A Phylogenetic analysis and taxonomy of iguanian lizards (Reptilia, Squamata)

NIVERSITY OF KANSAS MISCELLANEOUS
M OF NATURAL HISTORY (cana rier

‘e Ut |
8
eS
eet
> ‘gh
~ > A Phylogenetic Analysis and
x a Taxonomy of Iguanian Lizards
+ »
: (Reptilia: Squamata)
)
78
Bed ur
oe
Sea Ge
SE ¢
Le
oie B
eee
Se S Darrel R. Frost and Richard Etheridge
ve
es
ai
re
ie

LAWRENCE September 28, 1989

THE UNIVERSITY OF KANSAS, MUSEUM OF NATURAL HISTORY
PUBLICATIONS

The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in
1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the
above series are now published as The University of Kansas Museum of Natural History Occasional
Papers. The University of Kansas Museum of Natural History Miscellaneous Publications began
with number 1 in 1946. Longer research papers are published in that series. Monographs of the
Museum of Natural History were initiated in 1970. Authors should contact the managing editor
regarding style and submission procedures before manuscript submission. All manuscripts are
subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion
of the Director.

Beginning with this number, this publication is printed on acid-free paper. Occasional Papers and
Miscellaneous Publications are typeset using Microsoft® Word and Aldus PageMaker® on a
Macintosh computer.

Institutional libraries interested in exchanging publications may obtain the Occasional Papers
and Miscellaneous Publications by addressing the Exchange Librarian, The University of Kansas
Library, Lawrence, Kansas 66045-2800, USA. Individuals may purchase separate numbers from
the Office of Publications, Museum of Natural History, The University of Kansas, Lawrence, Kansas
66045-2454, USA.

Museum of Comparative Zoology Library
Harvard University

THE UNIVERSITY OF KANSAS

MUSEUM OF NATURAL HISTORY

MISCELLANEOUS PUBLICATION No. 81
September 28, 1989

A Phylogenetic Analysis and Taxonomy of
Iguanian Lizards (Reptilia: Squamata)

By

DARREL R. Frost! AND RICHARD ETHERIDGE?

'Museum of Natural History, The University of Kansas,
Lawrence, Kansas 66045-2454, USA
*Department of Biology, San Diego State University,
San Diego, California 92187-0057, USA

THE UNIVERSITY OF KANSAS
LAWRENCE
1989

MISCELLANEOUS PUBLICATIONS

Editor: Richard F. Johnston
Managing Editor: Joseph T. Collins
Design and Typesetting: Kate A. Shaw and Joseph T. Collins

MCZ
LIBRARY
OCT 24 1989

HARVARD
UNIVERSITY
Miscellaneous Publication No. 81
Pp. 1-65; 24 figures; 3 appendices
Published September 28, 1989

ISBN: 0-89338-033-4

MUSEUM OF NATURAL HISTORY
THE UNIVERSITY OF KANSAS

LAWRENCE, KaANsAsS 66045-2454, USA

PRINTED BY
UNIVERSITY OF KANSAS PRINTING SERVICE
LAWRENCE, KANSAS

CONTENTS

WPesaT PERC) IBY (6 Cea LO) IN eco, oor ee nse ca cess cesses tee eelctdace ase ts ee euniy Saari ey Suda eoc seeks sScoveaccatatececoes Pe 1
pene O) yD Gr IES Ni Secor coach anes Gusta toy anncsbon Nadavastecstactn.cehcsveate ces teas ideauedencesoessaeentneactonct some 2
iy TESTE QD Saas te ire are Regence cnn Re Or eee Rear as nnn nee ene er ae Pee ee ENP EB 3
cE RLOUICIEI 0) Bie 21 2a] D Me 2 ig 07, © 2 nae ed OE ae i ret ne 4
Wiican Si OPER IAAT ONS RS Si Boe 8 BS, seassstre nth ac sctiocesudatiuds Gua vadeaoseadege acto nave andeneorsedeiorle once Stee ee v
SLE SULTS ais pera os Re Or mi ne PR ore re ACP Aen 16
mcrononts (Agamidae® + AC NamacleOmid ac) ..sss.d.ctveress-sanccadessecseessacssecsvevasecessssacteeesaeacenes sobssets 18
PEMA CONAS Peseta aE Re ag Osh sara tsk MD Maeda ans std gu vaneshlscd Seaoantesadhowsa cease olnes beandedesna te eoeresacee 21
BAAS Md SOUTT EG Sic ee eet eee acne toca nea oes coarae tase area oes Pes ewan secu a Re owcoseecensnducl tena vee en Aa ana eEEE 22

BE eM cMM INULIN reac cccs cia wat te asnsaats sate ch sacs htantaneiSuehtne ck sicauecatcaee vans usdessvotesatevesteetieesotarcartetere ree 23
MMU MRMNTT ESS seen tind, Ate tenes cts A OAPs «Moises Saas avaccmimuce sacs cu cuved oe numioawesetacavisancssuceuse sis sake eee see dene 23

RY HOIANISAGSAUERG eave teens, (bot Sta ete here tet, Sie UE go Ne, SU ee Nes Bote Leen ea eee 23
CONG) ITTY SSS) SS AR AR aes an gt RP SF Ap PEPE HIDE IRSA AR. 7 Hal ntl nt tres, 24

BORG Cote) [ROTI S tee teste reset ere seine es en cL sgus ahha bec stass se 5sses duseaw need weed masses atau cd sevossuneet oesseseameee ss 24
“LUTRS) SYCGUT Er GLAS Pea ane AAR ANAS rane Ss ERE RENE euch PLUME O APR ei 24 ot 25
io PI MUNIMES 2 101 CE CIMS STOW) cheat coca ncticcscessaseaccbacvarecvscavoicaccossssteent easestertomte otcone eee eee 26
Scelopornes + Oplurines + Tropidurines Anoloids) 2.c.:...s<c..sqssecsso0s. ons sesivocccesce-sesse-staenese PH
MINIT AT AO MICS SUIL CS Ao eee cele Aree cack og ROIS. Bs Ae EIS OB os. cos Auch raoasob eee eeen 29
iO NOMIC ACCOUNTS AND CHARACTERIZATIONS: oicccct.-cesesconsseosesetevsssstessttessesteras se ail
Be rramin GC OPern SOA, 21., eteN os ese ua mncemeh s, 5. Jot 2 Spaces ane ates sai oe attack Sd tac Bese neous avene sce 31
Chamacleonidac Ratinesque;, US USX..... .cccsetencesadt oscseee eeescone eee eee eee ee 31
Jade N UCN OVE Fe? S50). Ue 72S fem eek ce I Wa Ne eh refed et 32
Chamacleonimae i atime Saue. 8 V5) asc -.- caters tcoekcoretesece seas cece dices ons ce ee 34
Lciolepidiniae Mitzinger,AB43 & secaceeisscsseseen ete tee ee ee nee 34

Cory tophiagdac Putz per. UGAG Se eee a occ. et secs eave uaciesteeesetaemoce eat csvset ek eae eles aeee 34
Crotaphyndac: Smithand Brodie. 1982 ca; ...:srssvor sels 00 Bictanass soc savess PEtactasce- vant Ae eee 36
Hoplocercidacmew-tamuly i. 0oe: Wael Wee, 2, EE EE 2 CU AS hd, Meee her) le 36

| (aT eee oT a STC (6) 0, TUG Ug a tine ees nn Aol I SRR le sl oma ie 37
Gpilieidac Needy OS 5 5. ee soca tO sce sdac sete ties es iuscactassuunenduscdanelaneneseand 39

PRY MOSOMANGACHIEZAT PET, NSA 3: ss coscsetececs ass ascaccee toe ek Mt 41
Oly c lente ACU UEZ IBS UNS 4 Stee Be Me oases cevaescecut cons cx cet Sceecesest ta cach tac ciasstoet i oshcovensserk 41

Blin UE AS MS CU OAS oc acc, cote sacs feces cance cutee ae AINE ccna aca ech ged eR nce ca OR ea 43
eiocephalinadeanew: subfamily maetecws, 1. Merc scrscseen sesso ves Sane waa roses ere eaecs 45

MEO AeTVMMTAC MEW SEND LATIN UY sea c cece cos sa98 Gack catia es os Seadicts ack cpss ces aetsa sc cdeccaseoeseastcecopeat ese seeeies 45

eUieerpyiilumratnt ae BCU A BAS sce oo cscs ces) cnvacss seesyaoeesat ee atta eee exe Sasa ya ciel Sep eck dae caes es 47

TRE RATURE CET Ds iccecsccceuseconh buzecet eeauschias. bozetad a! cadiuscsscssttensuacconsthassbseucsssseeerecnces sactecsec meee 48
PACE PSINI IEG ca ceaes cect secre eura eet enc tenentenaesyactaesathsccescusssesehve, Sauk se A6oSe tina. cern erate eonseseee eee ee ee 53
PAP PEND IZ cereale dasccesesassottencorsscnaceoota sense cnccaauaszetesuuustioe savek usescuodecneasespeonsdaseuaestecereoteees eee ee ame Y/

NPP NID 3 stvectstesecntsonstanacceesetihon Mente aes coed sae Uaeent , Mea tava eens danasmsaweseats ane sacs cavataloudeseses eaters semen 62

INTRODUCTION

Iguanidae*! is a large family (ca. 54 genera
and 546 species) of lizards found in the Ameri-
cas, the Fiji Island group and the Tongatapu and
Va‘vau groups in the Tonga Islands in the south-
west Pacific Ocean, and Madagascar and the
Comoro Archipelago in the western Indian
Ocean (Fig. 1). The unusual distribution of the
family may be an artifact of paraphyly; to date,
no apomorphies have been presented to support
the monophyly of Iguanidae* exclusive of the
Australo-Afro-Asian Acrodonta (Agamidae* [35
genera, 319 species; Wermuth, 1967; Moody,
1980] + Chamaeleonidae? [6 genera, 128 species;
Klaver and Boéhme, 1986]) (Fig. 1). Indeed,
Schwenk (1988) has suggested that one group of
iguanids (anoles) may be more closely related to
Agamidae* + Chamaeleonidae than to other

'We use an asterisk beside a taxonomic name (the
metataxon convention—Gauthier, 1986; Gauthier et
al., 1988 [cf. Donoghue, 1985]) to denote a nominal
supraspecific taxon for which evidence of either
monophyly or paraphyly is ambiguous or absent.
Although this practice cannot be applied to unitary
lineages (=species) (Frost and Hillis, 1990; but see
Donoghue, 1985, and de Queiroz and Donoghue,
1988) we “flag” some monotypic fossil genera in this
way to denote the lack of character evidence for
grouping specimens under these binomials. Although,
as used here, the metataxon convention is
substantively the same as the shutter quotation
convention of Wiley (1979), in this paper quotations
surround names that represent nominal taxa that are
demonstrably not monophyletic, but whose correction
is outside of the scope of this paper. Because the
historical reality of metataxa is questionable, their
treatment as entities rather than sets in text is arbitrary.
Casual collectives (e.g., agamids) are not asterisked
because they are not formal names and as such are
treated as are other casual collectives (e.g., lizards).

*Because this family-group name is based on the
Latin Chamaeleo, rather than the Greek Chamaeleon,
the formation of the family-group name must be
Chamaeleonidae, rather than the oft-used
Chamaeleontidae.

iguanids. Both Agamidae* and Chamaeleonidae
have been recognized universally by modern sys-
tematists, although monophyly of Agamidae*
remains ambiguous (contra Moody, 1980; Bor-
suk-Bialynicka and Moody, 1984).

Iguania (Iguanidae* + [Agamidae* + Cha-
maeleonidae]) has been established as the sister-
taxon of Scleroglossa (=Scincogekkonomorpha
[Sukhanov, 1961]), the non-iguanian members of
Squamata (1.e., other lizards and snakes) (Camp,
1923; Estes et al., 1988; but see Northcutt, 1978)
and, as such, can be assumed to be of great
antiquity. Even though the earliest fossil iguanid
is from the Upper Cretaceous (Estes and Price,
1973), fossil acrodonts (e.g., +Mimeosaurus*)
are also known from the Upper Cretaceous, and
several scleroglossan squamate groups were
well-diversified by the Upper Jurassic (Estes,
1983a,b). Thus, if the hypotheses of squamate
phylogeny are correct, Iguania must have been
present in the Jurassic.

Major, likely monophyletic, groups within
Iguanidae* were first recognized by Etheridge
(1959, 1964, 1966, 1967), Etheridge in Paull et
al. (1976), and Etheridge and de Queiroz (1988).
As currently understood, Iguanidae* is com-
posed of eight monophyletic groups of uncertain
relationships to each other or to Agamidae* +
Chamaeleonidae. These groups are: (1) anoloids
(11 genera; >200 species); (2) basiliscines (3
genera; 9 species); (3) crotaphytines (2 genera; 7
species); (4) iguanines (8 genera; 25 species); (5)
morunasaurs (3 genera; 11 species); (6) oplurines
(2 genera; 7 species); (7) sceloporines (10 gen-
era; 105 species); and (8) tropidurines (14 gen-
era; 182 species).

The purpose of this study is to reevaluate the
evidence for relationships within Iguania as well
as to investigate the possible paraphyly of Igua-
nidae* with respect to Agamidae* + Chamaele-
onidae, and Agamidae* with respect to Chamae-
leonidae, and to provide a taxonomy logically
consistent (Hull, 1964; Wiley, 1981a) with the
recovered phylogeny within Iguania.

2 MISCELLANEOUS PUBLICATIONS

= Iguanidae*
Agamidae*

oxy
IRS
Xe
OO
dy 4,

i

iv,
XY
OS

Fig. 1. Distribution of Iguanidae*, Agamidae*, and Chamaeleonidae.

ACKNOWLEDGMENTS

For loan of and/or access to specimens in their
care we thank: Charles W. Myers and Richard G.
Zweifel, American Museum of Natural History
(AMNH); Robert C. Drewes, Jacques Gauthier,
Alan E. Leviton, and Jens Vindum, California
Academy of Sciences (CAS); Raymond Laurent,
Fundaci6n Miguel Lillo, Tucuman, Argentina
(FML); José M. Cei, Universidad Nacional de
Rio Cuartos, Cordoba, Argentina (JMC); Wil-
liam E. Duellman, Joseph T. Collins, and John
Simmons, Museum of Natural History, Univer-
sity of Kansas (KU); Robert L. Bezy and John W.
Wright, Natural History Museum of Los Angeles
County (LACM); Douglas A. Rossman, Museum
of Zoology, Louisiana State University
(LSUMZ); Pere Alberch, José Rosado, and
Ernest E. Williams, Museum of Comparative
Zoology, Harvard University (MCZ); Teresa C.
de Avila Pires, Museu Paraense Emilio Goeldi,
Belém, Brazil (MPEG); David B. Wake, Harry
W. Greene, and Stephen D. Busack, Museum of
Vertebrate Zoology, University of California
(MVZ); Paulo E. Vanzolini, Museu de Zoologia,
Universidade de Sao Paulo, Brazil (MZUSP);
Gregory K. Pregill, San Diego Natural History
Museum (SDSNH); Mark Norell, San Diego
(MN); Walter Auffenberg, John B. Iverson, and
Peter Meylan, Florida State Museum, University
of Florida (UF-FSM); Arnold G. Kluge and

Ronald A. Nussbaum, Museum of Zoology,
University of Michigan (UMMZ); Tom Fritts and
Norman Scott, University of New Mexico
(UNM); W. Ronald Heyer, Roy W. McDiarmid,
George R. Zug, Ronald I. Crombie, and Frances
Irish, National Museum of Natural History
(USNM); Jonathan A. Campbell, University of
Texas at Arlington (UTA). The herpetological
collection of San Diego State University (SDSU)
and the osteological collection of R. Etheridge
(REE) were also used extensively in this study.

William E. Duellman, Lynne Frost, Jacques
Gauthier, Philip S. Humphrey, Arnold G. Kluge,
Mathias Lang, Kevin de Queiroz, Gregory K.
Pregill, William Presch, Olivier Rieppel, Linda
Trueb, John Wiens, E. O. Wiley, Ernest E. Wil-
liams, John W. Wright, and George R. Zug read
various parts and editions of the manuscript and
made helpful comments. Anne Musser executed
some of the illustrations. William Presch gener-
ously spent considerable time comparing our
results with those produced by PHYSYS, as did
Arnold Kluge with Hennig86. H. D. Cameron,
Alain Dubois, Philip Tubbs, and Jay M. Savage
assisted with nomenclatural questions, either in
discussion or in correspondence, although this
statement does not necessarily imply their agree-
ment. Errors of logic, observation, and judgment
are ours alone.

IGUANIAN LIZARD PHYLOGENY 3

METHODS

The principles guiding this study are those of
phylogenetic systematics (Hennig, 1966;
Eldredge and Cracraft, 1980; Wiley, 1981b). Out-
group comparison has been selected as the most
general means of deciding on the polarity of
character transformation series (Stevens, 1980;
Watrous and Wheeler, 1981; Armold, 1981; Far-
ris, 1982; Maddison et al., 1984; Kluge, 1985;
Brooks and Wiley, 1985; de Queiroz, 1985).
Following the evidence of Camp (1923),
Gauthier et al. (1988), and Estes et al. (1988),
Scleroglossa (=Scincogekkonomorpha = all non-
iguanian squamates) was selected as the first
taxonomic out-group, and Rhynchocephalia
(including +Gephyrosaurus Evans, 1980, 1981)
was selected as the second taxonomic out-group.
These taxa were used to determine, where pos-
sible, the polarity of particular transformation
series.

Using information from the literature (e.g.,
Moody, 1980; Borsuk-BiaJynicka and Moody,
1984; Arnold, 1984; de Queiroz, 1987; Etheridge
and de Queiroz, 1988; Estes et al., 1988; Lang,
1989), checked and augmented by our observa-
tions (see “Acknowledgments” for reference to
collections in which specimens were examined),
we constructed a character matrix of 67 transfor-
mation series for the operational taxonomic units
selected (see next section). A few traditionally
used characters were excluded from this analy-
sis, either because they were autapomorphies of
the taxonomic units, or because the characters
within the transformation series suffered from
insurmountable characterization problems, i.e.,
they could not be evaluated with any success
across all taxonomic units. The data matrix was
subjected to analysis using PAUP (Phylogenetic
Analysis Using Parsimony) version 2.4.1 (Swof-
ford, 1985) and, late in the development of the
manuscript, Hennig86 version 1.5 (Farris, 1988).
Within PAUP the data matrix was analyzed using
the multiple parsimony (MULPARS) and global
swapping (SWAP = GLOBAL) procedures.
Global swapping allows the program to search
for more parsimonious trees by global (as op-
posed to “nearest neighbor”) swapping of
branches. MULPARS allows the swapping pro-
cedure to be performed on all topologically dis-
tinct trees of a given length, rather than the first

tree found of any particular length. Although
character optimization followed the method of
Farris (1970) (the default in PAUP), use of the
BLRANGE (that calculates maximum and mini-
mum branch links) and CSPOSS (that notes
character ambiguity on all but terminal stems)
options, as well as comparison with the
DELTRAN option (that prefers convergence to
reversal) output, and evaluation of the distribu-
tion of “unknown” character assignments al-
lowed characters of ambiguous placement to be
detected. Within Hennig86, the branch-breaking
(bb) heuristic approach was employed. Tree
optimization was evaluated using the consist-
ency index (C.I.) of Kluge and Farris (1969).
Results from multiple runs of both PAUP and
Hennig86 were evaluated, and alternative root-
ings (i.e., placement of the hypothetical “ances-
tor”) checked using MacClade version 2.1
(Maddison and Maddison, 1987).

We did not regard characters of variable place-
ment to constitute particular evidence of rela-
tionship. That is not to say that we regard the
various character optimization procedures as
equally “likely”; in this case, we were merely
trying to find evidence of relationship that did not
require additional assumptions about how char-
acter evolution proceeds.

Some transformation series were used that
could not be polarized by appeal to out-groups. In
these cases, the “ancestor” cells in the data ma-
trix were coded as unknown (“9” in PAUP; “?” in
Hennig86). These unpolarized transformations
were polarized within the analytical programs by
correlation with the most parsimonious trees
generated by the independently polarized trans-
formations. Because the polarity of a transforma-
tion affects only rooting of the overall network
rather than efficiency of the network, inclusion
of these kinds of transformations is required if we
are trying to find the most parsimonious explana-
tion for all of the data.

Additionally, in some multiple-step transfor-
mations, additivity of the steps was not main-
tained because of lack of evidence of order of
transformation; these transformations were
treated as if any transformation between charac-
ters was one step (“unordered”). In some in-
stances of these “unordered” transformations,

4 MISCELLANEOUS PUBLICATIONS

the ancestral condition could be deduced even
though the polarity of transformations beyond
this initial condition could not be; in this case the
“ancestor” was coded, but the transformation
was treated as unordered. This allowed setting
the initial condition of the transformation(s)
without affecting subsequent transformations
within the unordered set.

In some instances, a character could not be
evaluated for a particular species. Or, in some
cases where we used deduced ancestors as taxo-
nomic units, the ancestral character was not
deducible. In these cases, the character was
coded as “unknown” for those taxa. The analyti-
cal programs allow for this contingency by

making “‘what if” assignments minimally in dis-
agreement with the otherwise most parsimonious
arrangement of the characters on the tree.

We have not employed differential character
weighting. This practice has been argued against
by Patterson (1982) and Novacek (1986). Thus,
the analysis was allowed to proceed as though the
probability of undetected homoplasy was distrib-
uted equally across all transformation series.
However, should the assumption that the plastic-
ity of transformation series was historically equal
prove to be wrong, or our a priori assessments of
homology prove to be wrong, our results will
have to be reevaluated.

CHOICE OF TERMINAL TAXA

A phylogenetic analysis is only as rigorous as
is allowed by: (1) the historical reality (=mono-
phyly) of the terminal taxa; (2) the quality of
characterization of the transformation series; (3)
the accuracy of a priori assessment of homology
and relative historical plasticity of character
transformation; and (4) the appropriateness of
the out-groups employed. Our out-group struc-
ture, although difficult to use because of wide-
spread homoplasy and relative apomorphy, is as
good as current understanding of phylogeny
within Lepidosauria allows (Gauthier e¢ al.,
1988; Estes et al., 1988). Although some charac-
terization problems remain in several of the
transformation series, these have been mini-
mized as much as possible. With regard to termi-
nal taxa, we could use the largest monophyletic
groups that we had confidence were substantially
corroborated. Traditionally, in a taxon as large as
Iguania, this would mean we might employ gen-
era as Our operational taxonomic units. However,
a number of these nominal genera are para-
phyletic, and using these with their derivative
taxa could cause unforeseen problems in data
analysis. Our solution was to use the largest
monophyletic generic and suprageneric groups
for which “ancestral” characters could be de-
duced adequately. To minimize the number of
“unknowns” (“9” in PAUP; “?” in Hennig86) in
the data matrix, we would subdivide terminal
taxa as far as necessary. However, in some cases

these “ancestral” characters could not be de-
duced and the character assignment was “un-
known.” The terminal taxa that we employed
were:

A. Acrodonts.—(1) +Priscagama*; (2) aga-
mas (agamids, excluding Uromastyx, Leiolepis,
and Physignathus); (3) Uromastyx; (4) Leiolepis;
(5) Physignathus; (6) chameleons. Acrodonta has
its monophyly supported by a number of unique
features, including maxillaries in broad contact
behind the premaxilla (Moody, 1980; Estes etal.,
1988). A number of other features support the
monophyly of the group, if it is assumed that the
first functional out-group of this taxon is Igua-
nidae*. Traditionally, within Acrodonta, Agami-
dae* and Chamaeleonidae have been recognized,
with Uromastycidae being recently resurrected,
but subsequently provisionally synonymized
with Agamidae* (see discussion in Borsuk-Bi-
alynicka and Moody, 1984).

Although Uromastycidae has been considered
monophyletic (Moody, 1980, 1983a), the genera
within this group, Uromastyx and Leiolepis, dif-
fer from each other in so many features that we
have treated them as separate taxonomic units.
Physignathus, although hypothesized as the sis-
ter-taxon of the remaining agamids (Moody,
1980 [unweighted analysis]), is also more easily
treated separately. For purposes of deducing
ancestral characters for agamas we provisionally
accepted the phylogeny of Moody (1980).

IGUANIAN LIZARD PHYLOGENY 5

The monophyly of chameleons is supported
by so many features, such as zygodactyl feet,
extremely extensile tongue, failure of the
pterygoid to meet the quadrate (Rieppel, 1981),
and reduction of the number of cervical ribs, that
their monophyly has never been questioned.
Hillenius (1986) and Rieppel (1981, 1987)
considered Brookesia to be the sister-taxon of the
remaining chameleons, but Klaver and BOhme
(1986) considered Brookesia + Rhampholeon to
be the sister-taxon of the remaining chameleons.
We have not entered this discussion and have
accepted only polarity decisions congruent with
both views.

Additionally, we have included the extinct
taxon, +Priscagama* (Borsuk-Bialynicka and
Moody, 1984) in the hopes that inclusion of this
putative agamid would allow a more clear reso-
lution of acrodont phylogeny. At least one fea-
ture, the presence of both the anterior and poste-
rior mylohyoid foramina in the splenial, may
support the monophyly of +Priscagaminae* of
Borsuk-Bialynicka and Moody (1984), includ-
ing, at least, +Priscagama* and +Mimeosaurus*.
However, it might be argued that the anterior
mylohyoid foramen is joined with Meckel’s
groove in Recent acrodonts (a further apomor-
phic condition that renders the monophyly of
+Priscagaminae* arguable). Additionally, the
pleurodont dentition of +Pleurodontagama* Bor-
suk-Bialynicka and Moody (1984) argues for
paraphyly of +Priscagaminae*.

B. Anoloids.—(7) Polychrus; (8) Enyalius;
(9) “Pristidactylus”; (10) para-anoles; (11)
anoles. The monophyly of the anoloids is
corroborated a priori by the character endolym-
phatic sacs extending into the nuchal muscula-
ture (Etheridge, 1959; Etheridge and Williams,
1985; Etheridge and de Queiroz, 1988); this
feature is found otherwise in some chameleons
within Iguania, and geckoes within Scleroglossa.

Polychrus is united by a large number of
characteristics (e.g., third and fourth toes of equal
length), but no hypotheses of phylogeny of the
species have been proposed. Etheridge and de
Queiroz (1988) regarded Polychrus as the sister-
taxon of the other anoloids, but this arrangement
is regarded as provisional.

Enyalius, “Pristidactylus,” Diplolaemus, and

Leiosaurus (including Aperopristis) form a
group, the leiosaurs, that has its monophyly cor-
roborated by the possession of subdigital scales
divided distally (Etheridge and Williams, 1985;
Etheridge and de Queiroz, 1988). Enyalius is the
likely sister-taxon of “Pristidactylus” and its
derivative taxa (Diplolaemus and Leiosaurus)
(Etheridge and de Queiroz, 1988). Following the
phylogenetic arrangement as posited by Eth-
eridge and de Queiroz (1988), ancestral charac-
ters for “Pristidactylus” are coextensive with the
characters of the non-Enyalius group of leio-
saurs, and can be deduced from the Chilean
group of “Pristidactylus” (alvaroi, valeriae, and
torquatus) and the first two in-groups from this:
“P.” casuhatiensis, and “P.” achalensis. Enyalius
was treated as a distinct terminal taxon in order to
avoid some deduced “unknowns” and because of
its provisional association with the other leio-
saurs.

