length distribution skewness: an indicator of phylogenetic information.
systematic ,Zoology 40: 257-270.
Irish FJ, Williams EE, Seling E. 1988. Scanning electron microscopy of changes in epidermal
structure occurring during the shedding cycle in squamate reptiles. Journal of Morphology 197:
105-126.
Jackson MK, Reno HW. 1975. Comparative skin structure of some fossorial and subfossorial
leptotyphlopid and colubrid snakes. Herpetologica 31: 350-359.
Jackson MK, Sharawy M. 1980. Scanning electron microscopy and distribution of specialized
mechanoreceptors in the Texas rat snake, Elaphe obsoletn lindheimeri (Baird and Girardj. Journal of
Moi$hology 163: 59-67.
220 R.1. B. HARVEY AND K.L. GUTBEKLET JR

Johnson NK, Zink RM,Marten JA. 1988. Genetic evidence for relationships in the avian family
Vireonidae. Condor 90: 428-445.
Kluge AG. 1989. A concern for evidence and a phylogenetic hypothesis of relationships among
Epicrates (Boidae, Serpentes). Systematic <oology 38: 7-25.
Kluge AG. 1991. Boine snake phylogeny and research cycles. Miscellaneous Publications, Museum o f
<oology, Universip ofMichigan 178: iv, 1-58.
Kluge AG. 1993. Aspidites and the phylogeny of pythonine snakes. Records ofthe Australian Museum 19
(Supplement): 1-7 7.
Kluge AG, Farris JS. 1969. Quantitative phyletics and the evolution of anurans. Systematic <oology
18: 1--32.
Lang M. 1989. The morphology of the Oberhautchen with the description and distribution of scale
organs in basiliscine iguanians. Amphibia-Reptilia 10: 423-434.
Mabee PM. 1989. Assumptions underlying the use of ontogenetic sequences for determining character-
state order. Transactions ofthe American Fisheries Sociep 118: 151-158.
Maddison WP. 1993. Missing data versus missing characters in phylogenetic analysis. Systematic Biology
42: 576-581.
Maddison WP, Donoghue MJ, Maddison DR. 1984. Outgroup analysis and parsimony. Systematic
<OO~OQ 33: 83-103.
Manzani PRYAbe AS. 1990. A new species of Tapinurns from the caatinga of Piaui, northeastern
Brazil (Squamata: Tropiduridae). Herpetologica 46: 462-467.
Matveyeva TN, Ananjeva NB. 1995. The distribution and number of the skin sense organs of
agamid, iguanid and gekkonid lizards. Journal of <oology, London 235: 253-268.
McGuire JA. 1996. Phylogenetic systematics of crotaphytid lizards (Reptilia: Iguania: Crotaphytidae).
Bulletin of Carnegze Museum ofNatura1 Histoy 32: 1-143.
Nixon KC, Carpenter JM. 1993. On outgroups. Cladistics 9: 413-426.
Olson EC. 1975. Permo-carboniferous paleo-ecology and morphotypic series. American <oologist 15:
37 1-391.
Perez-Mellado V, De la Riva I. 1993. Sexual size dimorphism and ecology: The case of a tropical
lizard, Tropidurus melanopleurus (Sauria: Tropiduridae). Copeia 1993: 969-976.
Peters JA, Donoso-Barros R. 1970. Catalogue of the Neotropical Squamata: Part II. Lizards and Am-
phisbaenians. Washington: Smithsonian Institution.
Peterson JA. 1983. The evolution of the subdigital pad of Anolis. 2. Comparisons among the iguanid
genera related to the anolines and a view from outside the radiation. Journal of Herpetology 17:
371-397.
Peterson JA. 1984. The microstructure of the scale surfaces in iguanid lizards. Journal ofHerpetology
18: 437-467.
Peterson JAYBezy RL. 1985. The microstructure and evolution of scale surfaces in xantusiid lizards.
Herpetologica 41: 298-324.
Peterson JAYWilliams EE. 1981. A case study in retrograde evolution: The Onca lineage in anoline
lizards. 11. Subdigital fine structure. Bulletin ofthe Museum of Comparative <oology 149: 2 15-268.
Platnick NI, Griswold CE, Coddington JA. 1991. On missing entries in cladistic analysis. Cladistics
7: 337-343.
Pregill GK. 1992. Systematics of the West Indian lizard genus biocephalus (Squamata: Iguania:
Tropiduridae). 'The Universip of Kansas Museum o f Natural Histoy Miscellaneous Publication 84: 1-69.
Price R. 1982. Dorsal snake scale microdermatoglyphics: Ecological indicator or taxonomic tool?
Journal of Herpetology 16: 294-306.
Price R,Kelly P. 1989. Microdermatoglyphics: basal patterns and transition zones.Journa1 ofHerpetology
23: 244-26 1.
Rodrigues MT. 1986. Um novo Tropidurus com crista dorsal do Brasil, com comentirios sobre suas
relaGdes, distribuiGZo e origem (Sauria, Iguanidae). Papiis Avulsos de <oolopza 36: 17 1-1 79.
Rodrigues MT. 1987. Sistematica, ecologia e zoogeografia dos Tmpidurns do grupo torquatus ao sul
do Rio Amazonas (Sauria, Iguanidae). Arqivos de <oologia 31: 105-230.
Rodrigues MT. 1988. Distribution of lizards of the genus Trofiidurns in Brazil (Sauria, Iguanidae). In:
Vanzolini PE, Heyer WR, cds. Proceedings .fa rnorkshofi on neotropical distribution patterns. Rio de Janeiro:
Academia Brasileira de Citncias, 305-3 15.
Schleich HH, Kastle W. 1979. Hautstrukturen als Kletteranpassungen bei Chamaeleo und Cophotis
(Reptilia: Sauria: Chamaeleonidae, Agamidae). Salamandra 16: 95-1 00.
Schmidt WJ. 1914. Studien am Integument der Reptilien. 5. Anguiden. <oolopzscheJahrbucher Abteilung
f u r Anatomie und Ontogenie der liere 38: 1-~ 120.
 PHYLOGENY OF TROPIDURINI 221

Slowinski JB. 1993. “Unordered” versus “ordered” characters. Systematic Biology 42: 155-1 65.
S m i t h HM. 1946. Handbook oflizards. Ithaca: Comstock.
Stewart GR, Daniel RS. 1972. Scales of the lizard Gekko gekko: surface sculpture examined with the
scanning electron microscope. Copeia 1972: 252-257.
Swofford DL. 1993. PAUP Phylagenetic anabsis uing parsimony. Version 3.1.1. Champaign, Illinois:
-

Illinois Natural History Survey.

