Eocene Lizards of the Clade Geiseltaliellus from Messel and Geiseltal, Germany, and the Early Radiation of Iguanidae (Reptilia: Squamata) Author(S): Krister T

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Eocene Lizards of the Clade Geiseltaliellus from Messel and Geiseltal, Germany, and the Early Radiation of Iguanidae (Reptilia: Squamata) Author(s): Krister T. Smith Source: Bulletin of the Peabody Museum of Natural History, 50(2):219-306. 2009. Published By: Peabody Museum of Natural History at Yale University DOI: http://dx.doi.org/10.3374/014.050.0201 URL: http://www.bioone.org/doi/full/10.3374/014.050.0201

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Eocene Lizards of the Clade Geiseltaliellus from Messel and Geiseltal, Germany, and the Early Radiation of Iguanidae (Reptilia: Squamata) Krister T. Smith

Abteilung Paläoanthropologie und Messelforschung, Senckenberg Research Institute and Natural History Museum, Senckenberganlage 25, 60325 Frankfurt am Main, Germany — email: [email protected]

Abstract The historical biogeography of the lizard clade Iguanidae is complicated. In addition to difficulties within the New World, where most of the more than 900 living species are found, two extant iguanid clades, Brachylophus and Oplurinae, occur well outside it. Moreover, there is a small set of Eocene species in Europe, most notably those placed in the genus Geiseltaliellus. To examine the relevance of Geiseltaliellus to iguanid biogeography, I redescribe several well-preserved specimens (nearly complete skeletons with epidermal scales) from the middle Eocene lake deposits of Messel, Germany. These specimens were previously referred to the type species, G. longicaudus, but comparison with the type material reveals differences that warrant specific distinction. Mes- sel Geiseltaliellus resembles extant Basiliscus in squamation and parietal growth. Phylogenetic analysis of morphological data using Bayesian and parsimony methods suggests the following about the evolution of pleurodont iguanians: (1) Iguanidae is monophyletic, its members united by unique features of the snout; (2) Iguanidae is divided into two major clades, one consisting of Polychrotinae* + Corytophaninae, Iguaninae + Hoplocercinae and Crotaphytinae (Clade A), the other of Phrynosomatinae, Tropidurinae* and Oplurinae (Clade B); (3) Polychrotinae* and Corytophaninae are sister taxa; and (4) Geiseltaliellus is on the stem of Corytophaninae. The presence of Geiseltaliellus in Europe during the warm, humid Eocene suggests dispersal from North America and a more northerly distribution of the corytophanine stem than the crown. Geiseltal- iellus represents a separate invasion by Iguanidae of the Old World and an evolutionary dead end. On the basis of its fossil record and modern distribution, Clade A is interpreted as ancestrally North American. Persistent conflict of morphological with molecular genetic data on iguanid relationships remains to be resolved.

Keywords Iguanidae, Corytophaninae, Messel, Geiseltal, historical biogeography, molecular-genetic.

Introduction of Gauthier et al. 1988 and Schulte et al. 2003; previously recognized, often Linnaean taxa whose The diverse clade Iguania comprises two major monophyly is in question—that is, they cannot subclades, Acrodonta and Iguanidae, which strongly be shown to be either monophyletic or have an evidently complicated biogeographic nonmonophyletic—are denoted with an asterisk.) history. Acrodonta (including Agamidae* and The Iguanidae is primarily a New World group Chamaeleonidae sensu Estes et al. 1988) is essen- (taxonomy of Iguanidae follows Schulte et al. tially an Old World clade, though in the green- 2003), diverse in both mainland North and South house of the Eocene at least one lineage America, which were isolated during most of the (Tinosaurus) found its way to North America and Cenozoic, and in the Caribbean. Furthermore, persevered until the close of the epoch (Hecht in Eocene fossils from Europe are referable to McGrew et al. 1959; Estes 1983a; Smith 2006a, Iguanidae (Kuhn 1944; Estes 1983a; Augé 1987; 2006b). (Here I follow the metataxon convention Rossmann 2000); species of the iguanine clade