The para-anoles (Urostrophus* and Anisole-
pis [including Aptycholaemus, fide Etheridge and
Williams, unpubl.]) may not form a mono-
phyletic group (although Etheridge and de
Queiroz, 1988, presented some evidence to sup-
port this conclusion); however, our analysis
cannot distinguish between them. They have
been treated together as the para-anoles.

The anoles (Chamaeolis,? Chamaelinorops,
Anolis, and Phenacosaurus) clearly form a
monophyletic group; this is corroborated by the
greatly elongated second ceratobranchials, and
having a distal pad raised under phalanges 2 and
3. Chamaeolis is likely the sister-taxon of the
remaining anoles because it retains a free angular
bone and palatine teeth (Etheridge, 1959). How-
ever, unlike Guyer and Savage (1986), we regard
the phylogenetic structure within the remainder
of the anoles to be problematic (Cannatella and
de Queiroz, 1989).

C. Morunasaurs.—(12) “Enyalioides.” Eth-
eridge and de Queiroz (1988) have documented
convincingly the monophyly of the morunasaurs
(“Enyalioides,” “Morunasaurus,” and Hoplocer-
cus), but have shown also that “Enyalioides” is
paraphyletic with respect to “Morunasaurus,”

3Usually unjustifiably emended to Chamaeleolis.

6 MISCELLANEOUS PUBLICATIONS

which, in turn, is paraphyletic with respect to
Hoplocercus. “Enyalioides” laticeps is the sister-
taxon of “E.” praestabilis + the remaining mo-
runasaurs (Etheridge and de Queiroz, 1988).
Therefore, the evaluation of ancestral characters
within the morunasaur group is based on com-
monalities of these two species. The a priori
assumption of monophyly of the morunasaurs is
supported by the possession of enlarged nasal
scales (Etheridge, 1969b) and greatly reduced
vomers.

D. Basiliscines.—(13) Basiliscus; (14) Co-
rytophanes; (15) Laemanctus. This phenotypi-
cally compact group is supported by one apomor-
phy, the posteriorly extended crest of the parietal
(Etheridge and de Queiroz, 1988; Lang, 1989).
Although other similar crests can be found in
some anoles, some iguanines, and chameleons,
the ontogeny of this crest makes it likely that
these are nonhomologous (Lang, 1989). Because
of the limitations of our deductive methodology
we have considered the three monophyletic gen-
era of basiliscines (Lang, 1989) to be terminal
taxa, although it is reasonably clear that Coryto-
phanes and Laemanctus form the sister-taxon of
Basiliscus (Etheridge and de Queiroz, 1988;
Lang, 1989). Corytophanes has three species,
Laemanctus has only two; and Basiliscus has
four in the relationship B. vittatus + (B. basiliscus
+ B. plumifrons) + B. galeritus (Lang, 1989).

E. Sceloporines.—(16) Petrosaurus; (17)
Sceloporus (including Sator); (18) Urosaurus;
(19) Uta; (20) Phrynosoma; (21) sand lizards
(Uma, Callisaurus, and Holbrookia). The sce-
loporines have their monophyly supported by the
sink-trap nasal apparatus (Stebbins, 1948),
which involves an elongated septomaxilla. Addi-
tionally, they have unique hemipenes (Frost,
1987). Although the cladogram of Presch (1969)
was supported by Etheridge and de Queiroz
(1988), it is clear that the rooting of the tree is
dependent on certain assumptions of out-group
comparison that we do not want to make in this
analysis. Therefore, for analytical reasons, we
treat Petrosaurus, Sceloporus, Urosaurus, Uta,
Phrynosoma, and the sand lizards as our terminal
taxa, each of which is demonstrably mono-
phyletic (Etheridge and de Queiroz, 1988).

F. Tropidurines.—(22) Phymaturus; (23)

Ctenoblepharys;* (24) Liolaemus; (25) Leio-
cephalus; (26) “Stenocercus” + Proctotretus;
(27) “Tropidurus”’; (28) Uranoscodon. The tro-
pidurines can not be supported as monophyletic
a priori by any features relative to all other
iguanians (Etheridge and de Queiroz, 1988), al-
though they are phenotypically similar. Within
the traditional tropidurines, however, a number
of monophyletic groups can be discerned.

The Liolaemus group (Phymaturus,
Ctenoblepharys, and Liolaemus) is supported by
the possession of preanal pores, and although
lacking in a few species, a recent analysis (Eth-
eridge, unpubl.) indicates that this absence is due
to loss. Etheridge and de Queiroz (1988)
regarded Phymaturus as the sister-taxon of all
others, the latter group now with more than 100
species. Although the majority of these are, and
always have been, allocated to the genus Lio-
laemus, in recent years species have been vari-
ously assigned or transferred to the new and
revived genera and subgenera Abas, Ceiolaemus,
Ctenoblepharys, Eulaemus, Ortholaemus,
Pelusaurus, Phrynosaura, Rhytidodeira, Velo-
saura, and Vilcunia (Cei, 1979a,b; Cei and Sco-
laro, 1982; Donoso-Barros, 1972, 1973; Laurent,
1983a,b, 1984, 1985; Nufiez and Yafiez, 1983).
Recent studies by Etheridge (unpubl.) confirm
Laurent’s (1984) decision to consider Ctenoble-
pharys as monotypic (C. adspersus), and the
decision of Etheridge and de Queiroz (1988) to
regard Ctenoblepharys as the sister-taxon of the
remaining species. Although the remaining spe-
cies form a monophyletic group, relationships
within this group are not yet resolved, and in
particular, there appears to be no support for the
continued recognition of Vilcunia. Therefore, we
include all species (i.e., we synonymize all of the
generic names) not in Phymaturus or Ctenoble-
pharys within Liolaemus, which, so constituted,
is monophyletic. Because of character-assign-
ment problems, Phymaturus and Ctenoblepharys
have been considered separately from Lio-
laemus.

Leiocephalus is monophyletic (Etheridge,
1966; Etheridge and de Queiroz, 1988; Pregill,

4 Usually unjustifiably emended to Ctenoblepharis.

IGUANIAN LIZARD PHYLOGENY 7

unpubl.); this is corroborated by unique character
distributions (e.g., parietal shelf shape and ante-
rior process of the interclavicle), and the unique
xiphisternal bars underlying the last sternal ribs?
(Etheridge, 1967).

The “Stenocercus” group (“Stenocercus,”
“Ophryoessoides,” and Proctotretus) 1s sup-
ported by one unique feature, extensive trans-
verse hemipenial musculature (Arnold, 1984).
Because “Stenocercus” is paraphyletic with re-
spect to “Ophryoessoides” (which may be poly-
phyletic) (Frost, 1987), large-scaled “Stenocer-
cus” and Proctotretus were used to determine
ancestral conditions; the fine-scaled species of
“Stenocercus” form a monophyletic group de-
rived from this assemblage (Frost, 1987).

The “Tropidurus” group (Uranoscodon,
“Tropidurus,” Tapinurus, Plica, Strobilurus, and
Uracentron) has its monophyly well corrobo-
rated by a number of apomorphies, none descrip-
tively unique (e.g., enlarged interparietal scale,
enlarged sternum), but not allowing paraphyly
with respect to any other group. For analytical
reasons, Uranoscodon (the sister-taxon of the
remaining “TJropidurus” group [Frost, 1987]) is
treated singly. Because “Tropidurus” west of the
Andes is the sister-taxon of “Tropidurus” east of
the Andes + Tapinurus, Plica, Strobilurus, and
Uracentron (Frost, 1987; B6hme, 1988), “Tro-
pidurus” was used to deduce the ancestral char-
acters for the “TJropidurus” group, excluding
Uranoscodon.

G. Crotaphytines.—(29) Crotaphytus; (30)
Gambelia. The monophyly of the crotaphytines
is not supported by any descriptively unique
features, although they share a combination of

features that, compared with any other iguanian
group, suggests their monophyly. Because of this
lack of a priori support of monophyly, we have
treated Gambelia and Crotaphytus as terminal
taxa.

H. Oplurines.—(31) Oplurus; (32) Chalaro-
don. The oplurines likely form a monophyletic
group, corroborated by the possession of
postxiphisternal inscriptional ribs forming
paired splints (Etheridge, 1965; Etheridge and de
Queiroz, 1988), and a black interparietal spot
(Etheridge, 1969a). Although it is reasonably
clear that Chalarodon and Oplurus are sister-
taxa (but see caveat in Etheridge and de Queiroz,
1988) we treat them separately because deducing
characters for the common ancestor of these taxa
was unclear for several transformation series.

I. Iguanines.—(33) Dipsosaurus; (34) Bra-
chylophus; (35) iguanas (other iguanines). The
monophyly of the iguanines, the large, herbivor-
ous iguanids, seems unassailable. Although
shared with Uromastyx and Hydrosaurus in
Agamidae* (Iverson, 1980, 1982), the presence
of colic septa are likely a synapomorphy of this
group, as is the position of the parietal process of
the supratemporal (de Queiroz, 1987; Etheridge
and de Queiroz, 1988). De Queiroz (1987) and
Etheridge and de Queiroz (1988) argued that
phylogenetically the iguanines have a basal tri-
chotomy with: (1) Dipsosaurus; (2) Brachylo-
phus; and (3) the remaining iguanines ([Ambly-
rhynchus + Conolophus] + Ctenosaura + Sauro-
malus + [Cyclura + Iguana]). These are the three
taxa that we have accepted as terminal for pur-
poses of our analysis.

TRANSFORMATION SERIES

Sources of the various transformation series
are given. Apomorphies of Iguania and a priori
autapomorphies of operational taxonomic units
were excluded from this analysis; monophyly is

assumed (see “Choice of Terminal Taxa’). The

‘ The apparently similar xiphisternal bars in Tapinurus
are associated with the myocommata of the m. pectoralis
major, which is not the case in Letocephalus.

coding “0” denotes the plesiomorphic, and “1”
(or higher) the hypothesized apomorphic charac-
ter, unless the transformation series has been
stated to be unpolarized or unordered, in which
case the integer assignment is arbitrary. A char-
acter assignment of “unknown” refers to the con-
dition being unobservable or of ambiguous as-
signment in that taxon. Character assignments
for the out-groups are shown in Appendix 1
(character matrix).

8 MISCELLANEOUS PUBLICATIONS

1. Premaxilla-nasal relationship (Etheridge,
1966; Etheridge and de Queiroz, 1988).—(0)
premaxillary spine overlaps nasal bones; (1)
nasal bones overlap premaxillary spine. Because
of interspecific variability Phymaturus is coded
as “unknown.”

2. Maxillae (Cope, 1864; Estes et al.,
1988).—(0) do not meet, separated by premax-
illa; (1) meet broadly anteromedially behind
palatal portion of premaxilla.

3. Maxilla, posterior extent (Moody, 1980;
Borsuk-Bialynicka and Moody, 1984).—(0) an-
terior to level of frontoparietal suture; (1) at, or
posterior to, level of frontoparietal suture. Our
observations indicate that Crotaphytus has the
plesiomorphic condition (contra Borsuk-Bialyn-
icka and Moody, 1984). Chameleons are coded as
“unknown” because Brookesia and some Cha-
maeleo have condition “0.”

4. Vomers (Borsuk-BiaJynicka and Moody,
1984).—(0) flat or convex; (1) ventrally con-
cave.

5. Lacrimal (Etheridge and de Queiroz,
1988).—((0) present; (1) absent. Chameleons are
coded as “unknown” because of interspecific
variability (Rieppel, 1981).

6. Lacrimal foramen (Etheridge and de
Queiroz, 1988).—(0) lacrimal foramen not much
larger than maxillopalatine foramen; (1) lacrimal
foramen very much larger than maxillopalatine
foramen.

7. Skull rugosity (Etheridge and de Queiroz,
1988).—(0) absent or restricted to frontal bone;
(1) extensive and matching outline of overlying
scales, found on other dermal skull bones besides
frontal bone and matching outline of overlying
scales. Although some Sceloporus (e.g., S. poin-
setti) have this condition, it is unlikely to be the
ancestral condition in that taxon. Leiocephalus is
coded as “unknown” because of interspecific
variation.

8. Jugal, squamosal contact (Moody, 1980;
Lang, 1989).—(0) not, or barely in contact with
squamosal; (1) broadly juxtaposed against
squamosal along a transverse suture. Chame-
leons are coded as “unknown” because of inter-
specific variability (Rieppel, 1981).

9. Postfrontal (Estes et al., 1988; Etheridge
and de Queiroz, 1988).—(O) present; (1) ex-
tremely small or absent. Phymaturus is coded as

“unknown” because of interspecific variability.

10. Parietal roof shape (Etheridge and de
Queiroz, 1988; Lang, 1989).—(0) trapezoidal;
(1) V or Y-shaped (posteriorly directed crest not
developed); (2) Y-shaped with posteriorly di-
rected median crest developed postembryoni-
cally; (3) Y-shaped with median crest developed
embryonically; roofed at proximal end. Anoles
have been treated as “unknown” because we
regard the assignment of either “0” or “1” to be
ambiguous. The peculiar vaulted parietal shape
of chameleons and the “helmeted” condition of
some anoles are regarded as nonhomologous
with either of the apomorphic conditions here hy-
pothesized.

11. Parietal foramen (Etheridge and de
Queiroz, 1988; Lang, 1989).—(0) at frontopari-
etal suture or in parietal; (1) entirely within the
frontal. Laemanctus is coded as “unknown” be-
cause of interspecific variability. Because they
lack a parietal foramen, chameleons are consid-
ered to be “unknown.”

12. Supratemporal (de Queiroz, 1987; Eth-
eridge and de Queiroz, 1988) (Fig. 2).—(0)
mostly on the lateral or ventral surface of the
supratemporal process of the parietal; (1) mostly
on the medial surface of the supratemporal pro-
cess of the parietal; (2) mostly in a groove in the
ventral margin of the supratemporal process of
the parietal. In addition to the two possible posi-
tions of the supratemporal described by the above
authors, we add a third, found in Liolaemus and
Ctenoblepharys. In these genera the supratem-
poral lies within a groove in the ventral margin of
the supratemporal process of the parietal so all
but its posterior extremity is hidden from medial
and lateral view. As an individual variant in a few
species of Liolaemus, the element is located
within a groove on the lateral surface of the
parietal. Because we cannot polarize this set of
transformations (other than hypothesizing that
“0” is plesiomorphic) we regard this transforma-
tion as unordered. Chameleons are coded as
“unknown” because the supratemporal is a small
splint on the mediocaudal edge of the ventral
ramus of the squamosal and has lost entirely any
connection with the parietal (Rieppel, 1981).

13. Osseous labyrinth (Etheridge and de
Queiroz, 1988).—(0) low to moderate elevation
of the osseous labyrinth above the general level

IGUANIAN LIZARD PHYLOGENY 9

Fig. 2. Supratemporal position. A: supratemporal (in black) substantially on lateral side of supratemporal process of parietal.
B: supratemporal substantially on medial side of supratemporal process of parietal (extent noted by dashed line). C:
supratemporal fits in groove on ventral side of supratemporal process of parietal.

of the opisthotics; (1) high elevation above the
general level of the opisthotics. Care must be
taken in the evaluation of this feature because all
very small iguanids have at least a moderately
elevated labyrinth.

14. Endolymphatic sacs (Etheridge and de
Queiroz, 1988).—(0) do not extend outside of
otic capsule into nuchal musculature; (1) extend
into nuchal musculature. Because some Brooke-
sia (Moody, 1983b), the possible sister-taxon of
other chameleons (Hillenius, 1986; Rieppel,
1987 [but see Klaver and Bé6hme, 1986]), have
the apomorphic condition, chameleons have
been coded as “unknown.”

15. Epiotic foramen (Moody, 1980; Borsuk-
BiaJynicka and Moody, 1984).—(0) absent; (1)
present. Although Moody (1980) hypothesized
that the otic depression in which the epiotic fora-
men sits is an apomorphy of slightly greater
universality, the depression is difficult to charac-
terize across all iguanian terminal taxa (1.e., it is
reasonably well developed in all taxa that have
elevated osseous labyrinths).

16. Dentary, expansion onto labial face of
coronoid (Borsuk-BiaJynicka and Moody, 1984)
(Fig. 3).—(0) dentary does not extend onto labial
face of coronoid; (1) dentary extends onto labial
face of coronoid. Out-group ambiguity (“1” in
rhynchocephalians, “0” in most scleroglossans)
requires that this transformation be treated as
unpolarized.

17. Dentary, posterior extent (Pregill, 1984;
Etheridge and de Queiroz, 1988) (Fig. 3).—(0)
not or only moderately extending posteriorly
beyond level of superior apex of coronoid; (1)
extending posteriorly well beyond apex of coro-
noid. Because rhynchocephalians have condition
“1” and scleroglossans “0,” this transformation is
treated as unpolarized. “Tropidurus” is coded as
“unknown” because of interspecific variability
(those west of the Andes have “0”; east of the
Andes, they have “1”’). Individual and interspeci-
fic variation does not allow further division of
this transformation.

18. Coronoid labial blade (Etheridge, 1966;
Etheridge and de Queiroz, 1988) (Fig. 3).—(0)

10 MISCELLANEOUS PUBLICATIONS

Fig. 3. Labial view of mandibles. Top: Physignathus
lesueuri, KU 69303 (scale=10 mm). Bottom: Leiocephalus
carinatus, UMMZ 149104 (scale=10 mm). Lettered arrows
show: (a) dentary extending onto labial face of coronoid
(Char. 16.1); (b) coronoid labial blade; (c) dentary extending
posterior and superior to anterior surangular foramen (Char.
19.1); (d) fused, acrodont teeth (26.1).

present, large; (1) small or absent. Out-group
ambiguity (“‘1” in many scleroglossans and rhyn-
chocephalians; “0” in most scleroglossans) re-
quires that this transformation be treated as unpo-
larized. Recognition of intermediate steps in this
transformation are obviated by intraspecific vari-
ation.

19. Anterior surangular foramen (Fig. 3).—
(0) anterior surangular foramen posterior to, or
dorsal to posterior extremity of dentary; (1) ante-
rior surangular foramen ventral to posterior ex-
tremity of dentary. In morunasaurs and acro-
donts, the dentary extends posteriorly above the
anterior surangular foramen. Because rhyn-
chocephalians also have this condition, out-
group ambiguity requires that this transforma-
tion be treated as unpolarized.

20. Meckel’s groove (Etheridge and de
Queiroz, 1988) (Fig. 4).—(0) not fused; (1)
fused. Out-group ambiguity (“O” in rhyn-
chocephalians; “1” or “0” in scleroglossans)
requires that this transformation be treated as
unpolarized. The apparently plesiomorphic con-
dition in two Pleistocene species of Leiocephalus
is regarded as a reversal (G. Pregill, pers.
comm.). Chalarodon madagascariensis is indi-
vidually variable and is coded as “unknown.”
Characterization of Phymaturus, Oplurus, Lio-

laemus, and Basiliscus is ambiguous because of
interspecific variation; they are coded as “un-
known.”

21. Splenial, anterior extent (Fig. 4).—(0)
extends anteriorly to or beyond 2 length of tooth
row; (1) does not extend anteriorly more than 2
length of tooth row; (2) does not extend anteri-
orly more than ’% length of tooth row. This trans-
formation series reflects the reduction of the
splenial anteriorly. Because rhynchocephalians
lack a splenial (Evans, 1980, 1981; Estes et al.,
1988) this transformation must be considered un-
polarized. We are unaware how previous authors
determined the identity of the angular (or angulo-
splenial?) in rhynchocephalians. Oplurus and
Liolaemus are sufficiently variable interspecifi-
cally that we have coded them as “unknown.”

22. Splenial, posterior extent (Fig. 4).—(0)
terminates posteriorly anterior to anterior edge of
mandibular fossa; (1) terminates posterior to, or
at anterior edge of mandibular fossa. See com-
ment under previous transformation series. Be-
cause of out-group ambiguity, this transforma-
tion series must be considered unpolarized.
“Enyalioides” is coded as “unknown” because of
in-group variability (“1” in “E.” laticeps; “O” in
other morunasaurs). Corytophanes is also coded
as “unknown” for the same reason (“O” in C.
percarinatus; “1” in C. hernandezi).

23. Angular, condition of contact with
splenial (Etheridge and de Queiroz, 1988) (Fig.
4).—(0) angular large; suture with splenial on
lingual face; (1) angular small; suture with
splenial on ventral or labial face. Because rhyn-
chocephalians lack a splenial, this transforma-
tion series must be treated as unpolarized.
Oplurus is coded as “unknown” because of inter-
specific variation.

24. Posterior mylohyoid foramen (Fig. 4).—
(0) anterior to or approximately at the level of
superior apex of coronoid; (1) between level of
superior apex of coronoid and anterior end of
adductor fossa; (2) posterior to anterior end of
adductor fossa. In rhynchocephalians and Uro-
mastyx, the posterior mylohyoid foramen ap-
pears to be united with the widely open Meckel’s
groove. In scleroglossans, the posterior mylohy-
oid foramen is anterior to the level of the peak of
the coronoid. Therefore, any posterior placement

IGUANIAN LIZARD PHYLOGENY 11

Fig. 4. Lingual view of mandibles. Top: Physignathus
lesueurt, KU 69303 (scale=10 mm). Middle top: Basiliscus
basiliscus, KU 93452 (scale=10 mm). Middle bottom:
Leiocephalus carinatus, UMMZ 149104 (scale=10 mm).
Bottom: Anolis petersi, KU 187446 (scale=10 mm). Lettered
arrows show: (a) unfused Meckel’s canal (Char. 20.0); (b)
splenial extending to or beyond 2 length of tooth row (Char.
21.0); (c) splenial extending less than % length tooth row
(Char. 21.2); (d) angular contacting splenial on lingual face
(Char. 23.0); (€) position of posterior mylohyoid foramen
(Char. 24).

of this foramen is considered derived. Petrosau-
rus and Oplurus are coded as “unknown” be-
cause of interspecific variability (“1” and “2”).

25. Crowns of marginal teeth (de Queiroz,
1987; Etheridge and de Queiroz, 1988).—(0) not
polycuspate; (1) polycuspate. Variation in the
shape of the crowns of the teeth is bewildering,
except for this transformation. Although Brachy-
lophus is variably polycuspate (and then weakly)
it has been coded as polycuspate (de Queiroz,
1987).

26. Posterior maxillary and dentary teeth
(Camp, 1923; Cooper et al., 1970; Estes et al.,
1988) (Fig. 3).—(0) pleurodont, replaced; not
fused to underlying bone; (1) acrodont, not re-

placed as adults; fused to underlying bone.
Because +Gephyrosaurus (the earliest rhyn-
chocephalian) does not have fused, acrodont
teeth, the acrodontan and rhynchocephalian con-
ditions are not considered homologous.

27. Palatine teeth (Moody, 1980; Etheridge
and de Queiroz, 1988).—(0) present; (1) absent.
Because of out-group ambiguity (“1” or “O” in
Scleroglossa; “0” in rhynchocephalians) this
transformation must be considered unpolarized.
Oplurus is coded as “unknown” because of inter-
specific variability.

28. Pterygoid teeth (Moody, 1980; Etheridge
and de Queiroz, 1988).—(0) present; (1) absent.
Polychrus and Leiocephalus are coded as “un-
known” because of interspecific variability.

29. Ceratobranchials (Etheridge and de
Queiroz, 1988).—(0) second not reaching clav-
icles; (1) second reaching clavicles. Chameleons
are considered “unknown” because ceratobran-
chials are lacking.

30. Clavicle (Moody, 1980; Etheridge and de
Queiroz, 1988).—(0) flat, with wide lateral
flange; (1) flange small or absent. Because the
clavicular flange is variably present in the out-
groups, the polarity of this transformation must
be considered unknown. Anoles and Liolaemus
are coded as “unknown” because of interspecific
variability. Chameleons are coded as “unknown”
because they lack clavicles.

31. Insertion of clavicle (Lang, 1989).—(0)
on suprascapula; (1) on scapula. Anoles are
coded as “unknown” because of interspecific
variability. Chameleons are coded as “unknown”
because they lack clavicles. We have treated this
transformation as unpolarized because rhyn-
chocephalians have condition “1” (Evans, 1981).

32. Interclavicle (Camp, 1923; Etheridge,
1966; Lécuru, 1968; Moody, 1980).—(0) ante-
rior process absent; (1) anterior process well
developed. This transformation is treated as
unpolarized because rhynchocephalians lack the
anterior process, whereas scleroglossans have it
plesiomorphically. Chameleons are assigned an
“unknown” because they lack an interclavicle.

33. Sternum, anterior extent (Fig. 5).—(0)
sternum does not approach junction of posterior
and lateral processes of interclavicle closely; (1)
sternum approaches junction of lateral and poste-

12 MISCELLANEOUS PUBLICATIONS

interclavicle

sternum

a b

Fig. 5. Sterna and interclavicles. (a) Dipsosaurus dorsalis,
KU 69107; sternum does not approach juncture of posterior
and lateral processes of interclavicle (Char. 33.0). (b) Plica
plica, M. A. Norell 76; sternum extends to juncture of
posterior and lateral processes of interclavicle (Char. 33.1).

rior processes of interclavicle closely. In some
iguanids the sternum extends anteriorly almost
all the way to the junction of the posterior and
lateral processes of the interclavicle. The more
widespread condition, found in the out-groups, is
for the posterior process of the interclavicle to be
free for a significant part of its length.

34. Caudal vertebral type (Etheridge, 1967;
Etheridge and de Queiroz, 1988).—(0) Scelop-
orus condition (transverse processes anterior to
fracture plane [if present], transverse processes
extend far down length of tail); (1) /guana condi-
tion (fracture plane passes between paired trans-
verse processes); (2) Basiliscus condition
(transverse processes generally not present, in
anomalous conditions when present they are
anterior to fracture plane); (3) anole condition
(transverse processes, if present, posterior to
fracture planes). Because of out-group compari-
son problems, these characters must be regarded
as unordered with respect to each other. Poly-
chrus, lacking fracture planes, could be either
condition “2” or “3,” and is therefore coded as
“unknown.” Para-anoles are coded as
“unknown” because they could be considered
S0COn 6S

35. Scapular fenestra (Etheridge and de
Queiroz, 1988).—(0) present; (1) absent. Out-
group comparison is ambiguous; that is,

scleroglossans are variable in the presence or
absence of a scapular fenestra, although rhyn-
chocephalians clearly lack them. For this reason,
this transformation is considered to be
unpolarized. Polychrus is coded as “unknown”
because of interspecific variability. “Enyalioi-
des” is coded as “unknown” because of deduc-
tive limitations (“1” in “E.” laticeps; “O” in other
morunasaurs). Chameleons are coded as “un-
known” because of interspecific variability and
dubious homology of the fenestrations.

36. Posterior coracoid fenestra (Etheridge
and de Queiroz, 1988).—(0) present; (1) absent;
(2) marginal and weak. Out-group comparison is
ambiguous (i.e., scleroglossans are variable,
even though rhynchocephalians clearly lack
coracoid fenestrae). Therefore, this transforma-
tion is considered unordered. However, many
iguanids that “lack” this fenestra have a thin area
of bone in the shape of this fenestra, which
implies to us that presence is the plesiomorphic
condition within Iguania. The leiosaurs and para-
anoles share the condition of having a small,
peculiar, marginal fenestra in the position of a
posterior coracoid fenestra (condition “2”). Be-
cause we lack any clear justification for the po-
larity between conditions “1” and “2,” we have
considered this transformation to be unordered.