Swofford DL, Berlocher SH. 1987. Inferring evolutionary trees from gene frequency data under
the principle of maximum parsimony. Systematic ~ o o l o g y36: 293-325.
Vanzolini PE, Gomes N. 1979. O n Tmpidurus hygomi: redescription, ecological notes, distribution,
and history (Sauria, Iguanidae). Pafiiis Auulsos <oologia 32: 243-259.
WiensJ. 1993. Phylogenetic systematics of the tree lizards (genus Umsaurus).Herpetologica 49: 399-420.
Wiens J.1995. Polymorphic characters in phylogenetic systematics. Systematic Biology 44: 482-500.
Wiens J,Reeder TW. 1995. Combining data sets with different numbers of taxa for phylogenetic
analysis. Systematic Biology 44: 548-558.
Wiens J,Reeder TW. 1997. Phylogeny of the spiny lizards (Sceloporus) based on molecular and
morphological evidence. Hmfietological Monographs 11: 1-10 1.
WiensJ, Servedio MR. 1997. Accuracy ofphylogenetic analysis including and excluding polymorphic
characters. Systematic Biology 46: 332-345.
Wiley EO. 1981. Phylogenetics: the theory andpractice ofphylagenetic systematics. New York: John Wiley and
Sons.
Williams EE. 1988. A new look at the Iguania. In: Vanzolini PE, Heyer WR, eds. Proceedings o f a
workshop on Neotmpical distribution patterns. Rio de Janeiro: Academia Brasileira de Citncias, 429-488.
Williams EE. 1989. Book Review (Estes R, Pregill G, eds. Phylogenetic relationsh$s ofthe lizard families.
Essays commemorating Charles L. Camp). Copeia 1989: 1 108-1 1 17.

APPENDIX 1

Summary o f ‘traditional‘ characters

‘Traditional’ morphological characters used in Frost’s (1992) analysis are summarized below. Readers
are referred to his publication for a more complete description of these characters.
1. Skull size: (0) adult males with head length <23% of snout-vent length; (1) adult males with
head length >23% of snout-vent length.
2. Skull elevation: (0) skull not elevated at level of orbits (skull height <39% of skull length); (1)
skull elevated at level of orbits (skull height >39% of skull length).
3. Skull compression: (0) not compressed; (1) compressed.
4. Rostrum length: (0) long; (1) shortened.
5. Premaxilla: (0) nasal spine narrow; (1) nasal spine broad.
6. Nasal bones: (0) not reduced; (1) reduced.
7. Nutritive foramen of maxillary: (0) small, inconspicuous; (1) enlarged, conspicuous.
8. Maxillopalatine foramen: (0)much smaller than lacrimal foramen; ( 1) enlarged and dorsoventrally
expanded.
9. Pineal foramen: (0) present; (1) absent.
10. Squamosal shape and skull width: (0) squamosal bone relatively straight; (1) squamosal bone
curved around the posterior end of the temporal fenestra.
1 1. Superior fossa of quadrate: (0) relatively small; (1) relatively large.
12. Alveolar shelf of mandible: (0) forming a well-defined ridge; (1) alveolar ridge rounded.
13. Lingual coronoid process of dentary: (0) not overlapping anterior lingual ‘leg’ of coronoid; (1)
overlapping anterior lingual ‘leg’ of coronoid.
14. Posterior extent of dentary: (0) dentary extending <50% of length from apex of coronoid to
anterior edge of articular; (1) dentary extending >50% of the length from the apex of the coronoid to
the anterior edge of the articular.
15. Anterior surangular foramen: (0) not ‘captured’ by contact of the coronoid and dentary posterior
to the foramen; (1) ‘captured’ by contact of thc coronoid and surangular posterior to the anterior
surangular foramen.
16. Angular condition: (0) large, distinct; (1) reduced or lost.
17. Posterior mylohyoid foramen, osseous contact: (0) posterior mylohyoid foramen penetrating
222 M. B. HARVEY AND R. I,. GUTBERLET JR

angular or between dentary and angular; (1) between angular and splenial; (2) between dentary and
splenial.
18. Posterior mylohyoid foramen, position: (0) at the level of anterior end of mandibular fossa; (1)
placed more posteriorly, about 33”/0of the length of the mandibular fossa back from the anterior end.
19. Premaxillary teeth, number: (0) 6-7; (1) 4-5.
20. Anterior maxillary teeth, enlargement: (0) not or only feebly enlarged in older adults; (I) enlarged
in older adults, forming caniniform teeth.
2 I . Posterior maxillary and dentary teeth, crown flaring: (0) shaft parallel-sided with crowns not or
weakly flared; (I) crowns flared.
22. Posterior maxillary teeth, elevation: (0) posterior maxillary teeth apparently hypsodont; (I)
posterior maxillary teeth brachydont.
23. Posterior maxillary teeth, orientation: (0) posterior maxillary teeth set obliquely on the maxilla;
(I) posterior maxillary teeth set more vertically on the maxilla.
24. Pterygoid teeth: (0) present; (1) absent.
25. Clavicle: (0) strongly flanged, frequently fenestrate; (1) weakly flared or cylindrical, never
fenestrate.
26. Sternum, fenestration: (0) single fenestration present; (I) fenestra absent.
27. Posterior process of the interclavicle anterior to the sternum: (0) ‘free’ part of the posterior
process of the interclavicle >25% of the total length of the sternum; (I) ‘free’ part <25%.
28. Interclavicle median process: (0) posterior process of the interclavicle extending as a broad
process posteriorly well beyond the posterolateral corners of the sternum; (I) not extending posteriorly
beyond the posterolateral corners of the sternum.
29. Scapular deflection and fenestration of scapulocoracoid (0) scapular fenestra present, large, and
scapula not bent; (I) scapular fenestra present, reduced, scapula weakly bent; (2) scapular fenestra
absent, with no room for fenestra in scapula because extremely bent.
30. Suprascapular fenestrations: (0) absent or very tiny; ( I ) large.
31. Rib formula: (0) five ribs in contact with the sternum and xiphisternum: 3 sternal ribs + 2
xiphisternal ribs; (I) six ribs in contact with the sternum and xiphisternum.
32. Recurved xiphisternal-pectoral ribs: (0) abscnt or present in short spurs; (1) present, long.
33. Humerus, head; (0) articular surface scroll-like; (1) somewhat elevated, ovate; (2) ball-shaped
and very elevated.
34. Medial centrale: (0) present; (1) absent.
35. Fourth metacarpal and first phalanx of fourth finger: (0) fourth metacarpal distinctly shorter
than third metacarpal-first phalanx of fourth finger distinctly shorter than first phalanx of third
finger; (I) fourth metacarpal equal in length to third metacarpal-first phalanx of fourth finger distinctly
shorter than first phalanx of third finger; (2) fourth metacarpal equal to length of third metacarpal-first
phalanx of fourth finger subequal to first phalanx of third finger.
36. Claw of first toe: (0) weakly flexed; (I) strongly flexed, recurved.
37. Fringe on fourth toe: (0) absent; (I) present.
38. Pubic symphysis, anterior margin: (0) acute; (I) flattened.
39. Anterior caudal vertebrae, neural spines: (0) moderate to high; (1) very depressed.
40. Caudal vertebrae, autotomy fracture planes: (0) present; (I) absent.
41. Nostrils: (0) exposed posterolaterally and key-hole shaped or some other modification thereoc
(1) exposed laterally and widely open; (2) directed anteriorly or anterolaterally and unconstricted.
42. Ventral thigh and preanal pigmentation region: (0) thighs without a well-defined ventral
pigmentation spot; ( I ) thigh and preanal region with a well-defined yellow or greenish spot; (2) thigh
and preanal region with a well-define brown or black spot.
43. Hemipenes, condition: (0) no terminal disks on hemipenial lobes; ( I ) terminal disks present on
hemipenial lobes.
44. Hemipenes, length of lobes: (0) short; (1) long.
45. Hemipenes, ornamentation: (0) calyces start below crotch between lobes; (I) calyces start at a
level well above the crotch between the hemipenial lobes.
46. Gular fold: (0) incomplete medially; ( I ) complete medially.
47. Antegular fold: (0) absent or weak and variable; (I) present, strong, well anterior to gular fold;
(2) present, strong, closely approximating or overlapping gular fold.
48. Antegular-oblique neck fold mite pockets, condition: (0) weak single mite pocket; (1) no mite
pocket; complex neck folding; (2) a well-developed mite pocket in the upper position; (3) ventrolateral
mite pockets; (4) no obvious mite pocket.
 PHYLOGENY OF TROPIDURINI 223