37. Median enlarged sternal fontanelle(s)
(Etheridge, 1964; Moody, 1980; Etheridge and
de Queiroz, 1988).—(0) absent or small, often
hidden by interclavicle; (1) present, large, and
not paired; (2) present, large and paired. Al-
though “0” clearly is the plesiomorphic condi-
tion, “1” and “2” are arguably polarized. There-
fore, these characters are treated as unordered
with respect to each other.

38. Cervical ribs (Etheridge, 1964; Moody,
1980; Etheridge and de Queiroz, 1988).—(0)
first pair on vertebra number 3; (1) first pair on
vertebra number 4; (2) first pair on vertebra
number 5. This transformation is considered
unpolarized because of ambiguous out-group
comparison. Liolaemus, Enyalius, and “Pristi-
dactylus” are coded as “unknown” because of
interspecific variability. Chameleons are coded
as “unknown” because numerical homology of
the cervical vertebrae cannot be determined.

39. Number of sternal ribs (Etheridge and de

IGUANIAN LIZARD PHYLOGENY 13

Queiroz, 1988).—(0) four; (1) three; (2) two or
fewer. Liolaemus is coded as “unknown” because
of interspecific variability (3 or 4 ribs), as are
anoles (3 or 2 ribs).

40. Postxiphisternal inscriptional ribs (Eth-
eridge, 1965; Etheridge and de Queiroz, 1988).—
(0) all attached proximally to dorsal ribs and
none are confluent midventrally; (1) one or more
pairs attached to dorsal ribs and are confluent
midventrally; (2) none attached to dorsal ribs or
continuous midventrally; present as pairs of iso-
lated elements. Although “0” is clearly ple-
siomorphic, the polarity of “1” to “2” cannot be
determined. Therefore, this transformation series
is considered unordered.

41. Tail autotomy fracture planes (Estes et
al., 1988; Etheridge and de Queiroz, 1988).—(0)
present; (1) absent. Uromastyx is assigned an
“unknown” because at least some specimens of
U. acanthinurus have functional fracture planes
(Etheridge, pers. observ.). Enyalius and anoles
are coded as “unknown” because of interspecific
variability.

42. Interparietal scale (Smith, 1946; Eth-
eridge and de Queiroz, 1988).—(0) small (or
absent); (1) large; as wide as interorbital space.
Sand lizards have been assigned an “unknown”
because of in-group variability (i.e., Uma has a
small interparietal). In spite of our coding, on
anatomical grounds it is questionable whether
the enlarged interparietal of sceloporines is ho-
mologous with the “enlarged” interparietal found
in some tropidurines; particularly in Urano-
scodon and “Tropidurus” west of the Andes,
evidence of edge-to-edge fusion of scales is fre-
quently obvious.

43. Interparietal coloration (Etheridge,
1969a).—(0) black spot absent; (1) black spot
present. Although the apomorphic condition
appears in some other iguanian species (e.g.,
Sceloporus nelsoni), it is not ancestral in any
other terminal taxon except the oplurines.

44. Superciliary scales (Etheridge and de
Queiroz, 1988).—(0) distinctly elongate and
imbricate; (1) not distinctly elongate and imbri-
cate. Ambiguous out-group comparison
(Sphenodon has condition “1,” but scleroglos-
sans are not really comparable) requires use of
this transformation as non-polarized. Because of

interspecific variation, Phymaturus is coded as
“unknown.”

45. Subocular scale (Etheridge and de
Queiroz, 1988).—(0) at least one scale below the
eye conspicuously enlarged; (1) scales below the
eye subequal. Out-group comparison does not
support a particular polarity of this transforma-
tion; it is therefore considered as unpolarized
(Sphenodon has subequal subocular squamation;
scleroglossans are variable). Because of inter-
specific variability (“O” in E. bilineatus; “1” in
other species), Enyalius is coded as “unknown”
as are the para-anoles (“0” in Anisolepis; “1” in
Urostrophus*), Liolaemus, and Phymaturus (“0”
in P. patagonicus; “1” in P. palluma).

46. Mid-dorsal scale row (Etheridge and de
Queiroz, 1988; Estes et al., 1988).—(0) present;
(1) absent. We have not addressed differences of
development of the median dorsal crest because
of complex intra- and interspecific variation.
Polychrus and anoles are considered “unknown”
because of interspecific variation. Because of
out-group ambiguity (“O” in Sphenodon; “1” in
Scleroglossa) this transformation is treated as
unpolarized.

47. Gular fold (Etheridge and de Queiroz,
1988).—(0) complete medially; (1) incomplete
medially or absent. Because some species (e.g.,
Polychrus femoralis) have “gular folds” that lack
any kind of distinctive change in squamation at
the fold line, we have restricted the use of “gular
fold” to those species that have a distinct change
in squamation at the level of the fold. For this
reason, the condition found in Polychrus fem-
oralis is considered “1” and the condition in
Laemanctus is considered “0.”

48. Femoral pores (Camp, 1923; Etheridge
and de Queiroz, 1988).—(0) present; (1) absent.
Out-group ambiguity (Sphenodon lacks femoral
pores but Scleroglossa has them plesiomorphi-
cally) requires the treatment of this transforma-
tion as unpolarized, even though this feature has
been considered a synapomorphy of Squamata
(Kluge, 1983; Gauthier et al., 1988).

49. Preanal pores (Laurent, 1984; Etheridge
and de Queiroz, 1988).—(0) absent; (1) present.

50. Distal subdigital scales (Etheridge and de
Queiroz, 1988).—(0) undivided; (1) divided.
Although Sphenodon lacks regular subdigital

14 MISCELLANEOUS PUBLICATIONS

scales, it clearly lacks the apomorphic condition
here specified, and is therefore regarded as hav-
ing condition “0.”

51. Subdigital scale surface macrostructure
(Peterson and Williams, 1981; Etheridge and de
Queiroz, 1988).—(0) carinate; (1) smooth. Out-
group ambiguity (Sphenodon has smooth sub-
digital scales even though scleroglossans usually
have carinate subdigitals) requires the treatment
of this transformation as_ unpolarized.
Chameleons, Enyalius, and “Pristidactylus” are
coded as “unknown” because of interspecific
variation.

52. Scale organs (Peterson, 1983; Etheridge
and de Queiroz, 1988; E. E. Williams, pers.
comm.).—(0) spinules absent; (1) spinules pres-
ent. We have simplified the transformation series
of Etheridge and de Queiroz (1988) in order to
obviate some out-group comparison problems.
Chameleons are assigned an “unknown” because
of interspecific variation.

53. Nasal chamber, sink trap (Stebbins,
1948).—(0) primitive condition (short vestibule,
concha well developed) or some apomorphic
condition not homologous with Character 53.1;
(1) sink-trap (elongate septomaxilla) (condition
*1 of following discussion).

Malan (1946), in her study of the comparative
anatomy of the lacertilian nasal capsule, pro-
vided a solid framework to which other contribu-
tions were made by Stebbins (1948) and Stimie
(1966). Although Malan’s work was the most
detailed, for purposes of this discussion we use
the nomenclature and points of reference of Steb-
bins (1943, 1948) because his work is most di-
rectly applicable to our own observations.

Primitively, saurians have a relatively long
vestibule leading from the external naris to the
nasal cavity. This vestibule is lined with erectile
tissue (Lapage, 1926; Malan, 1946), which has
been hypertrophied to form nasal valves in vari-
ous lineages. The vestibule attaches anterodor-
sally to the nasal cavity, which is divided sagit-
tally by a “tongue” of tissue, the concha, that
projects medially into the nasal cavity. More or
less hidden from dorsal view, beneath the con-
cha, the slit-like internal nares communicate with
the oral cavity. Behind the internal choana and
the concha is a blind cavity, the antorbital cham-
ber. This condition obtains in Sphenodon and in

various degrees of modification in most
scleroglossans.

In iguanians there are five major deviations
from this pattern (discussed under subsequent
transformation series): (1) sink-trap; (2) “S”
condition; (3) fusion of nasal concha to chamber
roof (=reduction of supraconchal part of nasal
chamber); (4) anole condition; (5) acrodontan
condition.

The sink-trap nasal apparatus of the sce-
loporines seems to have been derived from the
primitive condition by more or less direct poste-
rior elongation of the vestibule (which is sup-
ported by an equally apomorphic elongate septo-
maxilla) to enter the nasal cavity at the postero-
dorsal end.

54. Nasal chamber, S-condition (Stebbins,
1948).—(0) primitive condition, or apomorphic
condition not homologous with Character 54.1;
(1) S-condition (septomaxilla plow-share
shaped) (condition *2 of discussion under “‘Trans-
formation Series 53”). In the S-condition the
vestibule is elongate, S-shaped, and overlies the
nasal cavity. In all cases, the septomaxilla ex-
tends dorsally to contact the osseous roof of the
nasal cavity, and is shaped like a plow-share, a
condition otherwise unknown in lizards. Phylo-
genetically, we hypothesize that the S-condition
was derived from the primitive condition by
simple elongation of the vestibule over the nasal
cavity, whereby the vestibule opens into the nasal
cavity dorsomedially rather than in the primitive
anterodorsal position.

55. Nasal chamber, fusion of nasal concha
to roof of nasal chamber.—(0) primitive condi-
tion, or other apomorphic condition not homolo-
gous with Character 55.1; (1) fusion of concha to
roof of nasal chamber. In the “Stenocercus” and
the “Tropidurus” groups the primitive condition
is largely retained, except that the concha is fused
to the roof of the nasal chamber (condition *3 of
discussion under “Transformation Series 53”). In
some species, the vestibule is slightly elongated
and enters the nasal chamber at a relatively high
level.

56. Nasal chamber, anole condition (Steb-
bins, 1948).—(0) primitive condition, or some
apomorphic condition not homologous with
Character 56.1; (1) nasal concha lost, with nasal
chamber otherwise retaining plesiomorphic or-

IGUANIAN LIZARD PHYLOGENY 15

ganization (condition *4 in discussion under
“Transformation Series 53”). In Anolis, and pre-
sumably their near relatives, the concha is lost
(Malan, 1946; Stimie, 1966), although otherwise
the plesiomorphic condition is maintained. The
concha also is missing in Polychrus, but present
(although weak) in Diplolaemus and “‘Pristidac-
tylus.” Para-anoles and Enyalius are coded as
“unknown” because rarity in museum collections
precludes dissection.

57. Nasal chamber, acrodontan condition
(Malan, 1946; Parsons, 1970; SJaby, 1981,
1984).—(0) primitive nasal condition, or some
apomorphic condition not homologous with
Character 57.1; (1) reduction of concha con-
comitant with elongation of the nasal vestibule
(condition *5 in discussion under “Transforma-
tion Series 53”). Agamidae* (except Physi-
gnathus which has the primitive iguanian pattern
of having a relatively short nasal vestibule and a
small nasal concha) and chameleons have an
unusual condition in which there is a long vesti-
bule extending from a lateral or dorsolateral naris
over the nasal chamber and enters the nasal cham-
ber posterodorsally (Parsons, 1970). The nasal
concha is very small (Leiolepis) or absent (e.g.,
Uromastyx, “Agama,” chameleons). In some
aspects, the acrodontan condition is intermediate
between the “S” condition and the sink-trap, but
the unusual septomaxillary anatomy in all three
conditions (very long in the sink-trap; plow-
share shaped in the “S” condition; and very small
or absent in the acrodontan condition) argue for
nonhomology. Chameleons have modified the
agamid condition in a number of ways that seem
to be correlated with enlargement of the eyes and
tongue (Malan, 1946).

58. Ulnar nerve pathway (Jullien and Re-
nous-Lécuru, 1972; Renous, 1979; Estes, 1983a;
Etheridge and de Queiroz, 1988).—(0) L-condi-
tion (superficial); (1) V-condition (deep). Out-
group ambiguity requires use as unpolarized.

59. Dorsal shank muscle innervation (Jul-
lien and Renous-Lécuru, 1972; Renous, 1979;
Etheridge and de Queiroz, 1988).—(0) A-condi-
tion (peroneus); (1) B-condition (interosseus).
Out-group ambiguity requires use as unpolar-
ized.

60. Hemipenis, posterior lobe (Fig. 6).—(0)
no enlarged posterior lobe; (1) enlarged posterior

—~ .

aJf~ SS
en a

SOS

Rayer
EA, (ewes ES
ON A CSI LOH
Soak, S85
AIAa OO 2

by
a3).

Fig. 6. Hemipenes. (a) Sceloporus torquatus, KU 91414,
showing enlarged posterior lobe (Char. 60.1). (b)
“Stenocercus” festae, KU 134588, showing bilobate,
bisulcate condition (Char. 61.1). (c) Plica umbra, KU
147946, showing bicapitate, bisulcate condition (Char. 61.2).

16 MISCELLANEOUS PUBLICATIONS

lobe. In sceloporines, a posterior eminence, nor-
mally present but small in other lizards, is en-
larged to the point that in superficial examination
of the sceloporine hemipenis it seems to have a
posterior median lobe. Because Sphenodon lacks
a hemipenis, this transformation must be re-
garded as unpolarized.

61. Hemipenis, capitation and sulci (Fig.
6).—(0) unicapitate or weakly bilobate without
distinctly divided sulci; (1) bilobate with dis-
tinctly divided sulci; (2) strongly bicapitate. The
only members of the first out-group similar
enough to be comparable, teiids, support the
polarity 01-42. However, because Sphenodon
lacks a hemipenis, this transformation must be
regarded as unpolarized.

62. Hemipenis, m. retractor lateralis
posterior (Arnold, 1984).—(0) not completely
divided; (1) completely divided. Because
Sphenodon lacks a hemipenis, this transforma-
tion must be regarded as unpolarized.

63. Hemipenis, m. retractor lateralis poste-
rior (Arnold, 1984).—(0) not substantially situ-
ated within the hemipenial sheath; (1) substan-

tially situated within the hemipenial sheath.
Because Sphenodon lacks ahemipenis, this trans-
formation must be regarded as unpolarized.

64. Hemipenis, dorsal accessory sheath
muscle (Arnold, 1984).—(O) absent; (1) present.
Because Sphenodon lacks a hemipenis, this trans-
formation must be regarded as unpolarized.

65. Colic septa (Lénnberg, 1902; El Taubi
and Bishai, 1959; Iverson, 1980, 1982).—(0)
absent; (1) present. Agamas are coded as “un-
known” because of the presence of colic septa in
Hydrosaurus. —

66. Paired ventrolateral belly patches in
males (Etheridge and de Queiroz, 1988).—(0)
absent; (1) present. “Stenocercus” is coded as
“unknown” because some large-scaled species
(e.g., “S.” rhodomelas) have paired ventrolateral
patches.

67. Reticular papillae on tongue (Schwenk,
1988).—(0) absent; (1) present. Because we
depended entirely on the literature for this trans-
formation (suggested to be synapomorphy of
anoles and acrodonts), a number of taxa had to be
coded as “unknown.”

RESULTS

A total of 225 alternative supported (as op-
posed to unrejected) tree topologies were discov-
ered (208 steps; C.I.=0.385). Twelve networks
were discovered that could be variously rooted to
produce 18 unique trees of nine major mono-
phyletic groups (acrodonts; anoloids; basilis-
cines; crotaphytines; iguanines; morunasaurs;
oplurines; sceloporines; and tropidurines) (Fig.
7). Within these monophyletic groups alternative
topologies exist that are variously independent to
dependent on intergroup topology. Within the
Liolaemus group of the tropidurines two topolo-
gies were discovered, three in the sceloporines,
two in the acrodonts, and three in the anoloids.

A strict consensus tree (Nelson, 1979) of the
discovered tree topologies is presented in Figure
8. This consensus tree is not a parsimonious
solution of the data, but only a figure showing the
commonalities among the discovered trees. The
extensive polytomies seen in the consensus tree
are due both to variation in rooting points within
networks and topological differences among

equally parsimonious unrooted networks. Re-
gardless of the impression given by the consen-
sus tree, notall of the phylogenetic trees logically
consistent with the strict consensus tree are, in
fact, allowed within the constraints of the discov-
ered network topologies. Many trees are ex-
cluded (e.g., those showing a sister-taxon rela-
tionship between acrodonts and crotaphytines).
If attainment of a single tree is the only measure
of progress in the understanding of iguanian rela-
tionships, we have failed egregiously. However,
the number of possible dichotomous trees for 35
in-group taxa is 4.89 x 10 *’ (Felsenstein, 1978).
Because we have rejected all but 549 dichoto-
mous 208-step trees (i.e., all but 1.12 x 10°*% of
the total possible), we consider that we have
made great progress, indeed.

It is clearly impossible because of space con-
siderations to discuss the character support for
each topology. Therefore, only evidence for
major monophyletic groups and organization
within them will be discussed. In order to docu-

IGUANIAN LIZARD PHYLOGENY 17

MO A BA MO 5

AC IG
TR

GR. SC TR

AN

MO AC C

IG BA
GRE Sse » TR

AN OP

MO E MO Ie

AC IG
BA IG CR BA AC CR

AN

MO AC
IG BA G
SC TR
CR
AN OP
IG |

MO
CR-SC TR

AC

MO
BA CR SC AN OP

AC TR

AC BA IG MO CR TR SC OP AN

1

Fig. 7. Discovered unrooted networks with discovered rooting points (-o-). AC=acrodont groups (agamines, uromastycines,
and chameleons); AN=anoloids; BA=basiliscines; CR=crotaphytines; IG=iguanines; MO=morunasaurs; OP=oplurines;
SC=sceloporines; TR=tropidurines. Arrow points to rooting point (position of ancestor vector) that results in tree 1.

18 MISCELLANEOUS PUBLICATIONS

tPriscagama
Agamas
Physignathus
Uromastyx
Leiolepis
Chameleons
Polychrus
Anoles
Para-anoles
Enyalius
"Pristidactylus"
"Enyalioides"
Basiliscus
Corytophanes
Laemanctus
Petrosaurus
Uta
Sceloporus
Urosaurus
Phrynosoma
Sand lizards
Phymaturus
Ctenoblepharys
Liolaemus
Leiocephalus
"Stenocercus"
"Tropidurus"
Uranoscodon
Oplurus
Chalarodon
Crotaphytus
Gambelia
Dipsosaurus
Brachylophus
Iguanas

Fig. 8. Strict consensus tree (Nelson, 1979) of terminal taxa.

ment the degree of homoplasy required by all
discovered trees, one tree, arbitrarily selected, is
completely documented in Figure 9 and Appen-
dices 2 and 3.

ACRODONTS

As expected, Agamidae* and Chamaele-
onidae together form a monophyletic group, al-
though two topologies were discovered (Fig. 10).
Topology 1 was discovered in intergroup Net-
works A-D and F-K, but Topology 2 was discov-
ered only in Networks B, E, and L (Fig. 7).

Topology 1.—The monophyly of the agama
operational taxonomic unit (=agamids,
excluding Physignathus, Uromastyx, and Leiole-
pis) may be supported by 32.1 (well-developed
anterior process of interclavicle), however, the
interclavicle is lost in chameleons and is un-
known in +Priscagama*. Therefore, the alterna-
tive must be entertained that 32.1 is plesiomor-
phic in this clade with a reversal in Leiolepis and
Physignathus. Physignathus apparently has a
reversal to 30.0 (flat clavicle with a wide lateral
flange) although this is also an ambiguous place-
ment because the clavicle is absent in chame-
leons and unknown in +Priscagama*. Stem 1
(agamas + Physignathus) is corroborated by 15.1
(epiotic foramen) and is in agreement with the
results of Moody (1980). Also, 31.1 (insertion of
clavicle on scapula) may belong here, but place-
ment is made ambiguous by the lack of a clavicle
in chameleons and being unknown in +Pris-
cagama*. Chameleon monophyly is supported
by a number of characters that are apomorphies
in all topologies: 39.2 (strong reduction of num-
ber of sternal ribs), 47.1 (loss of gular fold), 48.1
(loss of femoral pores), and 58.1 (V-condition of
ulnar nerve pathway). In the networks that sup-
port this topology, 7.1 (extensive skull rugosity)
and 40.1 (midventrally confluent postxiphister-
nal inscriptional ribs) also are apomorphic.

Stem 2 carries unambiguously only one char-
acter that supports the monophyly of the chame-
leons + agamids, excluding Leiolepis and Uro-
mastyx: 6.1 (extremely enlarged lacrimal fora-
men). Although this condition as characterized is
shared with morunasaurs, both chameleons and
agamids (other than the uromastycines) have
these foramina more enlarged than in moruna-

IGUANIAN LIZARD PHYLOGENY

5
3
6 4 e
11
9
10
14
8
7
13
30
31 28
a 16
18 15
17
27
25
26
23
22
24
2
20
19

tPriscagama
Agamas
Physignathus
Uromastyx
Leiolepis
Chameleons
"Enyalioides"
Enyalius
"Pristidactylus"
Para-anoles
Polychrus
Anoles
Basiliscus
Corytophanes
Laemanctus
Iguanas
Brachylophus
Dipsosaurus
Crotaphytus
Gambelia
Petrosaurus
Uta
Urosaurus
Sceloporus
Sand lizards
Phrynosoma
Oplurus
Chalarodon
Phymaturus
Ctenoblepharys
Liolaemus
Leiocephalus
"Stenocercus"
"Tropidurus"
Uranoscodon

19

Fig. 9. A 208-step tree selected arbitarily from among those discovered. Character shifts for this tree are documented in

Appendices 2 and 3.

20 MISCELLANEOUS PUBLICATIONS

Topology 1

Agamas
Physignathus
Chameleons
Uromastyx

Leiolepis

'Priscagama

Topology 2

Agamas
Physignathus
Uromastyx
Leiolepis

Chameleons

Priscagama

Fig. 10. Alternative topologies dicovered for the acrodont
groups.

saurs and extremely enlarged in contrast to the
condition found in uromastycines, which have
the primitive condition. Another character that
“falls out” on this stem is 39.1 (reduction of
sternal ribs from 4 to 3), although in some topolo-
gies this is made ambiguous by the sternal rib
number being unknown in +Priscagama*.

The monophyly of Uromastyx is supported
by: 5.1 (loss of the lacrima]) and 65.1 (appear-
ance of colic septa—also in Hydrosaurus and
iguanines). Character 36.0 (anterior process of
interclavicle) is placed here ambiguously; see
previous discussion. Leiolepis is corroborated by

62.1 (completely divided m. retractor lateralis
posterior). Stem 3 (uniting Leiolepis and Uro-
mastyx) is supported by only one unique, un-
reversed character, 4.1 (cup-shaped vomers),
although in some arrangements three other fea-
tures “fall out” on this stem: 3.0 (reduction of
posterior extent of maxilla), 39.0 (four sternal
ribs), and 46.1 (mid-dorsal enlarged dorsal scale
row). 3.0 is rendered ambiguous by variation
within chameleons, 39.0 is rendered ambiguous
by network topology, and 46 is rendered ambigu-
ous by being unknown in +Priscagama*.

Stem 4 (acrodonts above +Priscagama*) is
supported by three unambiguously placed apo-
morphies: 16.1 (expansion of dentary onto labial
face of coronoid), 17.1 (far posterior extension of
the dentary), 21.1 (shortening of the splenial),
and 28.1 (loss of pterygoid teeth).

+Priscagama* has no unambiguously placed
apomorphies. Stem 5 (the acrodont groups), as
noted in “Choice of Terminal Taxa,” is well-
corroborated by 2.1 (maxillae meet anteromedi-
ally behind palatal portion of the premaxilla) and
26.1 (acrodont maxillary and dentary teeth, fused
in adults). Additionally, several characters of
ambiguous placement may belong on this stem:
3.1 (posterior extent of maxilla posterior to fron-
toparietal suture) is variable in chameleons and
likely reversed in Uromastyx and Leiolepis; 9.1
(postfrontal reduced) in some topologies is of
greater universality; 19.1 (anterior surangular
foramen ventral to posterior extremity of den-
tary) in some topologies has a greater level of
universality (shared with morunasaurs); 37.1
(large, paired sternal fontanelles) is ambiguous
because it is unknown in +Priscagama* and be-
cause of the extreme sternal modification in
chameleons; 57.1 (acrodontan nasal condition
[reversed in Physignathus]) and 67.1 (reticular
lingual papillae) are also unknown in +Pris-
cagama*.

Topology 2.—Agamas, Physignathus,
chameleons, Uromastyx, Leiolepis, and stems 4
and 5 are as in Topology 1. Stem 6 (Leiolepis +
Uromastyx) in this topology is nearly the same as
Stem 3 as in Topology 1, except that the lacrimal
foramina (6.0) are secondarily reduced (enlarge-
ment being an apomorphy of possible extra-acro-
dont universality). Stem 7, supporting a mono-

IGUANIAN LIZARD PHYLOGENY 21

phyletic Agamidae, carries 37.2 (paired, enlarged
sternal fontanelles), although this is ambiguous
because of strong modification of chameleon
sterna and being unknown in +Priscagama*, and
a reversal to 40.0 (short postxiphisternal inscrip-
tional ribs) that is dependent on the topology of
the intergroup network.

For no reason other than the comparative rar-
ity of topologies that allow the chameleons to
form the sister-taxon of the remaining acrodonts,
we suspect that chameleons are nested within the
traditional Agamidae*.

There are, of course, some general iguanian
topologies that do not preclude the acrodont taxa,
Agamidae* and Chamaeleonidae, from forming
the sister-taxon of Iguanidae* (Fig. 7). In these
trees no putative synapomorphies of Iguanidae*
are unambiguously placed, and the characters
that are placed ambiguously are all unordered or
unpolarized characters that were assigned arbi-
trarily by the computer program: 19.0 (anterior
surangular foramen posterior to, or dorsal to
posterior extremity of dentary [Ancestor as-
signed 19.1—posterior mylohyoid foramen ven-
tral to posterior extremity of dentary]), 30.0
(clavicular flange flat, with wide lateral flange
[Ancestor assigned 30.1—clavicular flange re-
duced or absent]), 38.1 (posterior coracoid
fenestra absent), and 63.1 (m. retractor lateralis
posterior not substantially situated within the
hemipenial sheath [Ancestor assigned 63.0]).

ANOLOIDS

The anoloid genera of Etheridge and de
Queiroz (1988) formed a monophyletic group in
all obtained trees and formed three topologies
(Fig. 11). Additionally, because para-anoles do
not have their monophyly supported unambigu-
ously it is conceivable that Urostrophus* and
Anisolepis are more closely related to other
anoloid genera than to each other. All three
anoloid topologies differ from the cladogram
presented by Etheridge and de Queiroz (1988),
primarily in the placement of Polychrus. Eth-
eridge and de Queiroz (1988) considered Poly-
chrus to be the sister-taxon of other anoloids (on
the basis of its having femoral pores and non-
spinulate scale organs), whereas we consider it to
be the sister-taxon of anoles. The change of

placement apparently requires the reacquisition
of femoral pores in Polychrus as well as the loss
of an otherwise ubiquitous feature of anoloids,
spinulate scale organs.

Topology 1 was discovered in Networks A-H
(Fig. 7); Topology 2 was discovered in Networks
A-B and E; and Topology 3 was discovered in
Networks A-B, J—-L (but not in A and B when
rooted such that anoloids are in polytomy with
the remaining iguanians). Polychrus was sup-

Topology 1

Polychrus
Anoles
Para-anoles

Enyalius

"Pristidactylus"
Topology 2
Polychrus
1 Anoles
2 Para-anoles
4 Enyalius
5 "Pristidactylus"
Topology 3
Polychrus
| Anoles
4 Para-anoles
Enyalius

5 "Pristidactylus"

Fig. 11. Alternative topologies discovered within anoloids.

22 MISCELLANEOUS PUBLICATIONS

ported in all trees by three unambiguously placed
characters: 5.1 (loss of lacrimal), 48.0 (regaining
of femoral pores), and 52.0 (loss of spinulate
scale organs). Additionally in Topologies 1 and 2,
27.1 (loss of palatine teeth) is placed on this stem.
Regaining of carinate subdigital scales (51.0)
possibly is a feature of Polychrus, but because
this feature is variable in “Pristidactylus” and
Enyalius, its placement is ambiguous.

The monophyly of anoles is supported by a
number of features in all three topologies: 17.1
(far posterior extension of dentary), 18.0 (regain-
ing of coronoid labial blade), 21.2 (little anterior
extension of splenial), 23.1 (reduced angular),
24.2 (far posterior placement of posterior my-
lohyoid foramen), and 67.1 (reticular lingual
papillae). Stem 1 (anoles + Polychrus) is cor-
roborated by four unambiguously placed charac-
ters: 29.1 (second ceratobranchial extends to
clavicles), 33.1 (anterior process of interclav-
icle), 38.2 (most anterior cervical ribs on verte-
bra 5), and 47.1 (loss of gular fold). Other, more
ambiguously placed features possibly on this
stem are: 30.1 (clavicular flange reduced), 31.1
(clavicular insertion on scapula), and 39.2 (< 2
sternal ribs), which are variable in anoles. Addi-
tionally, Ernest E. Williams (pers. comm.) has
noted that Polychrus and the anoles share a di-
vided mental scale.

The para-anoles (Urostrophus* and Anisole-
pis) are not united by any apomorphies whose
placement is independent of network, but para-
anoles, Polychrus, and anoles may have a rela-
tionship supported in Topologies 1 and 2 (Fig.
11—Stem 2) by 39.1 (3 sternal ribs), 41.1 (loss of
tail autotomy—teversed in anoles), and ambigu-
ously by 34.3 (anole caudal vertebral type),
which is difficult to evaluate in para-anoles
(which are either “O” or “3”) and Polychrus
(either “2” or “3”). Alternatively, when oplurines
or acrodonts are considered the sister-taxon of
anoloids (Fig. 11—Stem 6), 36.2 (marginal and
weak posterior coracoid fenestra), may support a
special relationship of para-anoles with Enyalius
+ “Pristidactylus.”

Enyalius is supported by 17.1 (far posterior
extension of dentary [also seen in anoles]) and
64.1 (hemipenial accessory sheath muscle). In

Topology 1 Enyalius is linked (Stem 3) with the
anoles, para-anoles, and Polychrus by an ele-
vated osseous labyrinth (13.1) (and possibly by a
widened frontal); in Topology 2 and 3 Enyalius is
linked (Stem 5) with “Pristidactylus” by 50.1
(divided terminal subdigital scales). Because the
condition is unknown in para-anoles and Enya-
lius, all that can be said about the nasal condition
(Transformation Series 56) is that either some-
where between “Pristidactylus” (56.0) and
anoles + Polychrus (56.1), the nasal concha is
lost, or, conversely, loss of the nasal concha may
be a synapomorphy of Polychrus + anoles.

Anoloid monophyly is supported (Stem 4) in
all networks and topologies by 14.1 (endolym-
phatic sacs penetrate nuchal musculature) and
61.2 (strongly bicapitate, bisulcate hemipenes).
Some notable features of anoloids are ambigu-
ously placed as apomorphies, either because of
the possibility of greater levels of universality, or
because of variability among anoloids: 7.1 (ex-
tensive skull rugosity—shared with chameleons
and morunasaurs), and 40.1 (midventrally con-
fluent postxiphisternal inscriptional ribs—
shared with chameleons, morunasaurs, Brachy-
lophus, and [in modified form] oplurines). In
Topologies 1 and 2, 52.1 (spinulate scale organs)
is considered an apomorphy for anoloids, but in
Topology 3, this is regarded as a synapomorphy
of anoloids + oplurines.

BASILISCINES

The topology of basiliscine relationships was
stable in all networks and trees, and agrees with
results presented by Etheridge and de Queiroz
(1988) and Lang (1989), i.e., Basiliscus is the
sister-taxon of Corytophanes + Laemanctus.
Apomorphies of the group include: 10.2 (Y-
shaped parietal roof with large median crest),
11.1 (parietal foramen in frontal [unknown in
Laemanctus}), and 34.2 (basiliscine-type caudal
vertebrae). In some topologies 63.1 (m. retractor
lateralis posterior substantially in hemipenial
sheath) falls on the ancestral stem, although it has
a greater level of universality in other topologies.
Basiliscus does not have apomorphies treated in
this analysis whose placement is independent of
network placement. In topologies where basilis-

IGUANIAN LIZARD PHYLOGENY 23

cines are considered the sister-taxon of acro-
donts, anoloids, or some combination, both su-
perciliary and subocular scales in Basiliscus must
return to the plesiomorphic condition (44.0 and
45.0), and in topologies where basiliscines are
considered to be the sister-taxon of acrodonts,
tail autotomy fracture planes (41.0) must be
regained in Basiliscus. In our opinion, these
hypothesized transformations reflect poorly on
the likelihood of a special relationship of acro-
donts and basiliscines, in light of the meager
evidence supporting this relationship.

Corytophanes is well-corroborated by 13.1
(elevation of osseous labyrinth) and 31.1 (clav-
icle insertion of scapula). In some out-group
topologies, 21.1 (shortened splenial) is also a
feature of this stem. Laemanctus was specified in
this analysis by development of extensive skull
rugosity (7.1) and by secondary enlargement of
the postfrontal (9.0), although placement of this
character is ambiguous because Basiliscus and
Corytophanes have reduced postfrontals (9.1).
Corytophanes + Laemanctus monophyly is sup-
ported by 8.1 (broadly juxtaposed squamosal and
jugal) and 10.3 (median parietal crest developed
embryonically).

CROTAPHYTINES

The crotaphytines are remarkably plesiomor-
phic in many respects and lack any descriptively
unique morphological features. Their mono-
phyly is demonstrable only against the back-
ground of their out-groups, which possibly are
either a group composed of sceloporines,
oplurines, and tropidurines (and possibly includ-
ing anoloids), or a group composed of acrodonts
and basiliscines. With respect to these, the crota-
phytines have arguably made three reversals:
27.0 (regain palatine teeth), 36.0 (regain poste-
rior coracoid fenestrae), and 38.0 (develop ribs
on the third cervical vertebra). More interest-
ingly, the S-condition nasal apparatus (54.1) in
the crotaphytines is only ambiguously consid-
ered homologous with that in the iguanines. Cro-
taphytus has only one unambiguous apomorphy
in this analysis (7.1—extensive skull rugosity in
older individuals) and Gambelia has none. Itmay
well be that Crotaphytus and Gambelia are preda-
tory relicts of a very old group.

IGUANINES

In no topology was there an unambiguous
dichotomous resolution of the three iguanine
taxa used in the analysis, although monophyly of
the group is highly corroborated, both results in
accordance with those of de Queiroz (1987) and
Etheridge and de Queiroz (1988). Characters that
support the monophyly of the iguanines in all
topologies are: 12.1 (supratemporal mostly on
the medial surface of the supratemporal process
of the parietal), 34.1 (iguanine caudal vertebrae),
and 65.1 (colic septa). Polycuspate teeth (25.1)
and 54.1 (S-condition nasal apparatus) in some
topologies have greater levels of universality.
Unambiguous apomorphies of the three terminal
taxa are: Dipsosaurus—11.1 (parietal foramen in
frontal), Brachylophus—none, and iguanas—
10.1 (V or Y-shaped parietal table). Our suspi-
cion, however, is that Dipsosaurus, with its sce-
loporine-like superciliaries and suboculars, as
well as unmodified postxiphisternal inscriptional
ribs, is the sister-taxon of Brachylophus + other
iguanines, which have broken-up suboculars,
undifferentiated superciliaries and modified in-
scriptional ribs.

MORUNASAURS

The morunasaurs were initially presumed to
be monophyletic (see “Choice of Terminal
Taxa”) on the basis of their very reduced vomers.
No other apomorphies were discovered whose
placement was independent of a particular net-
work placement. In Networks A-B, F-G, I-L the
morunasaurs were placed as the sister-taxon of
the iguanines and in this topology three apo-
morphies obtained: 6.1 (lacrimal foramen en-
larged), 7.1 (extensive skull rugosity), and 19.1
(anterior surangular foramen ventral to posterior
extremity of dentary). In Network C (moruna-
saurs as the sister-taxon of iguanines) and in
Network H (morunasaurs as the sister-taxon of
anoloids + iguanines) only two characters are
placed unambiguously on the morunasaur stem:
6.1 (enlarged lacrimal foramen) and 19.1 (ante-
rior surangular foramen position). In Network D,
the morunasaurs formed the sister-taxon of the
anoloids and four unambiguously placed apo-
morphies obtained: 6.1 (lacrimal foramen en-
larged), 18.0 (coronoid labial blade present), 19.1

24 MISCELLANEOUS PUBLICATIONS

(anterior surangular foramen position), and 25.1
(polycuspate marginal teeth). In Network E,
morunasaurs are regarded as the sister-taxon of
the acrodonts and the only unambiguously placed
characters on the morunasaur stem are: 18.0
(regain coronoid labial blade) and 25.1 (poly-
cuspate marginal teeth).

OPLURINES

In all topologies, the oplurines were supported
as monophyletic by: 12.1 (supratemporal sits
mostly on the medial surface of the supratem-
poral process of the parietal), 43.1 (interparietal
black spot), and 58.1 (V-condition of ulnar nerve
pathway). Other notable characteristics of
oplurines, 40.2 (postxiphisternal inscriptional
ribs forming paired splints) and 52.1 (spinulate
scale organs) are only arguably synapomorphic
or have greater levels of generality when
oplurines and anoloids are considered sister-taxa
(Fig. 7—Networks I-J); in other networks these
features are unambiguously placed as apomor-
phies of the oplurines. Oplurus is supported as
monophyletic by a reversal (39.0—four sternal
ribs) and 38.2 (first pair of cervical ribs on verte-
bra 5); Chalarodon by 9.1 (postfrontal lost), 30.1
(clavicular flange reduced or absent), and a re-
versal (46.0—regaining of median enlarged dor-
sal scale row).

SCELOPORINES

Three, equally parsimonious sceloporine to-
pologies were discovered that were independent
of network (Fig. 12). In all three, monophyly of
the sceloporines (Stem 1) was supported by: 28.1
(pterygoid teeth lost), 30.1 (clavicular flange
reduced), 33.1 (posterior process of interclavicle
invested by sternum anteriorly), 53.1 (sink-trap
nasal apparatus), 60.1 (enlarged posterior lobe of
hemipenis), and 62.1 (m. retractor lateralis pos-
terior completely divided). Also common to all
three topologies were: (1) Uta with no apomor-
phies; (2) Petrosaurus with two reversals, 38.0
(ribs on cervical vertebra 3) and 39.0 (4 sternal
ribs); (3) Phrynosoma and the sand lizards (Stem
2) supported as monophyletic by 5.1 (lacrimal
absent) and 9.1 (postfrontal absent).

Topology 1.—In the most common topology,
Stem 3 (Sceloporus, Urosaurus, Uta, and Petro-

Topology 1

Uta
Petrosaurus
Sceloporus
Urosaurus
Phrynosoma

B) Sand Lizards

Topology 2

Uta
Petrosaurus
Urosaurus
Sceloporus
Phrynosoma

Sand Lizards

Topology 3

Uta
Petrosaurus
Phrynosoma
Sand Lizards

Sceloporus

Urosaurus

Fig. 12. Alternative topologies discovered within the
sceloporines.

IGUANIAN LIZARD PHYLOGENY 25

saurus) was supported by a greatly enlarged in-
terparietal scale (42.1). This is ambiguous be-
cause if the small interparietal of Uma and
Phrynosoma is secondary (and the large inter-
parietal of Callisaurus and Holbrookia homolo-
gous), then this stem would be unsupported.
However, a character of ambiguous placement,
35.1 (loss of scapular fenestra; reversed in Sce-
loporus), might support this stem. Also, taxa
subtended by this stem have relatively well-
developed frontal scales that, if independent of
the interparietal scale development in this clade,
would lend support to either this arrangement or
Topology 2. Stem 4 (Sceloporus + Urosaurus) is
supported by 59.1 (B-condition of shank inner-
vation) and 66.1 (paired belly patches), the paired
belly patches in sand lizards apparently being
nonhomologous. However, the homologies of
66.1 are suspect, because “incipient” patches are
seen in Petrosaurus mearnsi and the axillary spot
of Uta may also be homologous. Sceloporus is
supported by 47.1 (loss of gular fold) and a
reversal, 35.0 (regaining scapular fenestra) and
Urosaurus was supported by no apomorphies in
this analysis. One character not included in this
analysis because of among-group characteriza-
tion problems, “hooked” clavicles (Etheridge,
1964) is congruent with this topology and would
serve to place Uta as the sister-taxon of Urosau-
rus + Sceloporus.

Topology 2.—Stems 1-3 areas in Topology 1.
Sceloporus is supported solely by 47.1 (loss of
gular fold), but Urosaurus is linked (Stem 5) with
Uta and Petrosaurus, rather than with Sce-
loporus, by 35.1 (loss of scapular fenestra). Uta
and Petrosaurus are linked (Stem 6) by a rever-
sal, 59.0 (A-condition of shank innervation).

Topology 3.—Stem 7 (Phrynosoma, sand
lizards, Petrosaurus, and Uta) is supported by a
reversal (59.0—A-condition of shank innerva-
tion) and Utaand Petrosaurus (Stem 8) are linked
by 35.1 (loss of scapular fenestra), a feature
shared convergently with Urosaurus in this to-
pology.

Each of the three topologies is at variance with
previously published cladograms (Presch, 1969;
Etheridge and de Queiroz, 1988). In these earlier
works, Petrosaurus was considered the sister-
taxon of the remaining sceloporines, whereas we

have no topologies in which this is the case. In
earlier studies, absence of a nasal valve and four
sternal ribs in Petrosaurus were considered ple-
siomorphic, but subsequently we concluded
(following Malan, 1946) that erectile tissue sur-
rounding the external naris is plesiomorphic for
squamates. Also, presumptive near-relatives
(e.g., “Tropidurus” west of the Andes) have nasal
valves. Therefore, the lack of a nasal valve in
Petrosaurus is only arguably plesiomorphic. The
4 sternal ribs of Petrosaurus are likely to be
apomorphic when compared with the 3-sternal-
ribbed tropidurines. Additionally, shimmy-bur-
ial, noted by Paull et al. (1976) and Etheridge and
de Queiroz (1988) as a possible synapomorphy
of the sceloporines, has been described in one
species of “Tropidurus” (Dixon and Wright,
1975) and may be found in at least one species of
“Agama” (Patterson, 1987). We suspect that this
behavior could be simply plesiomorphic within
Iguania.

TROPIDURINES

In this group, our results correspond reasona-
bly closely to the results of Etheridge and de
Queiroz (1988). In all networks (Fig. 7) the tro-
pidurines (Fig. 13—Stem 1) are supported by
two unambiguously placed features: 23.1 (re-
duced angular) and 47.1 (gular fold incomplete
medially). In Network J, these features are joined
by 22.1 (posterior extension of splenial) and in
Network K by 37.1 (enlarged, median sternal
fontanelle). Although this character list is not
impressive, because these characters occur inde-
pendently elsewhere in the tree, bear in mind that
this resolution is not particularly sensitive to out-
group placement. Even if the oplurines are ex-

Liolaemus group

Leiocephalus
"Stenocercus"
"Tropidurus"

Uranoscodon

Fig. 13. Tropidurine topology.

26 MISCELLANEOUS PUBLICATIONS

cluded from the analysis (possibly the first out-
group of the tropidurines), this resolution still
obtains. Stem 2 (Fig. 13—the Liolaemus group)
is supported in all topologies by a reversal, 24.0
(posterior mylohyoid foramen anterior to level of
apex of coronoid); and 35.1 (scapular fenestra
absent), 49.1 (preanal pores present), and 54.1
(S-condition nasal apparatus). Alternative to-
pologies exist in the Liolaemus group but this is
discussed following the other tropidurines.

Stem 3, the “northern tropidurines” of Eth-
eridge and de Queiroz (1988) are supported in all
topologies by: 20.1 (fusion of Meckel’s groove),
21.1 (splenial extends not more than 50% length
of tooth row), and a reversal to 59.0 (B-condition
of shank innervation). We regard the association
of Leiocephalus with the “Stenocercus” and
“Tropidurus” groups to be arguable. Leiocepha-
lus shares with the Liolaemus group the premax-
illary spine overlapped by the nasals (1.1—-vari-
able in Phymaturus) and an enlarged coronoid
labial blade (8.0), which in this “neighborhood”
of the cladogram is likely apomorphic. Leio-
cephalus is clearly monophyletic, supported by
the characters mentioned in “Choice of Terminal
Taxa” (10.1—shape of parietal roof; and 32.1—
presence of an anterior process of the interclav-
icle):

Stem 4, supporting the monophyly of the
“Stenocercus’ + “Tropidurus” groups carries
four unambiguously placed apomorphies: 24.2
(extreme posterior position of the posterior my-
lohyoid foramen), 36.0 (regaining of a posterior
coracoid fenestra), 55.1 (fusion of the nasal
concha to the roof of the nasal cavity [=reduction
of the supraconchal cavity]), and 61.1 (bisulcate
hemipenes). The “Stenocercus” group is weakly
corroborated by 23.0 (secondary enlargement of
the angular) and a feature noted in “Choice of
Terminal Taxa,” extensive hemipenial sheath
musculature. Because of the poor resolution
within the “Stenocercus” group, however, we
regard its monophyly as not well documented.

Stem 5, the “Tropidurus” group (“Tropidu-
rus” + Uranoscodon) is well-corroborated by
four features: 33.1 (posterior process of the inter-
clavicle invested by sternum far anteriorly), 42.1
(fused interparietal scales), 61.2 (strongly bi-
capitate hemipenes), and 64.1 (presence of a

hemipenial dorsal accessory sheath muscle). No
features analyzed support the monophyly of
“Tropidurus” (but see “Choice of Terminal
Taxa”), but the obviously highly apomorphic
Uranoscodon has lost distinctive superciliaries
and suboculars (44.1 and 45.1), regained a gular
fold (47.0), and developed the B-condition of
shank innervation (59.1).

TROPIDURINES: LIOLAEMUS GROUP

Within the tropidurines, the Liolaemus group
has two topologies that are independent of net-
work (Fig. 14). In both topologies, Phymaturus
carries unambiguously three reversals: 21.0
(splenial extends anteriorly more than 50%
length of tooth row), 38.0 (ribs on cervical verte-
bra 3), and 39.0 (sternal ribs 4 [although Lio-
laemus was coded “unknown” because it has 3 or
4]), plus 30.1 (clavicular flange reduced); and
Liolaemus has a reversal (36.0—posterior cora-
coid fenestra present). In Topology 1, Phyma-
turus is the sister-taxon of Ctenoblepharys +

Topology 1
Liolaemus

| Ctenoblepharys

Phymaturus

Topology 2
Liolaemus

Ctenoblepharys
9 Phymaturus

Fig. 14. Alternative topologies discovered within the
Liolaemus group.

IGUANIAN LIZARD PHYLOGENY 27

Liolaemus. Ctenoblepharys + Liolaemus (Stem
1) is supported by the supratemporal fitting in a
groove on the supratemporal process of the parie-
tal (12.2). In this topology Ctenoblepharys is
apomorphic in character 45.1 (subocular di-
vided), although Phymaturus was coded as
“unknown” for this feature because of internal
variability. In Topology 2, Liolaemus is the sis-
ter-taxon of Ctenoblepharys + Phymaturus, Stem
2, Ctenoblepharys + Phymaturus being sup-
ported by 45.1 (divided subocular). A feature
noted by Arnold (1984) as possibly synapomor-
phic for the Liolaemus group, m. retractor later-
alis posterior with a well-defined fleshy inser-
tion, is also more well-developed in Liolaemus
and Ctenoblepharys than in Phymaturus. Degree
of development of this muscle supports Topol-
ogy 1.

Although Topology 2 is analytically equal to
Topology 1, it is important to note that it takes
advantage of an “unknown” in Phymaturus (45.0
in Phymaturus patagonicus; 45.1 in P. palluma).
Because P. patagonicus is otherwise more ple-
siomorphic than P. palluma, and because of the
hemipenial musculature character of Arnold
(1984) mentioned above we support Topology 1
as the most likely.

SCELOPORINES + OPLURINES +
TROPIDURINES (+ ANOLOIDS)

In Networks A-H, and L (Fig. 7), sceloporines
form the sister-taxon of oplurines + tropidurines
(Fig. 15—Topology 1). In Network I, this ar-
rangement is augmented by anoloids being
placed as the sister-taxon of the oplurines (Fig.
15—Topology 2). In Network J, oplurines +
anoloids form the sister-taxon of sceloporines +
tropidurines (Fig. 15—Topology 3), and in Net-
work K, the topology that obtains is sceloporines
+ (anoloids + [oplurines + tropidurines]) (Fig.
15—Topology 4). The “fence lizard” habitus of
the sceloporines, tropidurines, and oplurines
cannot be denied, and we think that this similar-
ity is due to synapomorphy rather than homo-
plasy or plesiomorphy. The association of the
anoloids with this group is more problematical,
but worthy of serious consideration.

Topology 1.—Stem 1 (subtending the entire
group) is corroborated by three characters that

singly do not promote confidence against the
backdrop of variability in Iguania: 39.1 (three
sternal ribs [C.I.=ca. 0.16]) and 44.0 and 45.0,
which are both reversals to likely plesiomorphic
conditions of the superciliaries and enlarged
suboculars. A fourth character (37.1), an en-
larged sternal fontanelle, may support this clade,
but condition 37.0 in oplurines makes placement
of this feature ambiguous.

The monophyly of oplurines + tropidurines
(Stem 2) is weakly supported by the widely
homoplastic features 22.1 (splenial terminates
posteriorly at the anterior edge of the mandibular
fossa) and 48.1 (loss of femoral pores). Addition-
ally, 17.1 (strong posterior extension of the den-
tary) may support this clade, but condition 17.0
in the Liolaemus group makes the character
placement ambiguous.

Topology 2.—The subtending Stem 3 is
roughly equivalent to Stem 1 of Topology 1, but
with the addition of 21.1 (splenial extends anteri-
orly only 4 length of tooth row) to 39.1, 44.0, and
45.0 (discussed under “Topology 1”). Also in
Stem 1 of Topology 1, 37.1 (enlarged sternal
fontanelle) is placed ambiguously on this stem
because of the absence of an enlarged fontanelle
(37.0) in oplurines and anoloids. The monophyly
of tropidurines + (oplurines + anoloids) (Stem 4)
is supported by only one unambiguously placed
character, 48.1 (loss of femoral pores), although
20.1 (fused Meckel’s groove) and 22.1 (posterior
position of posterior mylohyoid foramen) can be
placed on this stem as one alternative.

The monophyly of anoloids + oplurines (Stem
5) is also supported by only one unambiguously
placed character, 52.1 (spinulate scale organs).
Assuming that the splint-like postxiphisternal
inscriptional ribs (40.2) and the mid-ventrally
continuous postxiphisternal inscriptional ribs
(40.1) of anoloids are homologous at a more
inclusive level than either is with 40.0 (nonelon-
gate postxiphisternal inscriptional ribs) would
add another unambiguous homologue on this
stem and would serve to make this, or Topology
3, the preferred topology of the relationships
between sceloporines, tropidurines, oplurines,
and anoloids.

Character 37.0 (loss of an enlarged sternal
fontanelle) is ambiguously considered a synapo-

28 MISCELLANEOUS PUBLICATIONS

Topology 1
Sceloporines
Tropidurines

2 Oplurines

Topology 3
Sceloporines
7 Tropidurines
6 Oplurines

8 Anoloids

Topology 2
Sceloporines
Tropidurines
Oplurines

5 Anoloids

Topology 4
Sceloporines
Anoloids
Oplurines

11 Tropidurines

Fig. 15. Alternative topologies of sceloporines + oplurines + tropidurines (+ anoloids).

morphy of anoloids + oplurines in this topology
because it depends on the enlarged median ster-
nal fontanelle of sceloporines and tropidurines
being a synapomorphy of the entire group (sce-
loporines, oplurines, tropidurines, and anoloids)
with a reversal, rather than the independent ac-
quisition of this feature in the sceloporine and
tropidurine clades.

Topology 3.—Stem 6 is supported unambigu-
ously by 21.1 (relatively short splenial), 39.1
(three sternal ribs), and 45.0 (subocular scale
enlarged). Characters 44.0 (superciliaries elon-
gate) and 48.1 (loss of femoral pores) are placed
on the stem, although this arrangement would
require sceloporines to develop femoral pores
convergently.

The evidence supporting the monophyly of
sceloporines + tropidurines (Stem 7) consists
solely of one unambiguously placed character,
37.1 (single, enlarged median sternal fontanelle),

although two other features can be placed on this
stem under different character optimization, 44.1
(distinctly elongate superciliaries) and 59.1 (in-
terosseus innervation of dorsal shank muscula-
ture).

Stem 8, subtending oplurines + anoloids, is
also supported by a single unambiguously placed
character, 52.1 (spinulate scale organs); al-
though, as discussed under Stem 5 of Topology 2,
assumption of homology between 40.1 (medially
confluent postxiphisternal inscriptional ribs) and
40.2 (splint-like postxiphisternal inscriptional
ribs) would add another character on this stem.
Two other characters of ambiguous placement
mightalso rest on this stem: 20.1 (fused Meckel’s
groove) and 48.1 (loss of femoral pores [which
would require convergent loss in tropidurines]).

Topology 4.—Stem 9 (subtending the entire
group) is corroborated by 21.1 (splenial extends
’ length of toothrow), 39.1 (three sternal ribs),

IGUANIAN LIZARD PHYLOGENY 29

and 45.0 (a reversal to having an enlarged sub-
ocular). Additionally, 44.0 (reversal to elongate
superciliaries) may belong here, although this
would require a change again to 44.1 in anoloids.
Stem 10 (anoloids + [oplurines + tropidurines])
is supported by 48.1 (loss of femoral pores),
although 20.1 (fused Meckel’s groove) and 52.1
(spinulate scale organs [requiring a loss in tro-
pidurines]) might be on this stem. Stem 11,
uniting oplurines and tropidurines, is also sup-
ported by only one unambiguously placed char-
acter, 22.1 (posteriorly extended splenial), al-
though this is convergent in anoles.