49. Oblique neck fold mite pocket, condition: (0) two mite pockets; ( I ) a single well-developed mite
pocket in the upper position indicated; (2) single, very enlarged mite pocket.
50. Mite pocket in antehumeral fold: (0) absent; (1) present laterally; (2) present ventrolaterally in
antehumeral-antegular fold.
51. Tufts of spines on side of neck: (0) absent; (1) present.
52. Rictal fold: (0) absent; (I) present.
53. Supra-auricular fold: (0) absent or poorly developed; ( I ) present, well developed.
54. Axillary pocket, presence: (0) absent; (1) present.
55. Axillary pocket, condition: (0) present, single; (1) multiple, usually 3, sometimes only 2.
56. Inguinal granular pocket: (0) absent or represented solely by a nongranular fold; ( I ) present.
57. Rostral scale, height: (0) rostral scale 1.5 to 2 x height of adjacent supralabials; (I) rostral scale
height reduced, less than 1.5 x height of adjacent supralabials.
58. Mental scale: (0) enlarged, extending posteriorly well beyond level of adjacent infralabials; (I)
reduced, not extending posteriorly well beyond level of adjacent infralabials.
59. Postmental series: (0) well defined; ( I ) poorly defined or absent.
60. Infralabial scales, number: (0) 6; ( I ) 8-9.
6 1. Infralabial expansion: (0) infralabials not ventrally expanded; ( I ) infralabials greatly expanded
ventrally.
62. Lateral gular scales: (0) imbricate posteriorly; ( I ) imbricate laterally.
63. Scales of frontonasal region: (0) imbricate posteriorly or no imbrication; ( I ) weakly imbricate
anteriorly; (2) all head shields strongly imbricate anteriorly.
64. Superciliary scales: (0) not or only weakly produced vertically to form a longitudinal crest; ( I )
produced vertically to form a longitudinal crest.
65. Circumorbital series: (0) in one row between the supraoculars and the median head shield; (1)
in two rows between the supraoculars and the median head shield.
66. Circumorbitals: (0) small; ( I ) enlarged at the expense of the supraoculars.
67. Interparietal: (0) not enlarged, smaller than interorbital distance; (1) enlarged, larger than
interorbital distance.
68. Interparietal length: (0) subequal to siLgnificantlyless than width; (1) significantly longer than
wide.
69. Rows of scales between subocular and supralabials: (0) 0-1; (I) 2 or more.
70. Subocular: (0) entire-0-1 preoculars; (1) divided-at least 2 preoculars in contact with the
orbit; (2) subocular-preocular series so fragmented as to be obscure.
7 I . Preauricular fringe: (0) present, ear canal deep, a continuous fringe of scales partially to nearly
completely covering ear opening; (I) reduced, ear canal deep; (2) auricular scales reduced, car canal
deep, a lower lobule with several short spines present; (3) auricular fringe absent, the ear canal shallow.
72. Middorsal scale row: (0) present; ( I ) absent.
73. Paravertebral scales: (0) keeled; (1) not keeled.
74. Lateral body scales: (0) imbricate, keeled; ( I ) granular and juxtaposed.
75. Caudal scales: (0) tail unarmed, longer than head + body; ( I ) tail armed with heavy mucrons,
+
roughly terete and subequal to length ofhead body; (2) tail armed with heavy mucrons, dorsoventrally
flattened and shorter than head + body.
76. Ventral scales: (0) smooth; (1) keeled.
77. Upper thigh scales: (0) not heavily mucronate; ( I ) heavily mucronate.
224 M. B. HARVEY AND K.L. GUTRERLET JR

APPENDIX 2

Data matrices of ingroup and outgroup taxa

Characters and states for species included in the phylogenetic analysis are provided. Characters
1L77 are those of Frost (1992) including modifications detailed in the phylogenetic section above. ‘?’
+
=missing, unknown, or not applicable. Characters 47,48 49, 54 55, and 75 were partially ordered. +
In the data matrix, presencelabsence coding of these characters is followed by coding of the condition
(CON) of their presence.