Although the evidence for a special relation-
ship among the tropidurines, sceloporines, and
oplurines is meager, alternatives, such as at-
tempting to ally the Madagascan oplurines with
the geographically proximate Afro-Australo-
Asian acrodonts are considerably less parsimoni-
ous.

SUMMARY OF RESULTS

Results of the analysis support the recognition
of the suprageneric groups of Etheridge (1959,
1964, 1966, 1967) and Etheridge and de Queiroz
(1988), as well as the acrodont group and its
constituent parts. Our results do not support the
hypothesized intragroup relationships of ano-
loids and sceloporines of Etheridge and de
Queiroz (1988), the unambiguously supported
monophyly of Agamidae* (Borsuk-BiaJynicka
and Moody, 1984), nor the metataxon status of
“Iguanidae” (because there is no evidence of
monophyly in the face of incongruent evidence
of paraphyly [Kluge, 1989]). No clear resolution
of intergroup relationships within iguanids was
obtained, although evidence of a relationship of
the Madagascan oplurines with the American
sceloporines and tropidurines (and possibly
anoloids) was presented.

Although Schwenk (1988) was the first to
present evidence that “Iguanidae” is paraphyletic
with respect to Agamidae* + Chamaeleonidae, it
appears from our analysis that his character evi-
dence, presence of lingual reticular papillae
(Char. 67.1), is homoplastic in anoles and acro-
donts. For this feature to be synapomorphic
would require either the paraphyly (or poly-
phyly) of the anoloids or the loss of the feature in
anoloids other than anoles.

The results of our analysis show that contin-
ued recognition of Agamidae* and “Iguanidae” is
not consistent with recovered historical relation-
ships. Rather than maintain the unsupported
collectives, Agamidae* and “Iguanidae,” we
propose to recognize as families, sedis mutabilis
(Wiley, 1979, 1981a), the largest historical
groups that are consistent with the strict consen-
sus tree generated in the phylogenetic analysis of
Iguania (Fig. 8). The taxonomy we have adopted
(followed in parentheses by former taxonomic or
informal equivalents) is listed below and also is
illustrated in tree form in Figure 16.

Iguania Cope, 1864: incertae sedis: +Aciprion*
Cope, 1873; ?+Arretosaurus* Gilmore,
1943; +Carduciguana* Augé, 1987;
+Cypressaurus* Holman, 1972; ?+Ericho-
saurus* Ameghino, 1899; ?+Geiseltaliel-
lus* Kuhn, 1944; +Harrisonsaurus Hol-
man, 1981; +Paradipsosaurus Fries, Hib-
bard, and Dunkle, 1955; +Parasauromalus
Gilmore, 1928; ?+Pleurodontagama* Bor-
suk-BiaJynicka and Moody, 1984; +Prist-
iguana* Estes and Price, 1973; ?+Swain-

, Chamaeleoninae
Chamaeleonidae

Leiolepidinae
Agaminae
Corytophanidae
Crotaphytidae
nasa Hoplocercidae
Iguanidae
Opluridae
Phrynosomatidae
Polychridae
Liolaeminae
Leiocephalinae

Tropiduridae Tropidurinae

Fig. 16. Taxonomic tree.

30 MISCELLANEOUS PUBLICATIONS

iguanoides* Sullivan, 1982.
Chamaeleonidae Rafinesque, 1815
(Acrodonta): incertae sedis: +Mimeo-
saurus* Gilmore, 1943; +Priscagama*
Borsuk-Bialynicka and Moody, 1984;
+Tinosaurus* Marsh, 1872.

Agaminae Spix, 1825 (Agamidae*: Agam-
inae)

Chamaeleoninae Rafinesque, 1815 (Cha-
maeleonidae)

Leiolepidinae Fitzinger, 1843 (Agami-
dae*: Uromastycinae; or Uromastyc-
idae)

Corytophanidae Fitzinger, 1843 (“Igua-
nidae”: basiliscines)

Crotaphytidae Smith and Brodie, 1982
(“Iguanidae”: crotaphytines)

Hoplocercidae new family (“Iguanidae”:

morunasaurs)

Iguanidae Oppel, 1811 (“Iguanidae”: igua-
nines)

Opluridae Moody, 1983 (“Iguanidae”:
oplurines)

Phrynosomatidae Fitzinger, 1843 (“Igua-
nidae”: sceloporines)
Polychridae Fitzinger, 1843 (“Iguanidae”:
anoloids)
Tropiduridae Bell, 1843 (“Iguanidae”: tro-
pidurines)
Leiocephalinae new subfamily
Liolaeminae new subfamily
Tropidurinae Bell, 1843

The salient differences between this taxon-
omy and the traditional one (e.g., Camp, 1923;
Estes et al., 1988) are: (1) Agamidae* and Cha-
maeleonidae have been combined and (2) the
informal groups within “Iguanidae” are accorded
formal, independent taxonomic status. With the
iguanid taxa this was unavoidable because these
are the largest groups whose historical reality is
well supported. Placing Agamidae* and Cha-
maeleonidae in one family was done because in
this case we have considerable evidence that this
is a single monophyletic group in a consensus
polytomy with the former iguanid groups. We
could have retained the name Acrodonta for this
group, with three families within it (Chamaele-

onidae, Leiolepididae [=Uromastycidae], and
Agamidae). In doing so, however, we would not
have been consistent in recognizing the largest
monophyletic group of acrodonts as a family in
symmetry with the other iguanian groups.

At this time we have chosen to recognize
formal subfamilies only in the new, enlarged
Chamaeleonidae (i.e., the former Acrodonta) and
within Tropiduridae. We have taken this step in
Chamaeleonidae because it is clear that not rec-
ognizing subfamilies would result in consider-
able confusion and because of the otherwise
long-standing nomenclatural stability of its con-
stituent groups. Within Tropiduridae we recog-
nize subfamilies in order to simplify our other
projects ongoing within these groups. Although
de Queiroz (1987) proposed a suprageneric phy-
logenetic taxonomy within his Iguaninae (our
Iguanidae), it was presented within a different
philosophical context, so, rather than enter a
philosophical discussion that is outside the scope
of this paper we, without prejudice, do not ad-
dress the subfamilial taxonomy of the iguanas.

In defense of our particular choice of rank-
ings, we could have enlarged Iguanidae to in-
clude the acrodont and iguanid groups as sub-
families (i.e., made Iguanidae equivalent to Igua-
nia). This would have rendered an enlarged Igua-
nidae coextensive (=redundant) with Iguania.
However, by using the Linnaean family-group
category as we have, we advertise the lack of
intergroup resolution, while avoiding nomencla-
tural redundancy. We recognize, of course, that
ranking is arbitrary and we see “families” not as
members of some natural class of comparable
entities, but as merely nomenclaturally internally
consistent “files” in the Linnaean book-keeping
system used throughout biology.

We realize that the new taxonomy and nomen-
clature will not be popular with those preferring
a Classification rooted in social tradition or in
some arbitrary measure of overall similarity.
And, if Agamidae were an older name than
Chamaeleonidae, we doubt that there would be
much controversy regarding that nomenclatural
change, because systematics as practiced by the
majority of workers has little to do with evolution
and much to do with a crude sort of essentialism.

IGUANIAN LIZARD PHYLOGENY 31

TAXONOMIC ACCOUNTS AND CHARACTERIZATIONS

The characterizations provided for the fami-
lies and subfamilies are not lists of apomorphies;
those data are available in Appendices 2 and 3
and in “Results.” The characterizations allow the
taxa to be differentiated from the other taxa of
equal rank in this section. Metataxon and quota-
tion conventions are suspended in synonymies
for nomenclatural clarity.

IGuANIA CopE, 1864

1864. Iguania Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:226.

Characterization.—(1) frontals fused em-
bryonically (JJollie, 1960; Estes et al., 1988); (2)
frontals constricted between the orbits (reversed
in some groups) (Estes et al., 1988); (3) broad
frontal shelf underlying nasals (Estes et al.,
1988); (4) postfrontal reduced (Estes et al., 1988;
Presch, 1988); (5) dracomorph brain-stem mor-
phology (Northcutt, 1978); (6) m. intercostalis
ventralis absent (Camp, 1923); (7) tongue mu-
cocytes mostly serous and sero-mucous (Gabe
and Saint Girons, 1969; Schwenk, 1988).

Content.—Those taxa that together form the
sister-taxon of Scleroglossa (=Scincogekkono-
morpha); traditional “Iguanidae,” Agamidae*,
and Chamaeleonidae; here recognized as Cha-
maeleonidae (including former Uromastycidae
and Agamidae*), Corytophanidae, Crotaphyti-
dae, Hoplocercidae, Iguanidae, Opluridae,
Phrynosomatidae, Polychridae, and Tropiduri-
dae.

Distribution.—All continental temperate and
tropical regions. Absent from most of Oceania.

Comment.—tThe attribution of the name
Iguania to Cuvier (1817) is in error; Cuvier used
the explicit (though non-Latinized) family-group
name Iguaniens. The first author to use the name
Iguania was Cope (1864), the same author that
coined the name Acrodonta for a group composed
of the former Agamidae* and Chamaeleonidae
(in the old sense), now equivalent to Chamaele-
onidae.

CHAMAELEONIDAE RAFINESQUE, 1815

1815. Camelonia Rafinesque, Analyse
Nat.:75. Type genus: “Camaeleo
Daud.” (=Chamaeleo Daudin, 1802
=Chamaeleo Laurenti, 1768).

1825. Agamae Spix, Anim. Nov. Spec. Nova
Lacert.:12. Type genus: Agama Daudin,
1802.

1825. Camelionidae Gray, Ann. Philos.,
(2)10:200. Type genus: “Chamelion,
Lin.” (=Chamelion Gray, 1825, a likely
incorrect subsequent usage of Chamae-
leon Gronovius, 1763, a rejected name
[Opinion 89]).

1825. Stellionidae Bell, Zool. J., London,
2:457. Type genus: “Stellio Daudin”
(not Stellio Laurenti, 1768). See
Stejneger in Smith (1932) and Smith
(1957) for discussion.

1826. Draconoidea Fitzinger, Neue Classif.
Rept.:11. Type genus: “Draco Kaup”
(=Draco Linnaeus, 1758).

1843. Gonyocephali Fitzinger, Syst. Rept.,
1:15. Type genus: “Gonyocephalus
Kaup (Cuv.)” (=Gonocephalus Kaup,
1825).

1843. Calotae Fitzinger, Syst. Rept., 1:15.
Type genus: “Calotes Kaup” (=Calotes
Cuvier, 1817).

1843. Semiophori Fitzinger, Syst. Rept., 1:15.
Type genus: “Semiophorus Wagl[er].”
(=Sitana Cuvier, 1829).

1843. Otocryptae Fitzinger, Syst. Rept., 1:15.
Type genus: “Otocryptis Wiegm[ann].”
(=Otocryptis Wagler, 1830).

1843. Lophurae Fitzinger, Syst. Rept., 1:15.
Type genus: “Lophura Wagll[er].
(Gray)” (=Lophura Gray, 1827 =Hydro-
saurus Kaup, 1828).

1843. Trapeli Fitzinger, Syst. Rept., 1:17.
Type genus: Trapelus Cuvier, 1817.

1843. Phrynocephali Fitzinger, Syst. Rept.,
1:18. Type genus: Phrynocephalus
Kaup, 1825.

32 MISCELLANEOUS PUBLICATIONS

1923. Brookesinae Nopsca, Fortschr. Geol.
Palaeont., 2:124. Type genus: Brooke-
sia Gray, 1865.

1984. Priscagaminae Borsuk-BiaJynicka and
Moody, Acta Palaeontol. Polon., 29:54.
Type genus: Priscagama Borsuk-Bi-
alynicka and Moody, 1984. See com-
ment.

Characterization.—(1) maxillae meet
broadly anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen variably en-
larged; (3) skull roof variably rugose (only cha-
meleons and +Priscagama*); (4) jugal and squa-
mosal broadly juxtaposed (not +Priscagama*);
(5) parietal roof shape quadrangular (or domed in
chameleons); (6) parietal foramen usually ab-
sent, if present, on frontoparietal suture; (7)
supratemporal on lateral side of supratemporal
process of parietal, except in chameleons in
which its reduced to a small splint on the medi-
ocaudal edge of the ventral ramus of the squa-
mosal and has lost entirely any connection of the
parietal (Rieppel, 1981); (8) nuchal endolympha-
tic sacs penetrating nuchal musculature only in
some Brookesia (Chamaeleoninae); (9) dentary
expanded onto labial face of coronoid (except
+Priscagama*); (10) no labial blade of coronoid;
(11) anterior surangular foramen inferior to pos-
teriormost extent of dentary; (12) Meckel’s
groove broadly open; (13) splenial short anteri-
orly, or absent; (14) dentary and maxillary teeth
acrodont, fused to underlying bone in adults; (15)
palatine teeth absent (present in +Priscagama*);
(16) pterygoid teeth absent (present in +Prisca-
gama*); (17) posterior process of interclavicle
not invested by sternum far anteriorly; (18) no
caudal autotomy (except some Uromastyx),
caudal vertebrae with single transverse processes
anteriorly; (19) posterior coracoid fenestra
absent, except in Uromastyx; (20) sternal fonta-
nelles present or absent; (21) number of sternal
ribs variable; (22) postxiphisternal inscriptional
ribs variable; (23) interparietal scale not en-
larged; (24) mid-dorsal scale row variable; (25)
gular fold complete medially, except chame-
leons; (26) femoral pores present or absent; (27)
spinulate scale organs absent (except in some
chameleons; the spiked scale organs of some
agamines are Clearly not homologous); (28) acro-
dontan nasal apparatus (except Physignathus),

vestibule long, concha reduced or absent; (29)
hemipenes variable, none known to be bicapitate
or bisulcate; (30) colic septa absent (except in
Uromastyx and Hydrosaurus).

Content.—Agaminae Spix, 1825; Chamaele-
oninae Rafinesque, 1815; and Leiolepidinae
Fitzinger, 1843.

Distribution.—Tropical and temperate re-
gions of Africa, Madagascar, southern Europe,
Asia, and Australia (Fig. 17).

Comment.—As here used, Chamaeleonidae
is equivalent to Acrodonta of Estes et al. (1988).
Former Agamidae* and Chamaeleonidae are
synonymized because the monophyly of Agami-
dae* is only ambiguously supported (Camp,
1923; Estes et al., 1988; contra Borsuk-BiaJyn-
icka and Moody, 1984), even though the tradi-
tional Chamaeleonidae (the chameleons) is well
supported. The constituent taxa of +Priscagami-
nae* Borsuk-BiaJynicka and Moody (1984) are
relegated to the status of incertae sedis within the
Chamaeleonidae because they are not tied to-
gether unambiguously by apomorphies, although
they are clearly plesiomorphic with respect to
any of the named suprageneric taxa within the
Chamaeleonidae.

AGAMINAE SPIX, 1825

1825. Agamae Spix, Anim. Nov. Spec. Nova
Lacert.:12. Type genus: Agama Daudin,
1802.

1825. Stellionidae Bell, Zool. J., London,
2:457. Type genus: “Stellio Daudin”
(not Stellio Laurenti, 1768). See
Stejneger in Smith (1932) for discus-
sion of unavailability of Stellio for any
member of Iguania.

1826. Draconoidea Fitzinger, Neue Classif.
Rept.:11. Type genus: “Draco Kaup”
(=Draco Linnaeus, 1758).

1843. Gonyocephali Fitzinger, Syst. Rept.,
1:15. Type genus: “Gonyocephalus
Kaup (Cuv.)” (=Gonocephalus Kaup,
1825).

1843. Calotae Fitzinger, Syst. Rept., 1:15.
Type genus: “Calotes Kaup” (=Calotes
Cuvier, 1817).

1843. Semiophori Fitzinger, Syst. Rept., 1:15.
Type genus: “Semiophorus Wagll[er].”
(=Sitana Cuvier, 1829).

IGUANIAN LIZARD PHYLOGENY 33

Agaminae

eed Hi
STPTT ETT Oy

os
s i
:
rt)
RANT é,
ss Seca Rx
RIES art z
' SSB, PETE x
QAO TOTTI ITTIOTIO

s)
OI Leiolepidinae

Kilometers

ES Chamaeleoninae

Fig. 17. Distribution of Chamaeleonidae, including subfamilies.

1843. Otocryptae Fitzinger, Syst. Rept., 1:15.
Type genus: “Otocryptis Wiegm[ann].”
(=Otocryptis Wagler, 1830).

1843. Lophurae Fitzinger, Syst. Rept., 1:15.
Type genus: “Lophura Wagl[er].
(Gray)” (=Lophura Gray, 1827 =Hydro-
saurus Kaup, 1828).

1843. Trapeli Fitzinger, Syst. Rept., 1:17.
Type genus: Trapelus Cuvier, 1817.

1843. Phrynocephali Fitzinger, Syst. Rept.,
1:18. Type genus: Phrynocephalus
Kaup, 1825.

Characterization.—(1) vomer flat or con-
vex; (2) lacrimal foramen extremely enlarged;
(3) skull roof not rugose or domed; (4) epiotic
foramen present (except in Moloch); (5) inter-
clavicle present; (6) paired, enlarged sternal
fontanelles; (7) postxiphisternal inscriptional
ribs short; (8) femoral pores present plesiomor-
phically; (9) normal feet.

Content®.—Acanthosaura
“Agama” Daudin, 1802;

Grays 1831;
“Amphibolurus”

Wagler, 1830; Aphaniotis Peters, 1864;
Caimanops Storr, 1974; “Calotes” Cuvier, 1817;
Ceratophora Gray, 1834; Chelosania Gray,
1845; Clamydosaurus’ Gray, 1825; Cophotis
Peters, 1861; Cryptagama Witten, 1984; Den-
dragama Doria, 1888; Diporiphora* Gray, 1842;
Draco Linnaeus, 1758; “Gonocephalus” Kaup,
1825; Harpesaurus Boulenger, 1885; Hydrosau-
rus Kaup, 1828; Hylagama Mertens, 1924; Japa-
lura Gray, 1853; Lophocalotes Giinther, 1872;
Lophognathus Gray, 1842; Lyriocephalus Mer-
rem, 1820; Mictopholis Smith, 1935; Moloch
Gray, 1841; Oriocalotes Giinther, 1864; Oto-
cryptis Wagler, 1830; Phoxophrys Hubrecht,

6 Until the controversy surrounding the
nomenclatural validity of names proposed by Wells
and Wellington (1983) is resolved, we refrain from
using their names.

7 Usually unjustifiably emended to Chlamy-
dosaurus.

34 MISCELLANEOUS PUBLICATIONS

1881; Phrynocephalus Kaup, 1825; Physi-
gnathus Cuvier, 1829; Psammophilus Fitzinger,
1843; Ptyctolaemus Peters, 1864; Salea Gray,
1845; Sitana Cuvier, 1829; “Tympanocryptis”
Peters, 1863; Xenagama Boulenger, 1895.

Distribution.—Temperate and tropical Eura-
sia, Africa, and Australia, including associated
islands (Fig. 17).

Comment.—As here used, Agaminae is
equivalent to Agamidae of previous authors, but
excluding Uromastyx and Leiolepis.

CHAMAELEONINAE RAFINESQUE, 1815

1815.Camelonia Rafinesque, Analyse
Nat.:75. Type genus: “Camaeleo
Daud.” (=Chamaeleo Daudin, 1802
=Chamaeleo Laurenti, 1768).

1825. Camelionidae Gray, Ann. Philos.,
(2)10:200. Type genus: “Chamelion,
Lin.” (=Chamelion Gray, 1825, a likely
incorrect subsequent usage of Chamae-
leon Gronovius, 1763, a rejected name
(Opinion 89).

1923. Brookesinae Nopsca, Fortschr. Geol.
Palaeont., 2:124. Type genus: Brooke-
sia Gray, 1865. See comment.

Characterization.—(1) vomer flat or con-
vex; (2) lacrimal foramen extremely enlarged;
(3) skull roof domed and rugose; (4) epiotic
foramen absent; (5) interclavicle absent; (6) no
sternal fontanelles; (7) postxiphisternal inscrip-
tional ribs elongate, fused medially; (8) femoral
pores absent; (9) zygodactyl feet.

Content.—Brookesia Gray, 1865; Calumma*
Gray, 1864; Chamaeleo Laurenti, 1768;
Bradypodion Fitzinger, 1843; Furcifer Fitzinger,
1843; Rhampholeon Giinther, 1874.

Distribution.—Extreme southwestern Eu-
rope, Africa (excluding the Sahara), southwest-
ern and northwestern Arabia, Madagascar, Sey-
chelles, India, and Sri Lanka, and associated
islands (Fig. 17).

Comment.—Our purpose here is not to evalu-
ate previous work on the phylogeny of chame-
leons, and although we have Brookesiinae Nop-
sca (Klaver and Bohme, 1986) in the synonymy
of Chamaeleonidae, this is only in recognition of
Brookesiinae as a family-group name. Because
our Chamaeleoninae is the equivalent of Cha-

maeleonidae of previous authors, we simply
regard Brookesiinae and Chamaeleoninae of
Klaver and BOhme (1986) to be tribes, Brooke-
siini (containing Brookesia and Rhampholeon)
and Chamaeleonini (containing Calumma*,
Furcifer, Bradypodion, and Chamaeleo).

LEIOLEPIDINAE FITZINGER, 1843

1843. Leiolepides Fitzinger, Syst. Rept., 1:18.
Type genus: Leiolepis Cuvier, 1829.

1868. Uromasticidae Theobald, J. Linn. Soc.
Zool., 10:34. Type genus: Uromastix
Merrem, 1820 (=Uromastyx Merrem,
1820).

Characterization.—(1) vomer concave; (2)
lacrimal foramen not enlarged; (3) skull roof not
rugose or domed; (4) epiotic foramen absent; (5)
interclavicle present; (6) paired, enlarged sternal
fontanelles; (7) postxiphisternal inscriptional
ribs short; (8) femoral pores present; (9) normal
feet.

Content.—Leiolepis Cuvier, 1829; Uro-
mastyx Merrem, 1820.

Distribution.—Deserts of North and East
Africa and Arabia to Iran, Afghanistan, Pakistan,
and western India; southern India and southern
China through Indochina to Sumatra (Fig. 17).

Comment.—Although this taxon has been
recognized previously (e.g., Borsuk-Bialynicka
and Moody, 1984), the name of priority is
Leiolepidinae rather than Uromastycinae.

CORYTOPHANIDAE FITZINGER, 1843

1843. Corythophanae Fitzinger, Syst. Rept.,
1:16. Type genus: “Corythophanes
Boie” (=Corytophanes Boie, 1827).
1900. Basiliscinae Cope, Annu. Rept. U.S.
Natl. Mus. for 1899:223. Type genus:
Basiliscus Laurenti, 1768.
Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof not strongly rugose (except in
Laemanctus); (4) jugal and squamosal broadly
juxtaposed in Corytophanes and Laemanctus; (5)
parietal roof Y-shaped with median crest formed
postembryonically in Basiliscus, embryonically
in Laemanctus and Corytophanes; (6) parietal

IGUANIAN LIZARD PHYLOGENY 35

foramen in frontal (parietal foramen absent in
Laemanctus); (7) supratemporal sits on lateral
side of supratemporal process of parietal; (8)
nuchal endolymphatic sacs not penetrating
nuchal musculature; (9) dentary not expanded
onto labial face of coronoid; (10) no labial blade
of coronoid; (11) anterior surangular foramen
superior to posteriormost extent of dentary; (12)
Meckel’s groove fused (except in some Basilis-
cus); (13) splenial relatively short (Coryto-
phanes) or long (Basiliscus and Laemanctus)
anteriorly; (14) dentary and maxillary teeth pleu-
rodont, not fused to underlying bone in adults;
(15) palatine teeth absent; (16) pterygoid teeth
present; (17) posterior process of interclavicle
not invested by sternum far anteriorly; (18) cau-
dal autotomy fracture planes present (except
Laemanctus), with transverse processes weak or
absent; (19) posterior coracoid fenestra absent;

1000
(Le eee)
Kilometers

Se :

Fig. 18. Distribution of Corytophanidae.

(20) sternal fontanelles very small or absent; (21)
sternal ribs 4; (22) postxiphisternal inscriptional
ribs short; (23) interparietal scale not enlarged;
(24) median dorsal scale row enlarged; (25) gular
fold complete medially; (26) femoral pores ab-
sent; (27) spinulate scale organs absent; (28)
nasal apparatus primitive, nasal vestibule short,
simple; concha present, free; (29) hemipenes
unicapitate, unisulcate; (30) colic septa absent.

Content.—Basiliscus Laurenti, 1768; Co-
rytophanes Boie, 1827; Laemanctus Wiegmann,
1834.

Distribution.—Western and eastern Mexico,
southward through Central America, to Ecuador
and Venezuela (Fig. 18).

Comment.—tThis family corresponds to the
“basiliscines” of Etheridge in Paull et al. (1976),
Etheridge and de Queiroz (1988), and Lang
(1989).

36 MISCELLANEOUS PUBLICATIONS

CROTAPHYTIDAE SMITH AND BRODIE, 1982

1982. Crotaphytinae Smith and Brodie, Guide
Field Ident. Reptiles N. Am.:106. Type
genus: Crotaphytus Holbrook, 1842.

Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof not strongly rugose, except in old
Crotaphytus; (4) jugal and squamosal not broadly
juxtaposed; (5) parietal roof trapezoidal; (6)
parietal foramen in frontoparietal suture; (7)
supratemporal sits on lateral side of supratem-
poral process of parietal; (8) nuchal endolymph-
atic sacs do not penetrate nuchal musculature; (9)
dentary not expanded onto labial face of coro-
noid; (10) labial blade of coronoid poorly devel-
oped or absent; (11) anterior surangular foramen
above posteriormost extent of dentary; (12)
Meckel’s groove not fused; (13) splenial rela-
tively long anteriorly; (14) dentary and maxillary
teeth pleurodont, not fused to underlying bone in
adults; (15) palatine teeth present; (16) pterygoid
teeth present; (17) posterior process of interclav-
icle not invested by sternum far anteriorly; (18)
caudal autotomy fracture planes present (except
in Crotaphytus), with transverse processes ante-
rior to fracture planes; (19) posterior coracoid
fenestra present; (20) sternal fontanelles very
small or absent; (21) sternal ribs 4; (22) postxi-
phisternal inscriptional ribs short; (23) interpari-
etal scale not enlarged; (24) mid-dorsal scale row
absent; (25) gular fold complete medially; (26)
femoral pores present; (27) spinulate scale
organs absent; (28) S-condition nasal apparatus;
nasal vestibule long, S-shaped, concha present;
(29) hemipenes unicapitate, unisulcate; (30)
colic septa absent.

Content.—Crotaphytus Holbrook, 1842;
Gambelia Baird and Girard, 1859.

Distribution.—Southwestern North America
from eastern Oregon to the Mississippi River and
south to northern Mexico (Fig. 19).

Comment.—Crotaphytidae corresponds to
the “crotaphytines” of Etheridge and de Queiroz
(1988).

HOPLOCERCIDAE NEW FAMILY

Type genus.—Hoplocercus Fitzinger, 1843.

Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen enlarged; (3)
skull roof strongly rugose (except in Hoplocer-
cus and “Morunasaurus”); (4) jugal and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal; (6) parietal foramen in fronto-
parietal suture (absent in some “Morunasau-
rus’); (7) supratemporal sitting on lateral side of
supratemporal process of parietal; (8) nuchal
endolymphatic sacs not penetrating nuchal mus-
culature; (9) dentary not expanded onto labial
face of coronoid; (10) labial blade of coronoid
large; (11) anterior surangular foramen inferior
to posteriormost extent of dentary; (12) Meckel’s
groove not fused; (13) splenial very large, pene-
trating far anteriorly; (14) dentary and maxillary
teeth pleurodont, not fused to underlying bone in
adults; (15) palatine teeth absent; (16) pterygoid
teeth present; (17) posterior process of interclav-
icle not invested by sternum far anteriorly; (18)
caudal autotomy fracture planes present (except
Hoplocercus), with transverse processes anterior
to fracture planes; (19) posterior coracoid
fenestra absent (except in “Morunasaurus” an-
nularis); (20) sternal fontanelles very small or
absent; (21) sternal ribs number 4; (22)
postxiphisternal inscriptional ribs long, conflu-
ent medially; (23) interparietal scale not en-
larged; (24) mid-dorsal scale row present (except
in “Morunasaurus” and Hoplocercus); (25) gular
fold complete medially; (26) femoral pores pres-
ent; (27) spinulate scale organs absent; (28)
primitive nasal apparatus; nasal vestibule short,
straight; concha present, free; (29) hemipenes
unicapitate, unisulcate; (30) colic septa absent.

Content.—‘‘Enyalioides” Boulenger, 1885;
Hoplocercus Fitzinger, 1843; “Morunasaurus”
Dunn, 1933.

Distribution.—Eastern Panama to the Pacific
lowlands of Ecuador; Upper Amazonian Basin of
Brazil, Colombia, Ecuador, and Peru; southeast-
em Brazil (Fig. 20).

IGUANIAN LIZARD PHYLOGENY 37

O 1000

Kilometers

Fig. 19. Distribution of Crotaphytidae.

Comment.—Hoplocercidae corresponds to
the “hoplocercines” of Smith et al. (1973),
“morunasaurines” of Estes and Price (1973), and
the “morunasaurs” of Etheridge and de Queiroz
(1988). We have employed Hoplocercus as the
type genus, rather than “Morunasaurus,”
because Hoplocercus is the oldest generic name
in the clade and the only name that does not refer
currently to a paraphyletic grouping (Etheridge
and de Queiroz, 1988).

IGUANIDAE OPPEL, 1811

1811. Iguanoides Oppel, Ordn. Fam. Gatt.
Rept.:26. Type genus: “Jguana Linné”
(=/guana Laurenti, 1768).

1843. Hypsilophi Fitzinger, Syst. Rept., 1:16.
Type genus: Hypsilophus Wagler, 1830
(=Iguana Laurenti, 1768).

1987. Amblyrhynchina de Queiroz, Univ.
California Publ. Zool., 118:160. Type

38

MISCELLANEOUS PUBLICATIONS

O etexe)

Kilometers

Fig. 20. Distribution of Hoplocercidae

i

"Hn

Il

IGUANIAN LIZARD PHYLOGENY 39

genus: Amblyrhynchus Bell, 1825. See
comment.

Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof not strongly rugose (except Am-
blyrhynchus); (4) jugal and squamosal not
broadly juxtaposed; (5) parietal roof variable; (6)
parietal foramen in frontoparietal suture (in fron-
tal in Dipsosaurus); (7) supratemporal sits on
medial side of supratemporal process of parietal;
(8) nuchal endolymphatic sacs not penetrating
nuchal musculature; (9) dentary not expanded
onto labial face of coronoid; (10) labial blade of
coronoid large; (11) anterior surangular foramen
superior to posteriormost extent of dentary; (12)
Meckel’s groove fused; (13) splenial relatively
short anteriorly; (14) dentary and maxillary teeth
pleurodont, not fused to underlying bone in
adults; (15) palatine teeth absent; (16) pterygoid
teeth present; (17) posterior process of interclav-
icle not invested by sternum far anteriorly; (18)
caudal autotomy fracture planes present (except
in Amblyrhynchus, Conolophus, Brachylophus,
and Jguana delicatissima), with transverse proc-
esses anterior and posterior to fracture planes
(when present) of anterior autotomic vertebrae;
(19) posterior coracoid fenestra present; (20)
sternal fontanelles very small or absent; (21)
sternal ribs 4; (22) postxiphisternal inscriptional
ribs variable (long and confluent medially in
some); (23) interparietal scale not enlarged; (24)
mid-dorsal scale row present (absent in Sau-
romalus and some Ctenosaura); (25) gular fold
complete medially; (26) femoral pores present;
(27) spinulate scale organs absent; (28) S-condi-
tion nasal apparatus; nasal vestibule long, S-
shaped; concha present (29) hemipenes unicapi-
tate, unisulcate; (30) colic septa present.

Content.—Amblyrhynchus Bell, 1825; Bra-
chylophus Cuvier, 1829; Conolophus Fitzinger,
1843; Ctenosaura Wiegmann, 1828; Cyclura
Harlan, 1824; Dipsosaurus Hallowell, 1854;
Iguana Laurenti, 1768; Sauromalus Duméril,
1856.

Distribution.—Tropical and subtropical
America from the southwestern United States
and eastern Mexico south to southern Brazil and

Paraguay; Galapagos Islands; Antilles; Fiji and
Tonga Islands (Fig. 21).

Comment.—lIguanidae corresponds to the
“iguanines” of Etheridge (1964) and Etheridge
and de Queiroz (1988) and Iguaninae of de
Queiroz (1987). For a formal infrafamilial taxon-
omy see de Queiroz (1987) (See discussion in
-IRESUIES: ):

OPLURIDAE Moopy, 1983

1843. Doryphori Fitzinger, Syst. Rept., 1:17.
Type genus: “Doryphorus (Cuv.).” See
comment.

1983a.Opluridae Moody, Adv. Herpetol.
Evol. Biol.:202. Type genus: Oplurus
Cuvier, 1829.

Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof strongly rugose; (4) jugal and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal; (6) parietal foramen in fronto-
parietal suture; (7) supratemporal sits on medial
side of supratemporal process of parietal; (8)
nuchal endolymphatic sacs not penetrating nu-
chal musculature; (9) dentary not expanded onto
labial face of coronoid; (10) labial blade of cor-
onoid poorly developed or absent; (11) anterior
surangular foramen above posteriormost extent
of dentary; (12) Meckel’s groove variably fused
or not; (13) splenial relatively short anteriorly;
(14) dentary and maxillary teeth pleurodont, not
fused to underlying bone in adults; (15) palatine
teeth present in some Oplurus, otherwise absent;
(16) pterygoid teeth present; (17) posterior pro-
cess of interclavicle not invested by sternum far
anteriorly; (18) caudal autotomy fracture planes
present, with transverse processes anterior to
fracture planes; (19) posterior coracoid fenestra
absent; (20) sternal fontanelles very small or
absent; (21) sternal ribs 3 or 4; (22) postxiphister-
nal inscriptional ribs appear in the form of paired
splints, isolated from the dorsal ribs and not
confluent medially; (23) interparietal scale not
enlarged; (24) mid-dorsal scale row absent
(Oplurus) or present, enlarged (Chalarodon);
(25) gular fold complete medially; (26) femoral

40 MISCELLANEOUS PUBLICATIONS

a

\ : sy! VAN
Re,
\\ y See.
Va \:

3000

Kilometers

Fig. 21. Distribution of Iguanidae, excluding Fijian and Tongan regions.

IGUANIAN LIZARD PHYLOGENY 41

pores absent; (27) spinulate scale organs present;
(28) primitive nasal apparatus; nasal vestibule
relatively short, straight; concha present, free;
(29) hemipenes unicapitate, unisulcate; (30)
colic septa absent.

Content.—Chalarodon Duméril and Bibron,
1837; Oplurus Cuvier, 1829.

Distribution.—Western and Central Mada-
gascar; Comoro Islands.

Comment.—Opluridae corresponds to the
“oplurines” of Smith et al. (1973), Etheridge in
Paull et al. (1976), and Etheridge and de Queiroz
(1988). The nomenclatural status of Doryphori-
dae is arguable. Fitzinger (1843) erected the
family Doryphori based on “Doryphorus
(Cuv.),” the parentheses around Cuvier meaning,
in Fitzinger’s words (1843:15) “Citata uncinis
inclusa auctores indicant, qui genus quidem
nominaverunt, sed non stricte in eodem sensu
proposuerunt” (=the parentheses enclose the
indicated authors of the genus name, although
the names are not used strictly in the original
sense as proposed). The problem lies in that
Doryphorus Cuvier has a type species set by
monotypy, Stellio brevicaudatus Latreille, 1802
(= Uracentron azureum), not included in Fitzin-
ger’s sense of the genus. Fitzinger regarded the
type species of Doryphorus to be Hoplurus max-
imiliani Duméril and Bibron, 1837 (=Oplurus
cyclurus Merrem, 1820). Article 65(b) of the
International Code of Zoological Nomenclature
(1985) requires that such “altered concept” prob-
lems be referred to the Commission for ruling.
However, pending application, we employ the
name Opluridae herein, rather than resurrecting a
name beset with nomenclatural difficulties.

PHRYNOSOMATIDAE FITZINGER, 1843

1843. Phrynosomata Fitzinger, Syst. Rept.,
1:17. Type genus: Phrynosoma Wieg-
mann, 1828.
1971. Sceloporinae Kastle, Grzimek’s Tierle-
ben, 6:181-182. Type genus: Scelo-
porus Wiegmann, 1828.
Characterization.—(1) maxillae not meet-
ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof not strongly rugose; (4) jugal and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal; (6) parietal foramen in fronto-

parietal suture; (7) supratemporal sits on lateral
side of supratemporal process of parietal; (8)
nuchal endolymphatic sacs not penetrating
nuchal musculature; (9) dentary not expanded
onto labial face of coronoid; (10) labial blade of
coronoid poorly developed or absent; (11) ante-
rior surangular foramen above posteriormost
extent of dentary; (12) Meckel’s groove not
fused; (13) splenial relatively long anteriorly;
(14) dentary and maxillary teeth pleurodont, not
fused to underlying bone in adults; (15) palatine
teeth absent; (16) pterygoid teeth absent; (17)
posterior process of interclavicle invested by
sternum far anteriorly; (18) caudal autotomy
fracture planes present (except in Phrynosoma),
with transverse processes anterior to fracture
planes; (19) posterior coracoid fenestra absent;
(20) sternal fontanelle enlarged and median; (21)
sternal ribs number 3 or 4 (Petrosaurus); (22)
postxiphisternal inscriptional ribs short; (23)
interparietal scale large (except in Phrynosoma
and Uma); (24) mid-dorsal scale row absent; (25)
gular fold complete medially (except Sce-
loporus); (26) femoral pores present; (27) spinu-
late scale organs absent; (28) sink-trap nasal
apparatus; nasal vestibule long, straight, sup-
ported by elongated septomaxilla; concha absent
(29) hemipenes unicapitate, unisulcate, with
enlarged posterior lobe; (30) colic septa absent.

Content.—Callisaurus Blainville, 1835;
Holbrookia Girard, 1851 (including Cophosau-
rus Troschel, 1852); Petrosaurus Boulenger,
1885; Phrynosoma Wiegmann, 1828; Sceloporus
Wiegmann, 1828 (including Sator Dickerson,
1919); Uma Baird, 1858; Urosaurus Hallowell,
1854; Uta Baird and Girard, 1852.

Distribution.—Southern Canada through the
USA to Panama (Fig. 22).

Comment.—Phrynosomatidae corresponds
to the “sceloporines” of Savage (1958), Eth-
eridge (1964), Presch (1969), and Etheridge and
de Queiroz (1988).

POLYCHRIDAE FITZINGER, 1843

1826. Pneustoidea Fitzinger, Neue Classif.
Rept.:11. Type genus: Not stated. See
comment.

1843. Polychri Fitzinger, Syst. Rept., 1:16.
Type genus: Polychrus Cuvier, 1817.

1843. Dactyloae Fitzinger, Syst. Rept., 1:17.

42 MISCELLANEOUS PUBLICATIONS

Fig. 22. Distribution of Phrynosomatidae.

Type genus: Dactyloa Wagler, 1830
(=Anolis Daudin, 1802).

1843. Draconturae Fitzinger, Syst. Rept.,
1:17. Type genus: “Dracontura Wagler”
(=Draconura Wagler, 1830 =Anolis
Daudin, 1802).

1864. Anolidae Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:227. Type genus: Ano-
lis Daudin, 1802.

Characterization.—(1) maxillae not meet-

ing anteromedially behind palatal portion of

premaxilla; (2) lacrimal foramen not enlarged;
(3) skull roof strongly rugose (except in Dip-
lolaemus and some Anolis); (4) jugal and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal or V or Y-shaped; (6) parietal
foramen normally in frontoparietal suture (in
parietal in some Anolis and lacking in some
Polychrus); (7) supratemporal sits on lateral side
of supratemporal process of parietal; (8) nuchal
endolymphatic sacs penetrating nuchal muscula-
ture; (9) dentary not expanded onto labial face of

IGUANIAN LIZARD PHYLOGENY 43

coronoid; (10) labial blade of coronoid variable,
from well-developed to absent; (11) anterior
surangular foramen above posteriormost extent
of dentary; (12) Meckel’s groove fused; (13)
splenial relatively to very short anterior to level
or apex of coronoid; (14) dentary and maxillary
teeth pleurodont, not fused to underlying bone in
adults; (15) palatine teeth present (except
Polychrus and anoles other than Chamaeolis);
(16) pterygoid teeth present (except some Poly-
chrus); (17) posterior process of interclavicle
variably invested by sternum far anteriorly; (18)
caudal autotomy fracture planes present or ab-
sent, with transverse processes anterior or poste-
rior to fracture planes, if present; (19) posterior
coracoid fenestra small or absent; (20) sternal
fontanelles very small or absent; (21) sternal ribs
number 2, 3, or 4; (22) postxiphisternal inscrip-
tional ribs long, confluent medially; (23) inter-
parietal scale not enlarged; (24) mid-dorsal scale
row variable; (25) gular fold complete medially
(except Polychrus and anoles); (26) femoral
pores absent (except Polychrus); (27) spinulate
scale organs present (except Polychrus); (28)
primitive nasal apparatus; nasal vestibule rela-
tively short, straight; concha present or absent;
(29) hemipenes variable—plesiomorphically
bicapitate, bisulcate (unicapitate in some Ano-
lis); (30) colic septa absent.

Content.—Anisolepis Boulenger, 1885
(including Aptycholaemus Boulenger, 1891, fide
Etheridge and Williams, unpubl.); Anolis
Daudin, 1802; Chamaeolis Cocteau, 1838; Cha-
maelinorops Schmidt, 1919; Diplolaemus Bell,
1843; Enyalius Wagler, 1830; Leiosaurus
Duméril and Bibron, 1837 (including Aperopris-
tis Peracca, 1897); Phenacosaurus Barbour,
1920; Polychrus Cuvier, 1817; “Pristidactylus”
Fitzinger, 1843; Urostrophus* Duméril and
Bibron, 1837.

Distribution.—Southern North America to
southern South America; West Indies (Fig. 23).

Comment.—Polychridae corresponds in
content to the “anoloids” of Etheridge and Wil-
liams (1985) and Etheridge and de Queiroz
(1988).

We select Polychri Fitzinger, 1843, to have
priority over Dactyloae Fitzinger, 1843, and
Draconturae Fitzinger, 1843, under the provi-

sions of Article 24 (“Principle of the First Revi-
sor’) and Recommendation 24A of the Interna-
tional Code of Zoological Nomenclature (1985).

Apparently owing to a typographic error, Fitz-
inger (1826) did not mention the apparent type
genus of Pneustidae, Pneustes Merrem, 1820
(type species: P. prehensilis Merrem, 1820).
Pneustes prehensilis is a nomen dubium (Smith,
1957), considered to be a synonym of Laemanc-
tus vautieri (=Urostrophus vautieri) by Fitzinger
(1843:62), but probably a synonym of Polychrus
acutirostris Spix, 1825 (P. E. Vanzolini, in litt.).
Note, however, that certain features noted in the
diagnosis of Pneustes prehensilis (e.g., four toes
on each foot) render it a nomen dubium. Not
having an “express reference” or “inference in
context” to a type genus in Fitzinger’s work
prevents Pneustoidea from being an available
family group name (Art. 11.f.i1.1; International
Code of Zoological Nomenclature, 1985), as
does the status of Pneustes as a nomen dubium.

We do not follow Guyer and Savage (1986) in
recognizing the nominal genera Ctenonotus Fitz-
inger, 1843, Dactyloa Wagler, 1830, Norops
Wagler, 1830, and Semiurus Fitzinger, 1843, as
distinct from Anolis because the phylogenetic
basis for recognition of these other taxa is argu-
able (Cannatella and de Queiroz, 1989).

TROPIDURIDAE BELL, 1843
1843. Tropiduridae Bell, Zool Voy. Beagle:1.

Type genus: Tropidurus Wied-
Neuwied, 1825. See comment under
Tropidurinae.

1843. Ptychosauri Fitzinger, Syst. Rept., 1:16.
Type genus: Ptychosaurus Fitzinger,
1843 (=Plica Gray, 1831).

1843. Steirolepides Fitzinger, Syst. Rept.,
1:17. Type genus: Steirolepis Fitzinger,
1843 (=Tropidurus Wied-Neuwied,
1825).

1843. Heterotropides Fitzinger, Syst. Rept.,
1:17. Type genus: Heterotropis Fitzin-
ger, 1843 (a nomen dubium) (=?Steno-
cercus Duméril and Bibron, 1837).

Characterization.—(1) maxillae not meet-

ing anteromedially behind palatal portion of
premaxilla; (2) lacrimal foramen not enlarged;

44 MISCELLANEOUS PUBLICATIONS

<}

ogi

ee
———————————————
SSS SS SSS
——
SSS
SS a SSS

So
~~
N

li
i
|

|

ll

O 3000

_——————SSSsSsSsSSSs

Kilometers

HI

on

“inp

--- A

v

/
f
(p

Fig. 23. Distribution of Polychridae.

IGUANIAN LIZARD PHYLOGENY 45

(3) skull roof not strongly rugose; (4) jugal and
squamosal not broadly juxtaposed; (5) parietal
roof trapezoidal (or V-shaped in Leiocephalus);
(6) parietal foramen in frontoparietal suture or
absent; (7) supratemporal sits on lateral or medial
side, or in ventral groove of supratemporal pro-
cess of parietal; (8) nuchal endolymphatic sacs
not penetrating nuchal musculature; (9) dentary
not expanded onto labial face of coronoid; (10)
labial blade of coronoid poorly developed or
absent (Tropidurinae) or well-developed (Lio-
laeminae, Leiocephalinae); (11) anterior suran-
gular foramen superior to posteriormost extent of
dentary; (12) Meckel’s groove variably fused
(except some Phymaturus, Ctenoblepharys, and
some Liolaemus); (13) splenial very short ante-
riorly (except in some liolaemines); (14) dentary
and maxillary teeth pleurodont, not fused to
underlying bone in adults; (15) palatine teeth
absent; (16) pterygoid teeth present (except some
Leiocephalus, some “Stenocercus”); (17) poste-
rior process of interclavicle not invested by ster-
num far anteriorly (except in the “Tropidurus”
group); (18) caudal autotomy fracture planes
present, with transverse processes anterior to
fracture planes; (19) posterior coracoid fenestra
present (except in Phymaturus and Ctenoble-
pharys); (20) sternal fontanelles median and
enlarged (except in some “TJropidurus’”); (21)
sternal ribs number 3 or 4; (22) postxiphisternal
inscriptional ribs short (long in some “Ophryoes-
soides” and some “Stenocercus”’); (23) interpari-
etal scale variable, enlarged only in the “Tropi-
durus” group of Tropidurinae; (24) mid-dorsal
scale row generally present (except in Liolaem-
inae, some “Stenocercus,” and some members of
the “Tropidurus” group); (25) gular fold incom-
plete medially; (26) femoral pores absent; (27)
spinulate scale organs absent; (28) generally
primitive nasal apparatus; nasal vestibule rela-
tively short, straight; concha present, generally
free, but fused to the roof of the nasal chamber in
Tropidurinae; (29) hemipenes variable; (30)
colic septa absent.

Content.—Leiocephalinae new subfamily;
Liolaeminae new subfamily; and Tropidurinae
Bell, 1843.

Distribution.—The Bahama Islands, Cuba
and Hispaniola and associated banks; Cayman
Islands; South America, excluding northern

Colombia and northern Venezuela, southward to
northern Tierra del Fuego; Galapagos Islands
(Fig. 24).

Comment.—Tropiduridae corresponds to the
“tropidurines” of Etheridge (1966) and Etheridge
and de Queiroz (1988).

LEIOCEPHALINAE NEW SUBFAMILY

Type genus.—Leiocephalus Gray, 1825.

Characterization.—(1) premaxillary spine
overlapped by nasals; (2) parietal table Y or V-
shaped (shared with some iguanids and anoles);
(3) large labial blade of coronoid; (4) anterior
extent of splenial extending more than % length
of precoronoid length of mandible; (5) well-
developed anterior process of interclavicle; (6)
interparietal scale not enlarged; (7) no preanal
pores; (8) primitive nasal condition, nasal vesti-
bule short and straight, nasal concha not fused to
roof of nasal chamber; (9) unicapitate, unisulcate
hemipenes.

Content.—Leiocephalus Gray, 1825.

Distribution.—Bahama Islands, Cuba and
Hispaniola and associated banks (extinct on
Jamaica, Puerto Rico, Barbuda, Antigua, An-
guilla, Martinique, and Guadeloupe [Pregill et
al., 1988]); Cayman Islands (Fig. 24).

LIOLAEMINAE NEW SUBFAMILY

Type genus.—Liolaemus Wiegmann, 1834.

Characterization.—(1) premaxillary spine
overlapped by nasals (except in some Phyma-
turus); (2) parietal table not Y or V-shaped; (3)
large labial blade of coronoid; (4) anterior extent
of splenial extending more than '% length of
precoronoid length of mandible; (5) poorly de-
veloped anterior process of interclavicle; (6)
interparietal scale not enlarged; (7) preanal pores
(except in some species of Liolaemus); (8) S-
nasal condition, nasal concha not fused to roof of
nasal chamber; (9) weakly bicapitate, unisulcate
hemipenes.

Content.—Ctenoblepharys Tschudi, 1845;
Liolaemus Wiegmann, 1834; Phymaturus Grav-
enhorst, 1838 (see comment).

Distribution.—Coastal and Andean Peru
southward through Bolivia, Chile, and Argentina
to northern Tierra del Fuego and the coasts of
Uruguay and southeastern Brazil (Fig. 24).

Comment.—We have not followed Cei and

46 MISCELLANEOUS PUBLICATIONS

Tropidurinae | =

O Kekexe)
_ za
Kilometers

Fig. 24. Distribution of Tropiduridae, including subfamilies. The solid line surrounding the distribution of Leiocephalinae
indicates that late Pleistocene—Early Holocene distribution of that taxon.

IGUANIAN LIZARD PHYLOGENY 47

Lescure (1985) and Cei (1986) in the use of
Centrura Bell, 1843, instead of Phymaturus
Molina, 1768. As noted by Cei and Lescure
(1985), the nomenclatural confusion was due to
misidentification of type species in the original
description of Phymaturus. In these cases it is
required that traditional usage be maintained and
that the International Commission of Zoological
Nomenclature be petitioned to resolve the prob-
lem (Art. 70b; International Code of Zoological
Nomenclature, 1985).

TROPIDURINAE BELL, 1843

1843. Tropiduridae Bell, Zool Voy. Beagle: 1.
Type genus: Tropidurus Wied-
Neuwied, 1825. See comment.

1843. Ptychosauri Fitzinger, Syst. Rept.,
1:16. Type genus: Ptychosaurus Fitzin-
ger, 1843 (=Plica Gray, 1831).

1843. Steirolepides Fitzinger, Syst. Rept.,
1:17. Type genus: Steirolepis Fitzinger,
1843 (=Tropidurus Wied-Neuwied,
1825).

1843. Heterotropides Fitzinger, Syst. Rept.,
1:17. Type genus: Heterotropis Fitzin-
ger, 1843 (a nomen dubium) (=?Steno-
cercus Duméril and Bibron, 1837).

Characterization.—(1) premaxillary spine
not overlapped by nasals; (2) parietal table not Y
or V-shaped; (3) no labial blade of coronoid; (4)
splenial extending anteriorly no more than %
length of precoronoid length of mandible; (5)
anterior process of interclavicle poorly devel-
oped or absent; (6) interparietal scale enlarged;
(7) no preanal pores; (8) nasal concha fused to
roof of nasal chamber, nasal vestibule relative
short and straight; (9) bisulcate, weakly to
strongly bicapitate hemipenes.

Content.—“Ophryoessoides” Duméril,
1851; Plica® Gray, 1831; Proctotretus Duméril
and Bibron, 1837; “Stenocercus” Duméril and
Bibron, 1837; Strobilurus Wiegmann, 1834; Tap-
inurus Amaral, 1933; “Tropidurus” Wied-
Neuwied, 1825; Uracentron Kaup, 1826; Urano-
scodon Kaup, 1825.

Distribution.—South America, excluding
northern Colombia and northern Venezuela,
southward to northern Chile and central Argen-
tina; Galapagos Islands (Fig. 24).

Comment.—We follow Smith and Grant
(1958) in regarding Bell’s (1843) publication of
Tropiduridae to have priority over Fitzinger’s
(1843) publication of Steirolepides, Ptychosauri,
and Heterotropides.

SUMMARY

A phylogenetic analysis of Iguania was per-
formed using 67 transformation series contain-
ing 147 characters of osteology, dentition, squa-
mation, internal nasal structure, musculature,
and hemipenes. For analysis, 35 taxonomic units,
representing all iguanians, were used. Data
analysis was performed using PAUP version
2.4.1 (Swofford, 1985) and Hennig86 version 1.5
(Farris, 1988). No evidence of monophyly was
discovered for “Iguanidae” and only ambiguous
evidence for the monophyly of Agamidae*, al-
though some lines of evidence support the view
that these nominal taxa are paraphyletic. The
historical reality (=monophyly) of Chamaele-
onidae was highly corroborated. A total of 225
alternative supported tree topologies were dis-
covered (208 steps, C.I.=0.385). These alterna-
tive topologies were produced by 12 unrooted

networks, that could be variously rooted to pro-
duce 18 trees of nine major monophyletic groups
(acrodonts [=Agamidae* + Chamaeleonidae],
anoloids, basiliscines, crotaphytines, iguanines,
morunasaurs, oplurines, sceloporines, and tro-
pidurines), and alternative topologies within
these monophyletic groups. These alternatives
are variably dependent on unrooted network
topology. Two topologies were discovered within

8 With the removal of the nomen oblitum rule (Art.
23b of the 1961 International Code) in the 1985
International Code, the oldest name available for this
taxon becomes Hypsibatus Wagler, 1830. However,
ongoing work by Frost will obviate this anomaly.
Therefore, we retain Plica for purposes of this
publication.