Species 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95

Leiocephalus personatus A A ? ? ? 0 1 0 ? 0 ? ? 0 0 0 0 0 0
Leio. schreibersi ? ? ? ? ? 0 1 0 ? 0 ? ? 0 0 0 0 0 0
Leio. semilineatus A A ? ? ? O I O ? ? ? 0 0 0 0 0 0 0
Liolaemus archPforur ~ ~ 0 0 0 0 0 1 0 0 00 0 0 0 0 1 0
Liol. chiliensis A A 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0
Liol. elongutus A A 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0
Stenocercus caducus K A I 0 1 1 ? 1 0 0 I 0 0 0 O ? ? 0
Steno. mumoratus c c i o ~ i o o o o i ~ o o o o i ~
[Link] A A 1 0 1 1 1 1 0 0 0 0 0 0 0 l 0 0
[Link] A A 1 0 0 0 0 0 0 ? 2 0 0 0 0 1 ? 0
Steno. iridescens G A 1 0 0 0 1 1 0 2 0 0 0 0 0 1 0 0

Species 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3

Ancestor 0 0 0 0 0 0 0 0 0 0 0 0 0
Uranosrodon 0 1 0 1 1 0 0 0 0 0 1 1 0
Micmlophus theresiae 0 0 0 0 0 0 0 0 0 0 1 I l
h% peruvianus 0 0 0 0 0 0 0 0 0 0 1 1 1
Plesiomicrolophus 0 0 0 0 0 0 0 0 0 0 1 1 1
M. occipitalis 0 0 0 0 0 0 0 0 0 0 1 1 1
M. stolzmanni 0 0 0 0 0 0 0 0 0 0 1 1 1
M. bivittatus 0 0 0 0 0 0 0 0 0 0 1 1 1
Tmpidnnis nanuzae 1 0 0 0 0 0 1 0 0 0 1 2 1
7: etheridgei 1 0 0 0 0 0 1 0 0 0 1 2 1
7: hygomi 1 0 0 0 0 0 1 0 0 0 1 2 1
7: vthrocephalus 1 0 0 0 0 0 1 0 0 0 1 2 1
7: hispidus 1 0 0 0 0 0 1 0 0 0 1 2 1
7: insulans 1 0 0 0 0 0 1 0 0 0 1 2 2
7: itambere 1 0 0 0 0 0 1 0 0 0 1 2 1
7 montanm 1 0 0 0 0 0 1 0 0 0 1 2 1
7: mucujensis 1 0 0 0 0 0 1 ~ 0 0 1 2 ~
7: oreadicus 1 0 0 0 0 0 1 0 0 0 1 2 1
7: torquatus 1 0 0 0 0 0 1 0 0 0 1 2 1
7: bogerti 1 0 0 0 0 1 1 0 0 0 1 2 1
7: melanopleuniJ 1 0 0 0 0 0 1 0 0 0 1 2 1
7: spinnlosus 1 0 0 0 0 0 1 0 0 0 1 2 I
7: semitaeniatus 1 0 1 0 0 2 1 0 0 0 1 2 1
7: plica 1 0 0 0 1 0 1 1 0 1 1 2 1
7: umbra 1 1 0 1 1 0 1 1 0 1 1 2 1
7: stmbilurus 1 0 0 0 1 0 1 1 0 0 1 2 1
7: azureus 1 0 0 0 I 0 I I I 0 1 2 1
7:flauiceps 1 0 0 0 1 0 1 1 0 0 1 2 1
7: callathe!yJ ] 0 0 0 2 7 7 ? ? 2 2 ? 2
7: chromatops 1 0 0 0 ? ? 2 3 > ? 2 ? ’
7: xanthochilus 1 0 0 0 ? ” ? ? ? ? 2 >
 PHYLOGENY OF TROPIDURINI 225

APPENDIX 2-continued

Species 14 15 16 17 18 19 20 21 22 23 24 25 26

Ancestor o ? o 0 0 o ? o o 0 o o
Uranoscodon 0 0 0 0 0 1 0 0 1 1 0 0 0
Microlophus thereJiae 0 0 0 0 0 0 0 0 0 0 0 0 0
[Link] 0 0 0 0 0 0 0 0 0 0 0 0 0
Plesiomicrolophus 0 0 0 0 0 0 0 0 0 0 0 0 0
hf. occipitalis 0 0 0 0 0 0 0 0 0 0 0 0 0
M. stolzmanni 0 0 0 0 0 0 0 0 0 0 0 0 0
hl. bivittatus 0 0 0 1 0 0 0 0 0 0 0 0 0
Tmpidums nauuzae 1 1 1 2 0 0 I 0 1 0 0 0 1
7: etheridgei 1 1 I 2 0 0 1 0 0 0 0 1 0
7: hygomi 1 1 I 2 0 0 1 0 0 0 0 1 0
7: eythmcephalus 1 1 I 2 0 0 1 0 0 0 0 I 0
7: hispidus I 1 1 2 0 0 1 I 0 0 0 1 0
7: insulans ? ? 1 ? ? 0 1 0 0 0 0 1 0
7: itambere 1 1 1 2 0 0 1 0 0 0 0 1 0
7: montanu 1 I I 2 0 0 I 0 0 0 0 1 0
7: mucujensis ? ? ? ? ? 0 1 0 0 0 0 I 0
7: oreadicus I 1 I 2 0 0 1 0 0 0 0 I 0
7: torquatus 1 1 1 2 0 0 1 0 0 0 0 I 0
7: bog& 1 0 I 2 0 0 1 0 0 0 1 1 0
7: melauopleurus 1 1 1 2 0 0 1 0 0 0 0 I 0
7: spinulosus 1 1 1 0 0 0 1 0 0 0 0 1 0
7: semitamiatus 1 1 1 2 I 1 1 1 0 0 1 1 0
7:plica 1 1 I 2 0 1 I 1 1 I 0 1 0
7: umbra I I 1 2 0 0 1 1 1 1 1 0 0
7: stmbilurus 1 I I 2 0 1 1 1 0 1 I 1 0
7: aznreu I I 1 0 1 1 1 1 0 1 1 0 0
7:Jlaviceps 1 1 1 0 1 1 I 1 1 1 0 0 0
7: callathehs ? ? ? ? 7 0 1 0 0 0 ? ? ?
7: chmmatops ? ? ? 2 ? 0 1 0 0 0 ? ? ?
7: xanthochilus ? ? ? ? ? 0 1 0 0 0 ? ? ?
~~

continued
226 M. B. HARVEY AND R. L. GUTBERLETJR

APPENDIX P--continued

~ ~~

Species 27 28 29 30 31 32 33 34 35 36 37 38 39

Ancestor 0 0 0 0 0 0 0 0 0 0 0 0 0
Uranosrodnn 1 1 0 1 0 0 0 0 0 0 1 0 0
Micmlophus theresiae 1 1 0 0 0 0 1 0 0 0 0 0 0
M. pemuianus 1 1 0 0 0 0 1 0 0 0 0 0 0
Plesiomicmlophus 1 1 0 0 0 0 1 0 0 0 0 0 0
M. occipitalis 1 1 0 0 0 0 1 0 0 0 0 0 0
M. stolzmanni 1 1 0 0 0 0 1 0 0 0 0 0 0
M. bivittatus 1 1 0 0 0 0 1 0 0 0 0 0 0
Tmpidums nanupae 1 1 0 1 0 0 2 1 1 0 0 0 0
7: ethmidgei 1 1 0 1 0 0 2 1 1 0 0 0 0
7: fygnmi 1 1 0 1 0 0 2 1 1 0 0 0 0
7: erythmcephalus 1 1 0 1 0 0 2 1 1 0 0 0 0
7: hispidus 1 1 0 1 1 0 2 1 1 0 0 0 0
7: insulans 1 1 0 1 1 0 2 1 1 0 0 0 0
7: itambere 1 1 0 1 0 0 2 1 1 0 0 0 0
T montanus 1 1 0 1 1 0 2 1 1 0 0 0 0
7: mucujensis 1 1 0 1 1 0 2 1 1 0 0 0 0
7: oreadicus 1 1 0 1 1 0 2 1 1 0 0 0 0
7: torquatu 1 1 0 1 1 0 2 1 1 0 0 0 0
7: dogerti 1 0 1 1 1 0 2 1 2 0 0 0 1
7: melanopleums 1 1 1 1 1 0 2 1 2 0 0 0 0
7: spinuloslls 1 0 1 1 1 0 2 1 2 0 0 0 0
7: semitaeniatus 1 1 2 1 0 1 2 1 2 0 0 0 1
7:plica 1 1 0 1 1 0 2 1 2 0 0 0 0
7: umbra 1 1 0 1 1 0 2 1 2 0 0 0 0
7: stmbilunu 1 1 0 1 1 0 2 1 2 0 0 1 0
7: azurells 1 0 0 0 1 0 2 1 2 1 0 1 0
7:Jauiceps 1 1 0 0 1 0 2 1 2 1 0 1 0
7: callatfzelys ? ? ? ? ? ? ? ? T O O ? ?
7: chmmalops ? ? ? ? ? ? ? ? ? 0 0 ? ?
7: xanthochilus ? ? ? ? ? ? ? ? ? 0 0 ? ?

continued
 PHYLOGENY OF TROPIDURINI 227

APPENDIX 2-continued

Species 40 41 42 43 44 45 46 47 C 48 C 50 51 52
o + o
N 49 N

Ancestor 0 0 0 0 0 ? 0 0 ? ? ? 0 0 0
Uranoscodon 0 0 0 0 1 1 1 1 1 1 6 0 1 0
Micmlophus theresiae 0 0 0 1 1 0 0 0 ? 1 1 0 0 0
M. peruuianus 0 0 0 1 1 0 0 0 ? 1 1 0 0 0
Plesiomim1ophu.r 0 0 0 0 1 0 0 0 ? 0 ? 0 0 0
M . occipitalis 0 0 0 1 1 0 0 0 ? 0 ? 0 0 0
M . stolzmanni 0 0 0 1 1 0 0 1 1 0 ? 0 0 0
M. bivittatus 0 0 0 1 1 0 0 0 ? 0 ? 0 0 0
Tmpidunu nanuzae 0 0 1 0 1 0 0 0 ? 1 3 1 0 0
T. etherulgei 0 0 2 0 1 1 0 0 ? 1 2 0 0 0
7: tygomi 0 0 2 0 1 1 0 0 ? 1 2 0 0 0
T. eythmcephalus 0 0 2 ? ? ? 0 0 ? 1 3 0 0 0
Z hkpidtcc 0 0 2 0 1 1 0 0 ? 1 3 0 0 0
T. insulans 0 0 2 0 1 ? 0 0 ? 1 3 0 0 0
T. itambm 0 0 2 0 1 1 0 0 ? 1 3 0 0 0
7: montanus 0 0 2 0 1 1 0 0 ? 1 4 0 0 0
T. mucujemi 0 0 2 ? ? ? 0 0 ? 1 3 0 1 0
T. oreadictcc 0 0 2 0 1 1 0 0 ? 1 4 0 0 0
T. torquatus 0 0 2 0 1 1 0 0 ? 1 2 0 1 0
T. bog&' 0 1 2 0 1 1 0 0 ? 1 1 0 1 1
I: melanophm 0 1 1 0 1 1 0 1 1 1 5 0 1 1
Z spinulosur 0 0 1 0 1 1 0 1 2 1 5 0 1 1
Z semitaeniatus 0 1 2 0 1 1 0 0 ? 1 1 0 1 1
Z plua 0 2 1 0 1 1 1 1 2 1 5 2 1 1
I: umbra 0 2 0 0 1 1 0 1 2 1 5 0 0 0
7: stmbilum 0 2 1 0 1 1 0 0 ? 0 ? 0 1 1
T. a z u w 1 2 0 0 1 1 1 1 1 1 1 0 0 1
7:jlapaviceps 1 2 0 0 1 1 1 1 1 1 1 0 1 1
'I:callathe&s ? 0 1 0 1 1 0 1 1 1 5 0 1 1
7:chmmatops ? 0 2 0 1 1 0 0 ? 1 2 0 0 0
Z xanthochilus ? 0 1 0 1 1 0 1 2 1 5 0 1 1

continued
228 hl. B. HARVEY AND R. I,. GUTBERLET JR

APPENDIX P--contznued

Species 53 54 C 56 57 58 59 60 61 62 63 64 65
+ 0
55 N

Ancestor 0 0 ? 0 0 0 0 0 0 0 0 0 a
CTranoscodon 0 0 ? 0 1 0 1 0 0 1 0 1 a
Micro1ophu.r themiae 1 0 ? 0 0 0 0 1 0 1 0 0 a
.\I. peruvianus 1 0 ? 0 0 0 0 1 0 1 0 0 a
PleJiomicrolophns 0 0 ? 0 0 0 0 0 0 1 0 0 a
,M. occipitali,\ 0 0 ? 0 0 0 0 0 0 1 0 0 a
'\'I. JtolZmanni 0 0 ? 0 0 0 0 0 0 I 0 0 a
'24. bivittatu 0 0 ? 0 0 0 0 0 0 1 0 0 a
TropiduruJ nanuzae o o ? o 0 0 o o 0 I 0 o a
7: etheridgei 0 0 ? 0 0 0 0 0 0 1 0 0 a
7: hygomi 0 0 ? 0 0 0 0 0 0 I 0 0 a
7: e9throcephaluJ 0 1 I 1 0 0 0 0 0 1 0 0 a
7: hirpidus 0 1 1 0 0 0 0 0 0 l 0 0 a
7: inalans 0 1 1 0 0 0 0 0 0 1 0 0 a
7: itambere 0 ? ? 1 0 0 0 0 0 1 0 0 a
7: montanus 0 1 2 1 0 0 0 0 0 1 0 0 a
7: mucujensis 0 1 2 1 0 0 0 0 0 1 0 0 a
7: oreadicu 0 0 ? 0 0 0 0 0 0 1 0 0 a
7: torquatus 0 1 2 1 0 0 0 0 0 1 0 0 a
7: bogerti 0 1 2 1 1 0 0 0 0 0 ? 0 a
7: melanupleumc 0 0 ? 0 0 0 1 0 0 1 1 0 c
7: .$inuloJuJ 0 0 7 0 0 1 1 0 0 1 I 0 y
7: smzitaeriiatus 0 1 2 i o n 0 0 0 I l o a
7: plica 0 0 ? 0 1 1 1 0 0 1 2 1 y
7: umbra 0 0 ? 0 1 1 1 0 0 1 1 1 y
7: stmbiluru3 0 0 ? 0 0 0 1 0 0 1 1 0 y
7: azureus 0 0 ? 0 1 0 0 0 1 0 1 O y
7:fiavzceps 0 0 ? 0 1 0 0 0 1 0 1 0 y
7: callathebs 0 0 ? 0 0 0 l O O l l 0 h
7: chromatops 0 0 ? 0 0 0 0 0 0 1 0 0 a
7: xanthochzlus 0 0 ? 0 0 I 1 0 0 1 I 0 y