48 MISCELLANEOUS PUBLICATIONS

the Liolaemus group of the tropidurines, three in
the sceloporines, two in the acrodonts, and three
in the anoloids. In order to obviate the possibility
of paraphyly and to reform named but misleading
groupings into historically real components, and
to demonstrate our ignorance of intergroup rela-
tionships, “Iguanidae” was partitioned into 8 taxa
sedis mutabilis, ranked as families: Corytopha-
nidae Fitzinger, 1843 (former basiliscines); Cro-
taphytidae Smith and Brodie, 1982 (former cro-
taphytines); Hoplocercidae new name (former
morunasaurs); Iguanidae Oppel, 1811 (former
iguanines); Opluridae Moody, 1983; Phrynoso-
matidae Fitzinger, 1843 (former sceloporines);
Polychridae Fitzinger, 1843 (former anoloids);
Tropiduridae Bell, 1843 (former tropidurines).

Within Tropiduridae three subfamilies were
recognized sedis mutabilis: Leiocephalinae new
name; Liolaeminae new name; Tropidurinae
Bell, 1843. Agamidae* may be paraphyletic with
respect to Chamaeleonidae; to correct this, three
monophyletic subfamilies (Chamaeleoninae
Rafinesque, 1815; Agaminae Spix, 1825; and
Leiolepidinae Fitzinger, 1843) were recognized,
sedis mutabilis, within a reconstituted Chamae-
leonidae Rafinesque, 1815 (equivalent to Acro-
donta Cope, 1864), that is a highly corroborated
monophyletic group. Relationships among the
family-groups are poorly resolved, and much of
the topological differences between discovered
trees was because of this lack of resolution.

LITERATURE CITED

ARNOLD, E. N. 1981. Estimating phylogenies at low
taxonomic levels. Z. Zool. Syst. Evolutionsforsch.,
19:1-35.

ARNOLD, E. N. 1984. Variation in the cloacal and hem-
ipenial muscles of lizards and its bearing on their
relationships. pp. 57-85 In: M. J. Ferguson (ed.),
The structure, development and evolution of rep-
tiles. Symp. Zool. Soc. London, 52.

AXTELL, R. W. 1958. A monographic revision of the
iguanid genus Holbrookia. Ph.D. Dissertation,
Univ. Texas, Austin. 222 pp.

AXTELL, R. W., AND A. O. WASSERMAN. 1953. Inter-
esting herpetological records from southern Texas
and northern Mexico. Herpetologica, 9:1-6.

BELL, T. 1825. Description of a new species of lizard.
Zool. J., London, 1825:457-460.

BELL, T. 1843. Zoology of the voyage of the H. M. S.
Beagle, under the command of Captain Fitzroy, R.
N., during the years 1832 to 1836. Edited and
superintended by Charles Darwin . . . naturalist to
the expedition. Part 5. Reptiles. vi + 51 pp., atlas.
Smith, Elder & Co., London.

Boume, W. 1988. Genitalmorphologie der Sauria:
Funktionelle und stammesgeschichtliche Aspekte.
Bonn. Zool. Monogr., 27:1-176.

Borsuk-Biatynicka, M., AND S. M. Moony. 1984.
Priscagaminae, a new subfamily of the Agamidae
(Sauria) from the Late Cretaceous of the Gobi
Desert. Acta Palaeontol. Polon., 29:51-81.

Brooks, D. R., AND E. O. Witey. 1985. Theories and
methods in different approaches to phylogenetic
systematics. Cladistics, 1:1—12.

Camp, C. 1923. Classification of the lizards. Bull. Am.
Mus. Nat. Hist., 48:289-481.

CANNATELLA, D. C., AND K. DE QuEIROZ. 1989. Phy-
logenetic systematics of the anoles: Is anew taxon-
omy warranted? Syst. Zool., 38:57-69.

Cer, J. M. 1979a. Remarks on the South American
lizard Liolaemus anomalus Koslowsky, and the
synonymy of Phrynosaura werneri Miiller (Rep-
tilia, Lacertilia, Iguanidae). J. Herpetol.,
13:183-186.

Cer, J. M. 1979b. A reassessment of the genus Cteno-
blepharis (Reptilia, Sauria, Iguanidae) with a de-
scription of a new subspecies of Liolaemus multi-
maculatus from western Argentina. J. Herpetol.,
13:297-302.

Cel, J. M. 1986. Reptiles del centro, centro-oeste y sur
de la Argentina. Herpetofauna de las zonas 4ridas y
semidridas. Mus. Reg. Sci. Nat. Torino, Monogr.,
4:1-526.

Cel, J. M., AND J. Lescure. 1985. Identité de Lacerta
palluma Molina, 1782, et révalidation de Centrura
flagellifer Bell, 1843 (Reptilia, Sauria). Bull. Mus.
Natl. Hist. Nat., Paris, (4)7(A)2:175-183.

Cel, J. M., AND J. A. ScoLaro. 1982. Anew species of
the Patagonian genus Vilcunia, with remarks on its
morphology, ecology, and distribution. J. Herpe-
tol., 16:354-363.

Cooper, J. S., D. F. G. Poo.e, AND R. Lawson. 1970.
The dentition of agamid lizards with special refer-
ence to tooth replacement. J. Zool., London,
162:85-98.

Cope, E. D. 1864. On the characters of the higher

IGUANIAN LIZARD PHYLOGENY 49

groups of Reptilia Squamata—and especially of
the Diploglossa. Proc. Acad. Nat. Sci. Philadel-
phia, 16:224—231.

Corr, E. D. 1900. The crocodilians, lizards and snakes
of North America. Annu. Rep. U.S. Natl. Mus. for
1898:153-1270.

Cuvier, G. 1817. Le régne animal distribué d’aprés
son organisation, pour servir de base d’histoire
naturelle des animaux et d’introduction 4 |’ anato-
mie comparée. Vol. 2, Les reptiles, les poissons, les
mollusques, et les aniélides. Deterville, Paris.

DE Queiroz, K. 1982. The scleral ossicles of the sce-
loporine iguanids: Areexamination with comments
on their phylogenetic significance. Herpetologica,
38:302-311.

DE Queiroz, K. 1985. The ontogenetic method for de-
termining character polarity and its relevance to
phylogenetic systematics. Syst. Zool., 34:280-299.

DE QuetRroz, K. 1987. Phylogenetic systematics of
iguanine lizards: Acomparative osteological study.
Univ. California Publ., Zool., 118:x1 + 203.

DE QuerRoz, K., AND M. T. DonocGuue. 1988. Phylo-
genetic systematics and the species problem. Cla-
distics, 4:317-338.

Drxon, J. R., AND J. W. WricuT. 1975. A review of the
lizards of the iguanid genus Tropidurus in Peru.
Contrib. Sci. Nat. Hist. Mus. Los Angeles Cty.,
271:1-39.

DonocuuE, M. J. 1985. A critique of the biological
species concept and recommendations for a phylo-
genetic alternative. Bryologist, 88:172-181.

Donoso-Barros, R. 1972. Contribucién al cono-
cimiento del género Ctenoblepharis Tschudi y
Phrynosaura Wermer (Sauria: Iguanidae). Bol. Soc.
Concepcion, 44:129-144.

Donoso-Barros, R. 1973. Un nuevo saurio de Bo-
livia. Neotropica, Buenos Aires, 19(60):132-134.

ELpreDcE, N., AND J. CRAcRaFT. 1980. Phylogenetic
patterns and the evolutionary process: Method and
theory in comparative biology. Columbia Univ.
Press, New York. 349 pp.

EL Tausi, M. R., AND H. M. Bisnar. 1959. On the
anatomy and histology of the digestive tract of the
lizard Uromastyx aegyptia (Forskal). Bull. Fac.
Sci. Cairo Univ., 34:13-50.

Estes, R. 1983a. Handbuch der Palaoherpetologie.
Encyclopedia of paleoherpetology. Part 10A, Sau-
ria terrestria, Amphisbaenia. Gustav Fischer,
Stuttgart and New York. xxii + 249 pp.

Estes, R. 1983b. The fossil record and early distribu-
tion of lizards. pp. 365-398 In: A. G. J. Rhodin and
K. Miyata (eds.), Advances in herpetology and
evolutionary biology: Essays in the honor of Ernest

E. Williams. Cambridge, Massachusetts.

Estes, R., J. GAUTHIER, AND K. DE QUEIROZ. 1988.
Phylogenetic relationships within Squamata. pp.
119-282 Jn: R. Estes and G. Pregill (eds.), Phylo-
genetic relationships of lizard families: Essays
commemorating Charles L. Camp. Stanford Univ.
Press.

Estes, R., AND L. I. Price. 1973. Iguanid lizard from
the Upper Cretaceous of Brazil. Science,
180:748-751.

ETHERIDGE, R. 1959. The relationships of the anoles
(Reptilia: Sauria: Iguanidae): An interpretation
based on skeletal morphology. Ph.D. Dissertation,
Univ. Michigan, Ann Arbor. xili + 236 pp.

ETHERIDGE, R. 1964. The skeletal morphology and
systematic relationships of sceloporine lizards.
Copeia, 1964:610-631.

ETHERIDGE, R. 1965. The abdominal skeleton of liz-
ards in the family Iguanidae. Herpetologica,
21:161-168.

ETHERIDGE, R. 1966. The systematic relationships of
West Indian and South American lizards referred to
the iguanid genus Leiocephalus. Copeia,
1966:79-91.

ETHERIDGE, R. 1967. Lizard caudal vertebrae. Copeia,
1967:699-721.

ETHERIDGE, R. 1969a. Tropidogaster blainvillii
Duméril & Bibron, 1837 (Reptilia, Sauria): Pro-
posed suppression under the Plenary Powers. Bull.
Zool. Nomencl., 25:224—226.

ETHERIDGE, R. 1969b. A review of the iguanid lizard
genus Enyalius. Bull. Br. Mus. (Nat. Hist.), Zool.,
18:231-260.

ETHERIDGE, R., AND K. DE QuEIROZ. 1988. A phylo-
geny of Iguanidae. pp. 283-368 Jn: R. Estes andG.
Pregill (eds.), Phylogenetic relationships of lizard
families: Essays commemorating Charles L. Camp.
Stanford Univ. Press.

ETHERIDGE, R., AND E. E. WILLIAMS. 1985. Notes on
Pristidactylus (Squamata: Iguanidae). Breviora,
483:1-18.

Evans, S. 1980. The skull of a new eosuchian reptile
from the Lower Jurassic of South Wales. Zool. J.
Linn. Soc., 70:203-264.

Evans, S. 1981. The postcranial skeleton of the Lower
Jurassic eosuchian Gephyrosaurus bridensis. Zool.
J. Linn. Soc., 73:81-116.

Farris, J. S. 1970. Methods for computing Wagner
trees. Syst. Zool., 19:83-92.

Farris, J. S. 1982. Outgroups and parsimony. Syst.
Zool., 31:328-334.

Farris, J. S. 1988. Hennig86, Version 1.5. Reference.
Privately printed.

50 MISCELLANEOUS PUBLICATIONS

FELSENSTEIN, J. 1978. The number of evolutionary
trees. Syst. Zool., 27:27-33.

Frrzincer, L. I. 1826. Neue Classification der Reptil-
ien nach ihren Natiirlichen Verwandtschaften nebst
einer Verwandts-Tafel und einem Verzeichnisse
der Reptilien-Sammlung des k. k. Zoologischen
Museum zu Wien. J. G. Hiibner, Wien. vii + 66 pp.

Firzincer, L. I. 1843. Systema Reptilium. Fasciculus
primus. Amblyglossae. Baumiiller and Seidel,
Wien. 106 pp.

Frost, D. R. 1987. A phylogenetic analysis of the
Tropidurus group of iguanian lizards, with com-
ments on the relationships within the Iguania
(Squamata). Ph.D. Dissertation, Univ. Kansas,
Lawrence. xi + 350 pp.

Frost, D. R., AND D. M. Hits. 1990. Species in
concept and practice: Herpetological applications.
Herpetologica, 46: in press.

Gasg, M., AND H. Saint Grrons. 1969. Données his-
tologiques sur les glandes salivaires des lépidosau-
riens. Mem. Mus. Natl. Hist. Nat., Paris, 58:1-112.

GauTHIER, J. A. 1986. Saurischian monophyly and the
origin of birds. pp. 1-55 Jn: K. Padian (ed.), The
origin of birds and the evolution of flight. Mem.
California Acad. Sci., 8:i—vii, 1-98.

GautTuigr, J. A., R. Estes, AND K. DE QUEIROZ. 1988.
A phylogenetic analysis of Lepidosauromorpha.
pp. 15—98 In: R. Estes and G. Pregill (eds.), Phylo-
genetic relationships of lizard families: Essays
commemorating Charles L. Camp. Stanford Univ.
Press.

Gorman, G. C., R. B. Hugy, anp E. E. WILLIAMS.
1969. Cytotaxonomic studies on some unusual
iguanid lizards assigned to the genera
Chamaeleolis, Polychrus, Polychroides, and
Phenacosaurus, with behavioral notes. Breviora,
316:1-17.

Gray, J. E. 1825. A synopsis of the genera of reptiles
and Amphibia, with a description of some new
species. Ann. Philos., (2)10:193-217.

Gray, J. E. 1827. A description of new species of
saurian reptiles; with a revision of the species of
chamaeleons. Philos. Mag. Ann., (2)2:207-214.

Guyer, C., AND J. M. Savace. 1986. Cladistic rela-
tionships among anoles. Syst. Zool., 35:509-531.

Hennic, W. 1966. Phylogenetic systematics. Univ.
Illinois Press, Urbana. 263 pp.

Hittentus, D. 1986. The relationship of Brookesia,
Rhampholeon and Chamaeleo (Chamaeleonidae,
Reptilia). Bijdr. Dierkd., 56:29-38.

Hu tt, D. L. 1964. Consistency and monophyly. Syst.
Foye este

INTERNATIONAL COMMISSION ON ZOOLOGICAL No-
MENCLATURE. 1961. International Code of Zoologi-

cal Nomenclature, adopted by the XV International
Congress of Zoology. International Trust for Zoo-
logical Nomenclature, London. 176 pp.

INTERNATIONAL COMMISSION ON ZOOLOGICAL No-
MENCLATURE. 1985. International Code of Zoologi-
cal Nomenclature, Third Edition, Adopted by the
XX General Assembly of the International Union
of Biological Sciences. International Trust for Zoo-
logical Nomenclature, London. 338 pp.

Iverson, J. B. 1980. Colic modification in iguanine
lizards. J. Morphol., 163:79-93.

Iverson, J. B. 1982. Adaptations for herbivory in
iguanine lizards. pp. 60-76 Jn: G. M. Burghardt
and A. S. Rand (eds.), Iguanas of the world. Noyes
Publs., New York.

Jotur, M. T. 1960. The head skeleton of the lizard.
Acta Zool., Stockholm, 41:1-64.

JULLIEN, R., AND S. RENous-Lécuru. 1972. Vari-
ations du trajet du nerf ulnaire (ulnaris) et de 1’in-
nervation des muscles dorsaux de la jambe chez les
lacertiliens (reptiles: squamates): Valeur systéma-
tique et application phylogénétique. Bull. Mus.
Natl. Hist. Nat., (Zool.), Paris, 23(29):207-246.

KAsTLeE, W. 1971. Die Leguan. pp. 181—206 Jn: H. C.
B. Grzimek, H. Hediger, K. Klemmer, O. Kuhn,
and H. Wermuth (eds.), Grzimek’s Tierleben. vol.
6. Kriechthiere. Kindler, Ziirich.

Kaver, C. J. J., AND W. BOumeE. 1986. Phylogeny and
classification of the Chamaeleonidae (Sauria) with
special reference to hemipenis morphology. Bonn.
Zool. Monogr., 22:1-64.

K.iuce, A. 1983. Cladistic relationships among
gekkonid lizards. Copeia, 1983:465-475.

K.iuce, A. 1985. Ontogeny and phylogenetic sys-
tematics. Cladistics, 1:13-28.

Kuuag, A. 1989. Metacladistics. Cladistics, 5: in press.

Kiuce, A. G., AND J. S. Farris. 1969. Quantitative
phyletics and the evolution of anurans. Syst. Zool.,
18:1-32.

Lana, M. A. 1989. Phylogenetic and biogeographic
patterns of basiliscine iguanians (Reptilia: Squa-
mata: “Iguanidae”). Bonn. Zool. Monogr.,
28:1-172.

LapacE, E. O. 1926. The septomaxillary of the Anura
and Reptilia. J. Morphol. Physiol., 46:399-431.

Larsen, K. R., AND W. W. TANNER. 1975. Evolution of
the sceloporine lizards ([guanidae). Gt. Basin Nat.,
35:1-20.

LaureENT, R. F. 1983a. Sinonimia del género Pelusau-
rus Donoso-Barros con Liolaemus (Sauria: Igua-
nidae). Bol. Assoc. Herpetol. Argentina, Buenos
Aires, 1(1):9-10.

Laurent, R. F. 1983b. Contribucién al conocimiento
del estructura taxonémica del género Liolaemus

IGUANIAN LIZARD PHYLOGENY yi t

Wiegmann (Iguanidae). Bol. Assoc. Herpetol.
Argentina, Buenos Aires, 1(3):16-18.

LavrENT, R. F. 1984. On some iguanid genera related
to or previously confused with Liolaemus Wieg-
mann. J. Herpetol., 18:357-373.

Laurent, R. F. 1985. Segunda contribucién al cono-
cimiento de la estructura taxondémica del género
Liolaemus Wiegmann (Iguanidae). Cuad. Herpe-
tol., Assoc. Herpetol. Argentina, Buenos Aires,
1(6):1037.

LaurENTI, J. N. 1768. Specimen medicum exhibens
synopsin reptilium emendatum cum experimentis
circa venena et antidota reptilium austriacorum.
Trattnern, Wien. 214 pp.

L&écuru, S. 1968. Remarques sur le scapulo-coracoide
des lacertiliens. Ann. Sci. Nat., Zool., 10:475-510.

LONNBERG, E. 1902. On some points of relation be-
tween the morphological structure of the intestine
and the diet of reptiles. Bih. Svenska Vetensk.
Akad. Handl., (4)28(8):3-53.

Luxe, C. 1986. Convergent evolution of lizard toe
fringes. Biol. J. Linn. Soc., 27:1-16.

Mapoison, W. P., M. J. DoNoGHUE, AND D. R. Map-
DISON. 1984. Outgroup analysis and parsimony.
Syst. Zool., 33:83-103.

Mapopison, W. P., AND D. R. Mappison. 1987. Mac-
Clade. Version 2.1. User’s Manual. Privately
printed.

Matan, M. E. 1946. Contributions to the comparative
anatomy of the nasal capsule and the Organ of
Jacobson of the Lacertilia. Ann. Univ. Stellen-
bosch, 24A:69-137.

Mepica, P. P., F. B. TURNER, AND D. D. Situ. 1973.
Hormonal induction of color change in female
leopard lizards, Crotaphytus wislizenii. Copeia,
1973:658-661.

MirTLemaNn, M. B. 1942. A summary of the iguanid
lizard genus Urosaurus. Bull. Mus. Comp. Zool.,
2:105-181.

Moony, S. M. 1980. Phylogenetic and historical bio-
geographical relationships of the genera in the
family Agamidae (Reptilia: Lacertilia). Ph.D. Dis-
sertation, Univ. Michigan, Ann Arbor. 373 pp.

Moony, S. M. 1983a. The rectus abdominis muscle
complex of the Lacertilia: Terminology, homology,
and assumed presence in primitive iguanian liz-
ards. pp. 195-212 In: A. G. C. Rhodin and K.
Miyata (eds.), Advances in herpetology and evolu-
tionary biology: Essays in honor of Ernest E. Wil-
liams. Cambridge, Massachusetts.

Moony, S. M. 1983b. Endolymphatic sacs in lizards:
Phylogenetic and functional considerations. Ab-
stract of paper presented as Soc. Stud. Amph. Rept.
and Herpetol. League annual meeting, Salt Lake
City, Utah.

NELson, G. 1979. Cladistic analysis and synthesis:
principles and definitions, with an historical note
on Adanson’s Familles des Plantes (1763-1764).
Syst. Zool., 28:1—21.

Nopsca, F. 1923. Die Familien der Reptilien. Fortschr.
Geol. Palaeont., Berlin, 2:1—210.

NortucuTt, R. G. 1978. Forebrain and midbrain or-
ganization in lizards and its phylogenetic signifi-
cance. pp. 11-64 Jn: N. Greenberg and P. D.
MacLean (eds.), Behavior and neurology of liz-
ards: An interdisciplinary colloquium. U.S. Dept.
Health, Educ. Welfare, Publ. No. (ADM) 77-491.

Novacrk, M. J. 1986. The skull of leptictid insecti-
vorans and the higher-level classification of euth-
erian mammals. Bull. Am. Mus. Nat. Hist.,
183:1-111.

Nunez, H., AND J. L. YANEz. 1983. Abas y Velosaura,
nuevos géneros de lagartos Iguanidae y proposi-
ciones sistemAticas respecto de los géneros aliados
(Reptilia: Squamata). Bol. Mus. Nac. Hist. Nat.
Chile, Santiago, 40:97-105.

OppeL, M. 1811. Die Ordnungen, Familien und Gat-
tungen der Reptilien als Prodrom einer Naturges-
chichte derselben. J. Lindauer, Miinchen.
xli+87pp.

Parsons, T. S. 1970. The nose and Jacobson’s Organ.
pp. 99-191 In: C. Gans (ed.), Biology of the Rep-
tilia. vol. 2. Academic Press, London and New
York.

PATTERSON, C. 1982. Morphological characters and
homology. pp. 21-74 In: K. A. Joysey and A. E.
Friday (eds.), Problems of phylogenetic recon-
struction. Academic Press, London.

PATTERSON, R. 1987. Reptiles of southern Africa. C.
Struik, Ltd., Cape Town. 128 pp.

PauLL, D., E. E. WILLIAMS, AND W. P. HALL. 1976.
Lizard karyotypes from the Galapagos Islands:
Chromosomes in phylogeny and evolution. Bre-
viora, 441:1-31.

Peterson, J. A. 1983. The evolution of the subdigital
pad in Anolis. I. Comparisons among the anoline
genera. pp. 245-283 In: A. G. C. Rhodin and K.
Miyata (eds.), Advances in herpetology and evolu-
tionary biology: Essays in honor of Emest E. Wil-
liams. Cambridge, Massachusetts.

Peterson, J. A., AND E. E. WiuiaMs. 1981. A case
history of retrograde evolution: the onca lineage in
anoline lizards. I. Subdigital fine structure. Bull.
Mus. Comp. Zool., 149:215-268.

PREGILL, G. K. 1984. An extinct species of
Leiocephalus from Haiti (Sauria: Iguanidae). Proc.
Biol. Soc. Washington, 97:827-833.

PREGILL, G. K., D. W. STEADMANN, S. L. OLSON, AND
F. B. Grapy. 1988. Late Holocene fossil verte-
brates from Burma Quarry, Antigua, Lesser An-

ay MISCELLANEOUS PUBLICATIONS

tilles. Smithson. Contrib. Zool., 463:i-111+1-27.

Prescu, W. 1969. Evolutionary osteology and rela-
tionships of the horned lizard genus Phrynosoma
(family Iguanidae). Copeia, 1969:250-275.

Prescu. W. 1988. Cladistic relationships within the
Scincomorpha. pp. 471-492 In: R. Estes and G.
Pregill (eds.), Phylogenetic relationships of lizard
families: Essays commemorating Charles L.Camp.
Stanford Univ. Press.

RAFINESQUE, C. S. 1815. Analyse de la nature ou Tab-
leau de l’univers et des corps organisés. J. Bar-
ravecchia, Palermo. 224 pp.

REEVE, W. L. 1952. Taxonomy and distribution of the
horned lizard Phrynosoma. Univ. Kansas Sci. Bull.,
34:817-960.

Renous, S. 1979. Application de principes cladis-
tiques a la phylogénése et la biogéographie des lac-
ertiliens. Gegenbaurs Morphol. Jahrb.,
125:376-432.

RiepPeL, O. 1981. The skull and jaw adductor muscu-
lature in chamaeleons. Rev. Suisse Zool.,
88:433-445.

RriEpPEL, O. 1987. The phylogenetic relationships
within the Chamaeleonidae, with comments on
some aspects of cladistic analysis. Zool. J. Linn.
Soc., 89:41-62.

SavaGE, J. M. 1958. The iguanid lizard genera
Urosaurus and Uta, with remarks on related
groups. Zoologica, New York, 43:41-54.

SCHWENK, K. 1988. Comparative morphology of the
lepidosaur tongue and its relevance to squamate
phylogeny. pp. 569-598 Jn: R. Estes and G. Pregill
(eds.), Phylogenetic relationships of lizard fami-
lies: Essays commemorating Charles L. Camp.
Stanford Univ. Press.

SLaBy, O. 1981. Morphogenesis of the nasal capsule,
the nasal epithelial tube and the organ of Jacobson
in a member of the family Agamidae. Folia Mor-
phol., Prague, 29:305-317.

SLaBy, O. 1984. Morphogenesis of the nasal apparatus
in amember of the genus Chamaeleon L. (Morpho-
genesis of the nasal capsule, the epithelial nasal
tube and the organ of Jacobson in Sauropsida—
VII). Folia Morphol., Prague, 32:225-246.

SmitH, H. M. 1946. Handbook of lizards. Comstock
Publ. Co., New York. xxi + 557 pp.

SmitH, H. M. 1957. Proposed use of the Plenary
Powers to validate the generic name “Phrynosoma”’
Wiegmann, 1828 (Class Reptilia, Order Squa-
mata), a name placed on the “Official List of Ge-
neric Names in Zoology” by the ruling given in

“Opinion” 92. Bull. Zool. Nomencl., 13:267-279.

SmiTH, H. M., AND E. D. Bropig, Jr. 1982. A guide to
field identification—Reptiles of North America.
Golden Press, New York. 240 pp.

SMITH, H. M., AND C. GRANT. 1958. The proper names
for some Cuban snakes: an analysis of dates of
publication of Ramon de la Sagra’s Historia Natu-
ral de Cuba, and of Fitzinger’s Systema Reptilium.
Herpetologica, 14:215—222.

SMITH, H. M., G. SINELNICK, J. D. FAWCETT, AND R. E.
Jones. 1973. A survey of the chronology of ovula-
tion in anoline genera. Trans. Kansas Acad. Sci.,
75:107-120.

SmiTH, M. A. 1932. Some notes on the monitors. J.
Bombay Nat. Hist. Soc., 35:615-619.

Sprx, J. B. von. 1825. Animalia nova sive species nova
lacertarum quas in itinere per Brasiliam annis
MDCCCXVII-MDCCCXX jussu et auspicius
Maximiliani Josephi I Bavariae Regis suscepto
collegit et descripsit Dr. J. B. de Spix. T. O. Weigel,
Lipsiae. 26 pp.

STEBBINS, R. C. 1943. Adaptations in the nasal pas-
sages for sand burrowing in the saurian genus Uma.
Am. Nat., 77:38-52.

STEBBINS, R. C. 1948. Nasal structure in lizards with
reference to olfaction and conditioning of the in-
spired air. Am. J. Anat., 83:183-222.

STEVENS, P. F. 1980. Evolutionary polarity of charac-
ter states. Annu. Rev. Ecol. Syst., 11:333-358.
Stmie, M. 1966. The cranial anatomy of the iguanid
Anolis carolinensis (Cuvier). Ann. Univ. Stellen-

bosch, 41A:243-268.