conhued
 PHYLOGENY OF TROPIDURINI 229

APPENDIX 2-continued

_____

Species 66 67 68 69 70 71 72 73 74 75 C 76 77 78
0
N

Ancestor 0 0 0 0 0 0 0 0 0 0 ? 0 0 a
Uranoscodon 0 1 0 0 2 3 0 0 0 0 ? 1 0 ?
Micmlophus theresiae 0 l O O O l l l l O ? O O a
M . pmvianus 0 1 0 0 0 1 0 1 1 O ? 0 0 a
Plesiomicmlophus 0 1 0 0 0 0 0 0 0 0 ? 0 0 a
M. occipifalis 0 1 0 0 0 0 0 0 0 0 ? 0 0 a
M. stolzmanni 0 1 0 0 0 0 0 0 0 0 ? 0 0 a
M. biuittatus 0 1 0 0 0 0 0 0 0 O ? 0 O ?
Tmpidurus nanu7a.e 0 1 0 0 0 0 0 0 0 0 ? 0 0 a
7: ethendgi 0 1 0 0 0 0 1 0 0 0 ? 0 0 a
7: lygomi 0 l O O O O l O O O ? O O a
7: qthmcephalus 0 1 0 0 0 0 1 0 0 0 ? 0 0 ?
7: hispidus O l O O O O l O O O ? O O a
7: insulans 0 1 0 0 0 1 1 0 0 0 ? 0 0 ?
7: itambere 0 1 1 0 0 0 1 0 0 0 ? 0 0 m
7: montanus 0 1 0 0 0 0 1 0 0 0 ? 0 0 a
T. mucujensis 0 1 0 0 0 0 1 0 0 0 ? 0 0 ?
7: oreadicus 0 1 0 0 0 0 1 0 0 0 ? 0 0 e
7: torquatus 0 1 0 0 0 0 1 0 0 0 ? 0 0 c
7: bogerti 0 1 0 0 0 2 1 0 0 0 ? 0 0 a
T. melanopleurus 0 1 0 0 0 2 0 0 0 0 ? 0 0 a
T. spinulosus 0 1 0 0 0 2 0 0 0 0 ? 0 0 y
7: semitaeniatus 0 1 0 0 0 2 1 1 0 0 ? 0 0 a
[Link] 0 1 0 1 1 2 0 0 0 0 ? 0 0 y
7: umbra 0 1 0 1 2 3 0 0 0 0 ? 1 0 ?
7: stmbilums 0 1 0 0 1 2 0 0 0 1 1 0 1 y
7: azureus 1 1 1 0 1 3 1 1 0 1 2 0 o y
7:Jlauiceps 1 1 1 0 1 3 1 0 0 1 2 o o y
7: callathehs 0 1 0 0 0 2 0 0 0 0 ? 0 0 y
7: chmmatops 0 1 0 0 0 0 1 0 0 0 ? 0 0 a
7: xanthochilus 0 1 0 0 0 2 0 0 0 O ? 0 0 y

contznued
230 M. B. HARVEY AND R. L. GUTBERLET JR

APPENDIX 2--continued

Species 79 80 81 82 83 84 85 86 87 88 89 90 91

Ancestor a ? 0 0 0 ? 0 0 ? 0 0 0 0
Uranoscodon a 1 1 1 1 1 1 0 0 2 1 1 1
hlicmlophus theresiae a 0 0 0 0 0 0 2 1 0 0 ? 0
M. peruvianus f 0 0 0 0 0 0 2 1 0 0 ? 0
Plesiomicrolophus a 0 0 0 0 0 0 0 1 0 0 0 0
hf. occipitah a 0 0 0 0 0 0 0 1 0 0 0 0
A4 stolzmanni a 0 0 0 0 0 0 0 1 0 0 0 0
hl. bizittatus ? ? ? ? ? ? ? ? ? ? ? ? ?
Tmpidurus nanuzae a 1 1 1 1 1 0 0 ? 2 0 0 0
7: etheridgei a 1 0 0 0 0 0 0 0 1 0 0 0
7: hygomi a 1 0 0 0 0 0 0 0 1 0 0 0
7:elythmcephalus T ? ? ? ? ? ? ? ? ? ? ? ?
7: hkpidus n l O O O O O 0 O l O O O
7: insulans T ? ? ? ? ? ? ? ? ? ? ? ?
7:itambere a 1 0 0 0 1 0 0 0 1 0 0 0
7:montanus a 1 0 0 0 0 0 0 0 1 0 0 0
7: mucujensis ? ? ? ? ? ? ? ? ? ? ? ? ?
7: oreadicus a l O l O l O O O l O O O
7: torquatus c 1 0 0 0 0 0 0 0 1 0 0 0
7: bogerti m 1 0 0 0 1 0 2 ? 2 0 0 0
7: melanopleurus y 1 0 0 0 0 0 1 0 1 0 0 0
7:spinulosus a 1 0 1 1 1 0 0 0 1 0 0 0
7: smitaeniatus q 1 0 0 0 0 0 2 0 1 0 ? 0
7:plica ? l l l l l l 0 0 0 l l O
7: umbra ? 1 1 1 1 1 1 0 0 0 0 1 1
7:strobilulur a 1 1 1 1 1 0 1 ? 2 0 0 0
7:azureus ? 1 1 1 1 1 1 2 0 0 0 0 0
7:jlauiceps ? 1 1 1 1 1 1 2 0 0 0 0 0
ir: callathebs y 1 0 0 0 0 0 l 0 0 0 0 0
7: chromatops b l O O O O 0 O O l O 0 O
7:xanthochilus a l O l l l O O 0 l O 0 O

continued
 PHYLOGENY OF TROPIDURINI 231

APPENDIX 2-continued

Species 92 93 94 95

Ancestor 0 0 0 0
Uranoscodon 0 1 0 0
Micmlophus h e s i a e 1 0 1 1
M. peruvianus 1 0 1 1
Plesiomicmlophus 1 0 0 0
M. occipitalir 1 0 0 0
M. stolrmanni 1 0 0 0
M. biuittatus ? ? ? ?
Tmpidunu nanurae 0 1 0 0
7:ethendgei 0 1 0 0
7:hygomi 0 1 0 0
7:qthmcephalus ? ? ? ?
7:hispidu 0 1 0 0
7:insulans ? ? ? ?
7:itambm 0 1 0 0
7:montanus 0 1 0 0
7:mucujmxi ? ? ? ?
7:oreaditus 0 1 0 0
7:tnrquatus 0 1 0 0
7:boguti 0 1 0 0
7:mehopleunu 0 1 0 0
7:spinulosus 0 1 0 0
7:semitoeniatus 0 1 0 0
7:plica 0 1 0 0
?: umbra 0 1 0 0
7:stmbilunu 0 1 0 0
7:azurm 0 1 0 0
7:Javiceps 0 1 0 0
Tmpidunu sp. I 0 1 0 0
Tmpidunu sp. 2 0 1 0 0
Tmpidunu sp. 3 0 1 0 0

APPENDIX 3

Apomorphy list
Numbered branch labels in the 50% majority consensus tree (Fig. 13) are followed by characters
supporting them unambiguously (bold) or under accelerated or delayed transformation optimization.
The derived character state for each character is placed after a decimal point. Weights (i.e. number
of steps) are supplied for polymorphic characters. In the analysis, all remaining characters were assigned
a base weight of 24 so that fixation of a polymorphic character was equivalent to a state change in a
nonpolymorphic character.
Unambiguous character to branch assignments (bold) and ambiguous character assignments under
accelerated transformation optimization:
Branch 1. - 11.1, 12.1, 13.1, 27.1, 28.1, 33.1, 44.1, 62.1, 67.1. Branch 2. - 1.1, 7.1, 12.2, 14.1,
15.1, 16.1, 17.2, 20.1,30.1,33.2,34.1,35.1,42.1,48.1,80.1,88.1,93.1,48+49condition.3. Branch
+
3. - 25.1, 42.2, 45.1, 72.1. Branch 4. - 31.1. Branch 5. - 54 55.1. Branch 6. - 56.1. Branch 7. 54 55
~ +
condition.2. Branch 8. - 51.1. Branch 9. - 29.1, 35.2, 52.1, 63.1, 71.2, 86.1, 48+49 condition.1.
Branch 10. - 42.1, 47+48.1, 54+55.0, 56.0, 59.1, 65.2, 72.0, 78.Y (24 steps), 48+49 condition.5,
86.1. Branch 11. - 65.Y (22 steps), 79.0, 82.1, 83.1, 84.1, 86.1, 47 condition.2. Branch 12. - 5.1, 8.1,
19.1, 21.1, 23.1, 29.0, 41.2, 70.1, 81.1, 88.0. Branch 13. 22.1, 46.1, 57.1, 64.1, 85.1, 90.1, 75
~

condition.2. Branch 14. - 17.0, 25.0, 42.0, 71.3, 47 condition.1. Branch 15. -2.1, 4.1, 52.0, 70.2, 76.1,
91.1. Branch 16. - 18.1, 30.0, 36.1, 38.1, 40.1, 59.0, 61.1, 62.0, 64.0, 66.1, 68.1, 72.1, 75.1, 86.2,
90.0, 48+49 condition.1. Branch 17. - 43.1. Branch 18. - 48.1, 53.1, 60.1, 71.1, 73.1, 74.1, 86.2,
232 M. B. HARVEY AND R. L. GUTBERLET JR

94.1, 95.1. Branch 19. 31.0, 78.M (12 steps), 84.1. Branch 20. - 6.1, 24.1, 39.1, 41.1, 79.M (12
-

steps), 86.2. Branch 21. - 79.Y (24 steps). Branch 22. 17.0, 28.0, 58.1. Branch 23. 87.1, 92.1.
- -

Uranoscodon. - 1.0, 7.0, 8.0, 12.1, 13.0, 14.0, 15.0, 16.0, 20.0, 21.0, 31.0, 33.0, 34.0, 35.0, 37.1,
41.0, 63.0, 65.A (24 steps), 88.2, 89.1, 4 8 f 4 9 condition.6. Tropidurus umbra. - 10.1, 17.2, 19.0,
24.1, 46.0, 51.0, 58.1, 69.1, 47 condition.2. Tmpidurus azureus. - 9.1, 22.0, 24.1, 28.0, 51.0, 73.1.
Tmpidurusplica. - 10.1, 50.2, 58.1, 63.2, 69.1, 89.1. Tmpidurus strobilurus. - 24.1, 38.1, 47+48.0, 75.1,
77.1, 86.1, 88.2. Tmpidurus melanopleurus. 41.1, 78.A (24 steps). Tmpidurus callathehs. - 65.B (1 step),
-

88.0. Tmpidurus bogerti. - 15.0, 28.0, 57.1, 62.0, 84.1, 88.2. Zopidurus semitaeniatus. 3.1, 6.2, 18.1,
-

19.1, 21.1, 29.2, 31.0, 32.1, 73.1, 79.Q (4 steps). Tmpidurus torquatus. 4 8 f 4 9 condition.2, 78.C
-

(2 steps), 79.C (2 steps). Tmpidurus montanus. - 48+49 condition.4. Sopidurns itambere. - 68.1.
Tmpidurus hkfiidus. - 21.1, 79.N (13 steps). Tmpidurus insulans. 71.1. Tipidurus oreadicus. - 78.E (4
-

steps), 82.1, 84.1, 48+49 condition.4. Tropidurus hygomi. - 4 8 t 49.2. Tmpidurus nanuzae. 22.1, -

26.1, 50.1, 81.1, 82.1, 83.1, 84.1, 88.2. Microlophus theresiae. - 72.1. Microlophus peruvianus. - 79.F (5
steps). Microlophus stolzmanni. - 47 t 48.1. Microlophus bivittatus. 17.1.
-

Ambiguous character to branch assignments under delayed transformation optimization:

Branch 2. 15.1, 48 f49.1, 80.1, 48+49 condition.3. Branch 3. - 42.2, 88.1. Branch 10. 48 f 4 9
- -

condition.5. Branch 11. - 78.Y (24 steps). Branch 13. 88.0. Branch 15. 64.1, 90.1. Branch 16. - 17.0,
- -

75 condition.2. Branch 20. - 48+49.1. Branch 21. 86.1. Branch 22. - 47 condition.2. Branch 23. -
-

87.1. Uranoscodon. 17.0. Tmpidurus umbra. -47 condition.2. Tmpidurusplica. -64.1, 90.1, 47 condition.1.
-

Tmpidurus strobilurus. - 86.1, 88.1. Tropidurus spinulosus. 17.0, 28.0. Zopidurus melanopleurus. - 65.C (1
-

step). Tropidurus callathe&s. 78.Y (24 steps). Tropidurus bogerti. - 6.1, 79.M (12 steps). Tmpidurus semitaeniatus.
-

- 6.2, 79.Q(16 steps). Tmpidurus itambere. - 78.M (12 steps), 84.1. Tmpidurus nanuzae. - 42.1, 88.2.