SUKHANOYV, V. B. 1961. Some problems of the phylo-
geny and systematics of Lacertilia (seu Sauria).
Zool. Zh., 40:73-83. [English translation by Scin-
tran, 1976. S. M. Moody and G. R. Zug (eds.).
Smithson. Herpetol. Inform. Serv., 38:1-15.]

SworrorD, D. L. 1985. PAUP—Phylogenetic Analy-
sis Using Parsimony. Version 2.4.1. User’s Manual.
Privately Published.

THEOBALD, W. 1868. Catalogue of the reptiles of Brit-
ish Birma, embracing the provinces of Pegu, Marta-
ban, and Tenasserim; with descriptions of new or
little-known species. J. Linn. Soc., London,
10:4-67.

Watrous, L. E., AND Q. D. WHEELER. 1981. The
outgroup comparison method of character analysis.
Syst. Zool., 30:1-11.

WELLS, R. W., AND C. R. WELLINGTON. 1983. A syn-
opsis of the class Reptilia in Australia. Austr. J.
Herpetol., 1(3-4):73-129.

IGUANIAN LIZARD PHYLOGENY 53

Wermuty, H. 1967. Liste der rezenten Amphibien
und Reptilien: Agamidae. Tierreich, 86:i—xiv,
1-127.

Wiey, E. O. 1979. An annotated Linnaean hierarchy,
with comments on natural taxa and competing
systems. Syst. Zool., 28:308-337.

Wiey, E. O. 1981a. Convex groups and consistent
classifications. Syst. Bot., 6:346-358.

Wuiey, E. O. 1981b. Phylogenetics: The theory and
practice of phylogenetic systematics. John Wiley
and Sons, New York. 439 pp.

APPENDIX 1

DaTA MATRIX OF IGUANIAN TERMINAL TAXA

SPHENO=Rhynchocephalia; SCLER=Sclero-
glossa; ANCES=Ancestor of ingroup (Iguania);
PRISC=+Priscagama*; AGAMI=agamids, excluding
other agamid terminal taxa; UROMA=Uromastyx;
LEIOL=Leiolepis; PHYSI=Physignathus; CHAME
=chameleons; POLYC=Polychrus; ENYAL=En-
yalius; PRIST=“Pristidactylus”; PARAA=Urostro-
phus* and Anisolepis,; ANOLE=anoles; ENYLD
=“Enyalioides”; BASIL=Basiliscus; CORYT=Cory-
tophanes; LAEMA=Laemanctus; PETRO=Petrosau-
rus; SCELO=Sceloporus; UROSA=Urosaurus;

UTA=Uta; PHRYN=Phrynosoma; SANDL=Sand
lizards;PH YMA=Phymaturus; CTENO=Cteno-
blepharys; LIOLA=Liolaemus, LEIOC=Leioceph-
alus; STENO=“Stenocercus”; TROPI=“Tropidurus”’;
URANO=Uranoscodon; CROTA=Crotaphytus;
GAMBE=Gambelia; OPLUR=Oplurus; CHALA
=Chalarodon; DIPSO=Dipsosaurus; BRACH=Bra-
chylophus; IGUAN=iguanines, excluding Dipso-
saurus and Brachylophus. Unordered transformations
are: 12, 34, 36, 37, 40.

MISCELLANEOUS PUBLICATIONS

54

Appendix 1 continued.

CHARACTER

27) 2 ee

a

Zz

q

a0

il

M203 G5 6

TAXON

1

1

Oo =0 e000" "0" 0" OO” 0 OF 0 0. 0 0

SPHEN

0 1

?

0F70" "07" 0" 0 0. 0 0r 0" 0 0 0: 8. 0 OO 0

SCLER

ied

Q
0-0, 0

O70. (0, 105 0) 0.05 Os 0 0. On 0 Oe 5 0

ANCES
PRISC

1-0 0358

1

tO <O gale Ope?

Le ten O00 <0), 1 30
Pel

0
0

AGAMI

oe GC Colo SoS Crt eS co
Bae eae A TH NO CO TO 4
COO On nA aA a TOO a AO
earn Oooconooe &
ee
aan tt Onoonooeo oe
maMaoinorinooocoooo°oce
eo .Se — Oo So oC oO oc, Qo 2 oc ©
CcoOe nA HTH TH OOO oO O&O
oooco~nxwn Oowrne oon o &
oooeoooojoo°ocncneo9o &
So oo Go. So OC Cro Ole Sc) ©
Se So: oC SO CG So SCO oO A om i -o
asHormamnmocooocnoocneoenat oO &
ee <> > oo
COOn nD AHA TH OO AO
cocoon n oooceontooe &
—srHooecnoococoeoeoceceo &
Sa SS CS CFO OO CFO co oc
Cone oooocoooeoeo }&
Ce Oe Pe ee)
eooooocoqcoocoqcocqc &
jaa)
CEEEEELECECEEL
P=
Bgees
Be E Seek e sae ode

Le O60
LT O30
1

1
1

OleeOre, ORWOr MO; 10400).0>. 0 0) OF OOO '0" °0.0
OF OOO 10-0 O10 0-0 600". 0 00 0
OO 0Y 00" 0207 0 OF 0 O07 0. 0 0. 0 0

02-07 10> 70
07 0 0" 0
2
1

1
1

SCELO

UROSA

UTA

0 0

0

0. On a
0

0° 0" 0-0) 0. * 0304.0
O° O70) 0, OF.40 207 O2).0,4,0

0 0..0 0, 04.0. 0,: 0

1
1
u

O05 a

1
1

PHRYN

0.0, 70

0 0 0. 0.0 0 ©

SANDL

1
1

0

1

?

PHYMA

CTENO
LIOLA

0:0: 0.10.0), 0. 10.05 05 0 2. 0 0. GO), 0:..0y O40

004.0. 40. On 20708220" OFZ) KOs O00

Og, 0-70)~ 10,40
Oy 0, 0R R00 (02 “OL 1058 OF 10% 405550. HO HOR TORSO M20

Le Oaee0
1
1

z
1

0 0 0" 0 0" 04.05 0
0. 0. 0 40. 0-00, (0.0

0 0 72
1
1
1
1

1
1

OF Oe “OPO: (0. O26

?

LEIOC

STENO
TROPI

0

0; MOstOn0- 0) 0.4105, O20) 0.0 0" 0 O00

00 080 SOO 0. 00-0. 0 0. 0 OO 0
OF0 0 50% 0 0

URANO

CROTA

0 0. O50

1
1

0

1

0.0. 0536

OnOF 40" 0 Os O05 s0

GAMBE

OPLUR

1

(ee

1

OF0e 200) 20.0. O:r0e 107 O80

OD ORO” 0 = 0.40,00:370

0

1

CHALA

On Oes 00720) 510: 0am On. Oar

DIPSO

eng

O® O70) 1.097042 04 OO NO20) 4020 + 1 oO ONO S70 0> OP VOetOm ae

OPPO Or 0) F0e OS ONO “0

BRACH

1

IGUAN

59

IGUANIAN LIZARD PHYLOGENY

Appendix 1 continued.

CHARACTER

4 4 4 4 4

7

6

2 Sy 4, 45

1

ooo 0 0

1

o 0 0

Oo 1

0

1

® 0.156

UROMA
LEIOL

PHYSI

CHAME

POLYC

0

2 0
Z

1
1

o 6 G0 OG 0 OO 0 @ 8 Oo O

ENYAL

1

O° 0,0 06 09 0-0 O09 GO GO GO

PRIST

u

7D O0050 0 0 GO GO GO 0

PARAA

ANOLE

ENYLD

BASIL

2 S0eOOre0

1

of 0 OG O O @ 0

1

oo O20 OO O @ 2B tf

1
1
1

Gg: 0 6 0

® O4;0-0
0 0 10750

1

1

0 0 @ GO O O 2

OF 10

1

0 O 0
0 ©

0
0

1

CORYT

LAEMA
PETRO

SCELO

UROSA

oo & OO

PHRYN

0

oC oO 0 0) a 0

SANDL

2

1

1
1

Oo oc HO oO 0 @ 1
1

1
1

Q. 0-6
))6§«0..0

1
1

PHYMA

CTENO

0 © C6 O00
0) sO O.Oy.0 120
co Oo @ 0 0
oO GO O0UCO

o Oo G&G OG @ 0

LEIOC

Dt

1

1
1

oo 0 0 0 0 &Y Oo 0
eo oo 0 & 0

1
1

0 2 *.6" 0
1
1
0

0
?

STENO
TROPI

oO o 0

1

2 0. 0

URANO

CROTA

0) 0

Ho oO Go GO Oo. & OG OO OO tI

1
1
?

1
0
0

OO 2 Oh OO OF GO 46

Omo 2 00 00 GO 0 tO OG GG Oo 0 8.0

0 0

GAMBE

OPLUR

O20" 2 Oe
1

1
1
1

bp OOO 0 Oo Oo O 0
0 0

1
1

Lt 2 Or

1

q 2, 0.6 por a Uo 6 0

CHALA

0

oo CG 0 0 GO ft

0

1

DIPSO

”

1

BRACH

1

IGUAN

MISCELLANEOUS PUBLICATIONS

56

Appendix 1 continued.

CHARACTER

a -& Gi A AS cS aS

5,3 a © 6
ain:

5

7

8 4-5

3 “OO: = "2. ~3 "4 "5

7

QO “1026

0 0 0 0OCert

TAXON

SPHEN

?

0 °oO "0°00 “0 0 °0 0 "OO 2

1

d,

SCLER

al

0 0 @°0 0 °O0 0° 0°02

oO %

?

ANCES

0
0

1
1
1

oo Oo 0-0 “O 0° OO. 0 90

1
1
1
1

o°O “0 0 OO 0 OO 0 0
O-o 0 O° 0 0 0 C0 0 0
Oo oO 0 °O 0°00 °C GO 0 G 0

1
1

1
?
1

LEIOL

0 "0 90
0 0-36

1
1

0 0) -0
0 2°0 QO © OC tac

1

PHYSI
CHAME

POLYC

0 1

0

1

o0 0 G0 0 0 0 @

% 1

1

oC. 0

ENYAL
PRIST

0 2 0 0 0 OO 3GeRe
2° 0 0 O00

1

0: O° CO. ¢ 30

Qf -2 11

1

?

0 0 0 °O -O “O S0mee

1
1
1
1

oo 0° 0. 0 0°O 8 0 0 CG OO

1
7 0 O

ENYLD

BASIL

Ll 0 °O 3Oes0¢
0 °0 "0
0 OF*¢

1
1

Q -0 @

"0 Oo Oo 0 O° O 0 0 9

1
1
1

?

oo
0. *0" ~0

cd -0- 0-6 0 0 0 0 0
no oo 0 'O0 0 0 0 CU 0

1) “O70
1

CORYT

?

0-0

LAEMA

Soaqeeoqqeqn oot eS eo owas
epi S) ere oe a ore eS Se SS SS
eee! Ge Se ee ee oe ose oe SY Oa ‘a
O'e SO Oo Oo Co So OS CG SO ao oC CO oO CCS
e160 "oe oo oS oe Oo Go oo oe co oo oo Cre
HAndntet ae wt Ooogoocooocoocoeceoeocoooe°s
eeooeoocooeoCceoqoutaqocooco Oo
aH oaota eS Ooocooooooqcocoqo cs
On nH COOH FH TH OOOH Fa FH DTH OOS
eeCceeceoecooocoqcoeonasocse
SooocoocoocecooqocoqooCcnoqlo
Saooeooeoeocoqcoqcooqooqcoo co >]
eeooeeocooocooomtnetoeooqcoqcco
Soe ooo 4] =| © COO SC aa OC Ce SS
aoa oso eS ocoeoeoeceooooqoocoo
Seeoooeeoooocooooeontaocse
Sooocoeoeoqoacooqcoococeqcoqooe
OooqocooocqcoeoqoCcoqoqcooqgooo
coocooonn nt oooaoaacocoeC eo
COCO CO On FA HF A DT OOH TH OC SO
Cn COC On a ae at TH DO OOCOCoCC SO
ae aos ato aS OSeSCeooae as Soo CS
Seq oro eoe sO CS So Oa aa Oo OC OC | =
Ono. Aso eve SkaHk {05
~ 4 Pe 4Zn645 Ze = Rag

PRO R eee Pope eee eee hae
POSSE Ae Oa oa e Doo eoa aS

IGUANIAN LIZARD PHYLOGENY 57

APPENDIX 2

Apomorphy lists for tree in Figure 9. See legend of Appendix 1 for abbreviations used. Asterisks note character
shifts that are of ambiguous placement. Characters from unpolarized or unordered transformations are noted by a
“U.” Daggers note other characters (not an exhaustive list) not used in the analysis.

TRANSFORMATION ANCESTRAL DERIVED
STEM SERIES CHARACTER CHARACTER
PRISC . 6 1 0
AGAMI U 32 0 1
UROMA 2) 0 1
U 32 0 1
Ui 36 1 0
65 0 1
LEIOL U 62 0 1
PHYSI U* 30 1 0
a7 1 0
CHAME 39 1 2
47 0 1
* 48 0 1
U 58 0 1
+supratemporal reduced to small splint not in contact with parietal (Rieppel, 1981).
+pterygoid fails to meet quadrate (Rieppel, 1981).
+zygodactylous feet.
POLYC =) 0 1
UU) 27 0 1
U 48 1 0
Dil 1 0
52 1 0
tacrocentric chromosomes acquired (Gorman et al., 1969).
tfourth toe reduced, equals third in length (Cope, 1900; Etheridge and de Queiroz, 1988).
ENYAL Ua 0 1
U 64 0 1
fthroat scales conical (Etheridge and de Queiroz, 1988).
PRIST ~~ as 1 0
+supradigital scales become transversely expanded, lamellar-like (Etheridge and de Queiroz,
1988).
+proximal subdigital scales of toes 1-3 become enlarged (Etheridge and de Queiroz, 1988).
PARAA tsexual dichromatism lost (Etheridge and de Queiroz, 1988).
ANOLE Usa, 0 1
uw as 1 0
U 21 1 2
U 2 0 1
Ui. 23 0 1
24 1 2
67 0 1
tdistal pad raised under phalanges 2 and 3 (Etheridge and de Queiroz, 1988).
ENYLD WU as 1 0
25 0 1
ynasal scale enlarged (Etheridge, 1969b).
BASIL U 44 1 0
WD 45 1 0
CORYT 13 0 1
UF 21 0 1
U_.3l 0 1
LAEMA 7 0 1
oe ge) 1 0

58 MISCELLANEOUS PUBLICATIONS

Appendix 2 continued.

TRANSFORMATION ANCESTRAL DERIVED

STEM SERIES CHARACTER CHARACTER
PETRO U.e3s 1 0

39 1 0

+neural spines shortened (Etheridge, 1964).

SCELO WU ~35 1 0

47 0 1
UROSA +secondary cusps of posterior marginal teeth reduced (Etheridge and de Queiroz, 1988).
UTA tdark axillary spot (Mittleman, 1942) (homologous with those in sand lizards?).
PHRYN 8 0 1

+skull and head scales strongly modified (Reeve, 1952; Presch, 1969).
+phalanges lost from digits 4 and 5 of manus (Etheridge and de Queiroz, 1988).

SANDL 66 0 1

tlabial scales elongated, obliquely oriented; lower jaw countersunk (Smith, 1946; Axtell,
1958).

PHYMA U 21 1 0

U 30 0 1

U 38 1 0

39 1 0

CTENO U 45 0 1

LIOLA Winey 0 1

U 36 1 0

LEIOC 10 0 1

U 32 0 1

tlong, free xiphisternal rods curve forward to underly xiphisternal ribs (Etheridge, 1966).
+postrostral scales lost (Etheridge, 1966).

STENO Us 23 1 0
+extensive transverse hemipenial musculature (Amold, 1984).
TROPI lingual coronoid process of dentary present (Frost, 1987).
+articular surface of humeral head elevated (Frost,1987).
URANO U 44 0 1
U 45 0 1
47 1 0
Wress 0 1
tgrebe-like toe fringes develop (Luke, 1986).
CROTA 1 0 1
41 0 1
GAMBE tclavicular fenestrae (Etheridge and de Queiroz, 1988).
tintermuscular dermal pit on posterior surface of thigh (weakly developed in many
Crotaphytus).
OPLUR U 38 1 2
39 i 0
CHALA 9 0 1
Ur 30 0 1
U 46 1 0
DIPSO 11 0 1
U 44 1 0
U 45 1 0
BRACH U 40 0 if
41 0 1
IGUAN 10 0 1
U 36 1 0
1 15 0 1
U3! 0 1

IGUANIAN LIZARD PHYLOGENY

Appendix 2 continued.

TRANSFORMATION ANCESTRAL DERIVED
STEM SERIES CHARACTER CHARACTER
. * 3 1 0
4 0 1
Ae 6 1 0
* 39 1 0
U 46 0 1
3 1 1 0
Ui 37 0 DZ
U 40 1 0
4 * 8 0 1
U 16 0 1
Wi al 0 1
U* 21 0 1
28 0 1
* 67 0 1
5 2 0 1
* 3 0 1
9 0 1
26 0 1
U* 30 0 1
U* 38 1 2
+39 0 1
* 41 0 1
eS) 0 1
U* 63 0 1
6 * 6 0 1
WaalS 0 1
U 20 1 0
U* 48 1 0
7 24 0 1
29 0 1
U* 30 0 1
U* 31 0 1
33 0 1
U 36 2, 1
U 38 1 2
ety Bie) 1 2
47 0 1
+division of mental scale (E. E. Williams, pers. comm.).
8 U* 34 0 3
39 0 1
41 0 1
U* 46 0) 1
* 56 0 1
9 U* 45 1 0
50 0) 1
10 el 0 1
14 0 1
U* 21 0 1
U 27 1 0
U 36 1 2
23 5)I 0 1
52 0 1
U 61 0 2

60 MISCELLANEOUS PUBLICATIONS

Appendix 2 continued.

TRANSFORMATION ANCESTRAL DERIVED
STEM SERIES CHARACTER CHARACTER

11 /

U 40

12 8

10

41

+sharp canthal ridge acquired (Etheridge and de Queiroz, 1988; Lang, 1989).
“Omer9

oN COO
SEP WR

5)

0
0
U 34 0

U_ 63 0

14 Uer21 1
15 Uirv9 0
0

RSPR OrFNR NKR

+dorsal scales enlarged relative to laterals (Larsen and Tanner, 1975).
16 U 35 0)
42 0
1 5) 0
9 0
+scleral ossicle 6 reduced or lost (de Queiroz, 1982).
trow of enlarged chinshields that increase in size posteriorly (Montanucci in Etheridge and
de Queiroz, 1988).
tanterior fibers of m. retractor lateralis anterior reflected outwards or posteriorly before
insertion (Arnold, 1984).
18 28
UN30

— — es pe

U 59
U 60
Ui 62
tscleral ossicle 8 reduced (de Queiroz, 198
19 33

ll el >

20

*
_
0°

Zl

eee ee, —e Me Me Se
ron

22 12
23

S

*
_
~

S

Ww

Nn —
SCOOFRPFORRKFrFHOOORF OF OF OCONOOrFCOCCOCSO

RFrRrROoOoNGONRF RFR ONK OF NRF

IGUANIAN LIZARD PHYLOGENY 61
Appendix 2 continued.

TRANSFORMATION ANCESTRAL DERIVED
STEM SERIES CHARACTER CHARACTER

24 eae

25 12

26

Zt

28 MS)

*
—
~
SBR Rr OrFrRrFOOCCOCOCOCOOCOOCORrRFOCcCoOrFr:®S

38
+females acquire gravid coloration (Medica et al, 1973) (also in some phrynosomatids [e.g.,
Holbrookia (Axtell and Wasserman, 1953), Petrosaurus mearnsi] and some tropidurids
[e.g., Tropidurus thoracicus (Dixon and Wright, 1975)]).
20

COCCOCORFROCO OR RP RP RP RP RP RrPREPNONFRrrRr OF

29

30

31

Bec. Cece, 3 "ey &
2 Ww
oo
OrPrOrRrFOOrFRCOOYF

FP OOrONR FOF FF OC

54
32 +frontals fused embryonically (Jollie, 1960).
+frontals constricted between the orbits (Estes et al., 1988) (modified in some taxa).
+broad frontal shelf underlying nasals (Estes et al., 1988).
tpostfrontal reduced (Estes et al., 1988; Presch, 1988).
+parietal foramen on frontoparietal suture (Estes et al., 1988) (modified in some taxa).
tiguanian brain morphology (Northcutt, 1978).
tm. intercostalis ventralis lost (Camp, 1923).
+tongue mucocytes mostly serous and sero-mucous (Gabe and Saint-Girons, 1969; Schwenk,
1988).

62 MISCELLANEOUS PUBLICATIONS

APPENDIX 3

Lists of changes within transformation series for tree in Figure 9. See legend of Appendix 1 for abbreviations
used. Asterisks note character shifts that are of ambiguous placement. Characters from unpolarized or unordered
transformations are noted by a “U.”

TRANSFORMATION CHANGED ALONG
SERIES From To STEM CONSISTENCY

1
20 0.500
2 =) 1.000
3

2 0.500
2 1.000

ns

UROMA 0.333

PRISC 0.333

LAEMA 0.250

PHRYN 0.333

LAEMA 0.167
10

LEIOC 0.600
11
DIPSO 0.500
12

22 0.667
13 *

PRIST 0.333

10 1.000
1 1.000
4 1.000

* * *
COOrCCCCOCOHCCOCCOCCCOCOONOKFCCCCCCCCOCOrorFrOCSCOOOroeorse
KEEP OPEB ESB RP OH ENE PEP BEEP WN OR PRP EP PRP PEP EPP OrOOr rE rrOoOrrOrF

—
Ww

ENYAL 0.167

IGUANIAN LIZARD PHYLOGENY

Appendix 3 continued.

TRANSFORMATION CHANGED ALONG

SERIES CONSISTENCY

SI
fe)
K
a |
°
N
Sy
is
K-4

*

SS SOS Sq Se) SSreey eye ne a — i SF al ea — aL EL ll I) ll

U18

ANOLE 0.200
U19 6 1.000

U20

21 0.333
U21

++ + & +

ANOLE 0.250
U22
ANOLE 0.500
U23

ANOLE 0.333
24

ANOLE 0.333
25
ENYLD 0.500
26 5 1.000

U27

POLYC 0.333
28
4 0.500
d 1.000
U30

PHYSI 0.167
U31 *

CORYT 0:333
U32

AGAMI 0.333
53

7 0.333
U34

WN RE Bee eee POR PE HP ee EE HE OORP PRE NFONNRFRFORP RF RFPNRFONOFF OF OCOFrCOFrOCS
N
Ww

8 1.000

64 MISCELLANEOUS PUBLICATIONS

Appendix 3 continued.

TRANSFORMATION CHANGED ALONG
SERIES FROM To STEM CONSISTENCY

U35

SCELO 0.250
U36

UROMA 0.286
U37

25 0.667
U38 ‘

PETRO 0.286
39

CHAME 0.222
U40

BRACH 0.500
41 *

CROTA 0.200
42
16 0.500
43 25 1.000

U44

BASIL 0.250
U45

OrrFROOCOCOFOCORP FP SE RF RFF FF rRPNORrFNOCCONFRFRFOONNONNONKF OCOF CON OFF O
~

DE > I SP > > a a ce ee ee ee oe ee ee ee ee ee ee ee 2 ee ee a ee

BASIL 0.167

IGUANIAN LIZARD PHYLOGENY

Appendix 3 continued.

TRANSFORMATION CHANGED ALONG
SERIES FROM To STEM CONSISTENCY

U46

CHALA 0.200
47

CHAME 0.200
U48

CHAME 0.200

23 1.000
9 1.000
POLYC 0.500

POLYC 0.333
18 1.000

0.333
20 1.000
8 1.000
PHYSI 0.500

CHAME 0.500

URANO 0.200
18 1.000

19 0.500
LEIOL 0.500
13 0.500
ENYAL 0.500
UROMA 0.500
66

SANDL 0.500
67 *

OO Ss BS SE BEB ND RE NR RK RK OO OR RF OR RP RE ROR RP OR RF OR RP RR OOF Or RF Or RF Or KF OF
No
Ww

Nn

lon

*
oqooooocoocoocoocoorocooonorrrOCO OFM COC CC OCOrF COFCO Or CCC OCOFrF Or OC OCOrFM COFCO OF Oo

ANOL 0.500

65

iit

716

|

62 3

iin

67

68.

69.

70.
fae
a2.

has

74.

72.
76.

rip

78.

ADs

80.

RECENT MIS

roe eS
An ecogeographic analysis of th / abn toa
1-75, 22 text-figures, 27 plates, “<< <7,

Internal oral features of larvae fi —
and ecological considerations.
Paper bound.

c, evolutionary
June 24, 1980.

The Eleutherodactylus of the A

: Leptodactyli-
dae). By John D. Lynch and Wi gust 29, 1980.
Paper bound. |
Sexual size differences in reptile ruary 27,1981
Paper bound.
Late Pleistocene herpetofaunas | 26 text-figures.

May 8, 1981. Paper bound.

Leptodactylid frogs of the ger
adjacent Colombia. By John D.

n Ecuador and
Paper bound.

Type and figured specimens of fossil vertebrates in the collection of The University of Kansas
Museum of Natural History. Part I. Fossil fishes. By H.-P. Schultze, J. D. Stewart, A. M. Neuner
and R. W. Coldiron. Pp. 1-53. October 6, 1982. Paper bound.

Relationships of pocket gophers of the genus Geomys from the central and northern Great Plains.
By Lawrence R. Heaney and Robert M. Timm. Pp. 1-59, 19 text-figures. June 1, 1983. Paper
bound.

The taxonomy and phylogenetic relationships of the hylid frog genus Stefania. By William E.
Duellman and Marinus S. Hoogmoed. Pp. 1-39, 30 text-figures. March 1, 1984. Paper bound.

Variation in clutch and litter size in New World reptiles. By Henry S. Fitch. Pp. 1-76, 15 text-
figures. May 24, 1985. Paper bound.

Type and figured specimens of fossil vertebrates in the collection of The University of Kansas
Museum of Natural History. Part II. Fossil amphibians and reptiles. By H.-P. Schultze, L. Hunt,
J. Chorn and A. M. Neuner. Pp. 1-66. December 3, 1985. Paper bound.

Type and figured specimens of fossil vertebrates in the collection of The University of Kansas
Museum of Natural History. Part III. Fossil birds. By John F. Neas and Marion Anne Jenkinson.
Pp. 1-14. February 5, 1986. Paper bound.

Type and figured specimens of fossil vertebrates in the collection of The University of Kansas
Museum of Natural History. Part IV. Fossil mammals. By Gregg E. Ostrander, Assefa Mebrate
and Robert W. Wilson. Pp. 1-83. November 21, 1986. Paper bound.

Phylogenetic studies of north american minnows, with emphasis on the genus Cyprinella
(Teleostei: Cypriniformes). By Richard L. Mayden. Pp. 1-189, 85 text-figures, 4 color plates.
June 1, 1989. Paper bound. ISBN: 0-89338—029-6.

;

Nise aes ait
itt eT

Ni
ih

wy Al
1 We ;