APPENDIX 4

Specimens examined
Each species examined is followed by the number of specimens in parentheses and their museum
numbers. Specimens for which scales were mounted for scanning electron microscopy are identified
with an asterisk. All specimens were examined for the distribution of scale organs. Museum acronyms
are CBF (Coleccion Boliviana de Fauna, La Paz), CM (Carnegie Museum, Pittsburgh), KU (Natural
History Museum of the University of Kansas, Lawrence), MCZ (Museum of Comparative Zoology,
Harvard University, Cambridge), NKR (Museo de Historia Natural ‘Noel Kempff Mercado’, Santa
Cruz, Bolivia), TCWC (Texas Cooperative Wildlife Collection, College Station), UMMZ (University
of Michigan Museum of Zoology, Ann Arbor), USNM (United States National Museum of Natural
History, Washington), and UTA (University of Texas at Arlington).
Leiocephalus personatus (1 1): TCWC 50670, 5067 1*, 50672-75, 50676*, 50677, 50678*, 50679,
50680*. Leiocephalus schreibersi (10): UTA 18055*, 18057*-59*, 18061*, 18063*-64*, 18066*-68*.
Leiocephalus sailineatus (10): TCWC 50682, 50684*, 50688*, 50692, 50694, 50697-98, 5071 1*, 50712,
507 13*. Liolaemus archeforus (2): TCWC 60042*-43*. Liolaemus chiliensis (2): UTA 30936*-37*. Liolaemus
elongatus (4): TCWC 54947*-48*, 57053*, 60037*. Microlophus occipitalis (27): TCWC 24099*, 24108,
241 10, 241 12, 241 15-16, 241 19-21, 28386-87, 28388*, 28392, 28399,28405, 28407, 28409, 28412,
28414*, 28422, 28424-25, 28426*, 28427, 28433, 28439, 53188. Microlophus peruvianus (31): TCWC
28610-12, 28613*, 28614-16, 28617*, 28618, 28622-23, 28625, 28636-38, 28642-44, 28646,
28650-51, 28654, 28656, 28657*, 28658-59, 53189, 53191-92; UTA 31924*. Micmlophus stolzmanni
(1 7): TCWC 28440*, 28446*, 28447-50,28453-54,28456-57,28460,28462*, 28465,28466*, 28467,
28469, 28472. Microlophus theresiae (17):TCWC 28588*, 28589, 28591-92, 28594*, 28595-96, 28597*,
28600-01, 28602*, 28603-05, 28607-08, 28920. Microlophus thoracicus (7): TCWC 28478-79, 28485,
28492, 28542, 28544, 28545. Micmlophus tigrzs (15): TCWC 28648, 28649*, 28652, 28662, 28667,
28669-70, 2867 1*, 28674, 28676, 28677*, 28678, 28682*, 28683. Plesiomicmlophus koepckeorum (26):
TCWC 2844142, 28444, 28685, 28686*-87*, 28688-93, 28694-97, 28699*, 2870C07, 53186.
Stenocercus caducus (5): CBF uncatalogued* (field tag M.B. Harvey 4738); UTA 38046-48, 39102.
Stenocercus marnoratus (10): UTA 380645*-66*, 391 39-45. Stenocercus fmbriatus (1): TCWC 65928*.
Stenocercus guentheri (1): TCWC 45382*. Stenocercus iridescens (4): UTA 18088*-91*. Tmpidurus azureus (9):
UTA 15388*-96*. Tmpidurus bogerti(2):USNM 300593*-94*. Tmpiduruscallathehs(l8: UTA 39610*-27*
holotype and paratypes. Zopidurus chromatops (14): UTA 39603*-05*, NK 1075*-85* holotype and
 PHYLOGENY OF TROPIDURINI 233

paratypes. Tmpidurus etheridgei (28): C h l ~4595,4595*, 4638-39, 35863-73; UMMZ 51274, 68107 (n=
6), 109104, 129362 (n=5). TmpidurusJavice,bs (2): UTA 3631*, 22583*. Tropidurus guarani (23): USNM
342058-73, 342075-76, 342082, 342084, 348087, 348089, 319758* Paratype. Eopidurus hygomi (2):
MCZ 172874*-75*. Eopidurus hispidus (20): TCWC 58385-86, 58388-92, 58394-96, 58520-2 1; UTA
3014*, 15584*, 32651*. EopiduruJ itambere(2):USNM 148773*, 148775*. Eopidurusmontanus(2):USNM
2 18206*-07*. Tropidurus melunopleurus (19): USNM 28 1589; UTA 37949*-59*; seven uncataloLped
specimens with field tags RLG 1394, 1536*, 1538*, MBH 4010*-1 I*, 4016*, 4771. Tropidurus nanuzae
(2): USNM 213515*, 213517*. Eopidurusoreadicus(4): UMkIZ 57699* (n=4, scale from four specimens
mounted for SEM). Eopidurus semitaeniatus (3):TCWC 53484*-86*. Tmpidurus spinu1osu.r (52): CM p955,
~4593,34836-38, 35874-77, 94223, 142520, MCZ 28633, MVZ 110961, 110974-75, 127091-97,
127099-100, 145148; UMMZ 94092, 109850,68106 (n=2), 169665, 109099-102; USNM 314206-07,
342040-41, 342044-45, 342047-48, 342050-57, UTA 34590*-91*. Tropidurus strobilurus (1): TCWC
53483*. Tmpidurus torquatus(54):UMMZ 55748 (n=2), 55749, 103573, 108599, 108604 (u=6), 108605*
(n=3, scales from two specimens mounted for SEM), 108606* ( n = 6 , scales from one specimen
mounted for SEM), 108607, I08608 ( n = 6 ) , 108609 (n=4), 108610* (n=4, scales from one specimen
mounted for SEM), 149180-89, 167140-41, 204095-101. Tropidurus umbra (13): UTA 3742, 3939,
5035, 521 1, 37991-93, 38016-17, 39606*-09*. 7ropiduru.r xunthochilus (8): UTA 38051*-56*, NK
1086*-87* holotype and paratypes. Tropidurus sp. from Matto Grosso do Sul, Brazil (2): USNM
302999-3000. Uranoscodon superciliosus (6): UTA 3519*, 3738*-41*, 3937*.