Bulletin of the Peabody Museum of Natural History 50(2):219–306, October 2009. © 2009 Peabody Museum of Natural History, Yale University. All rights reserved. • http://www.peabody.yale.edu 220 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Brachylophus inhabit genetically related islands de Queiroz 1987; Lang 1989; Sites et al. 1996; in the South Pacific (e.g., Gibbons 1981; Pregill Frost 1992; Pregill 1992; McGuire 1996; Reeder and Steadman 2004); certain iguanines and and Wiens 1996; Titus and Frost 1996; Poe 1998, tropidurines are endemic to the Galápagos; and 2004; Wiens and Hollingsworth 2000; Frost et al. the clade Oplurinae is endemic to Madagascar 2001; Wiens and Etheridge 2003; Torres-Carvajal (Blanc 1965). et al. 2006; Torres-Carvajal and de Queiroz 2009) The existence of iguanids on Pacific islands or anatomical systems other than the skeleton may well be explained by sweepstakes dispersal (e.g., Renous-Lécuru and Jullien 1972; Peterson on favorable ocean currents (on Brachylophus, see 1984; Schwenk 1988). Macey et al. (1997) and Cogger 1974; Gibbons 1981; and Sites et al. 1996), Schulte et al. (1998, 2003) presented phylogenetic but the existence of an endemic clade on Mada- analyses of Iguania based on molecular-genetic gascar is a conundrum. Are oplurines relicts of a data. Divergent ingroup topologies have been previously much more widely distributed inferred in recent morphology-based phyloge- Iguanidae (or Iguania), whose range included netic studies of Iguania (Conrad and Norell 2007; Africa and Madagascar before the sundering of Conrad et al. 2007; Conrad 2008). It is minimally Gondwana in the Early Cretaceous (Jullien and fair to say that there is no consensus on morpho- Renous-Lécuru 1972; Estes 1983b; Williams 1988; logical evidence for even iguanid monophyly. Rossmann 2001)? Or did they arrive later from Hoffstetter had (1942) refuted all earlier refer- the south, perhaps through Antarctica before its ences to Iguanidae of European fossil taxa. Shortly refrigeration (Blanc 1982; Noonan and Chippin- thereafter, however, Kuhn (1944) briefly described dale 2006)? Or did they take a northerly route, Geiseltaliellus longicaudus from the middle into Eurasia during the greenhouse of the Eocene, Eocene lignite deposits of Geiseltal near Halle thence south through Africa (Boulenger 1918) or (Saale), Germany. The first mention of a speci- India (Rage 1996)? The presence of Paleogene men from what became the type locality seems to iguanids in Europe makes the last possibility plau- be that of Weigelt (1931:69): sible. A Holarctic origin of afrotherian mammals has also been proposed (Zack et al. 2005), several Bei weitem der wichtigste Fund ist eine außeror- dentlich langschwänzige Eidechse, deren Schwanz early Tertiary dispersal events between Africa and fast dreimal so lang ist wie Kopf und Rumpf zusam- Asia are documented (e.g., Gheerbrant and Rage men. Das Tier ist vollständig mit Schädel und allen 2006; Smith et al. 2008), and the fossil record of Extremitäten erhalten und zeigt in der Rückenge- the African Paleogene is notoriously poor (e.g., gend wie an den Extremitäten Reste einer sehr feinen Estes 1983a). Beschuppung. Es handelt sich ganz offensichtlich um einen hochspezialisierten Baumbewohner, dessen The European iguanids are therefore quite rel- Bearbeitung sehr viel Interessantes verspricht. evant to our understanding of iguanian historical biogeography, for the nature of their relationships Since the original description of Geiseltaliel- to other iguanids will affect what importance we lus, several new skeletons of the taxon have been ascribe to this particular incursion into the Old discovered in the early middle Eocene lake World. Yet the relationships of iguanids to each deposits of Messel near Darmstadt, Germany. other remain problematic quite apart from those Rossmann (2000) has already provided partial of the fossil forms. Knowledge of Iguanidae was descriptions of these specimens. Additionally, a greatly advanced by the work of Etheridge (1959, significant number of fragmentary remains sim- 1964, 1965, 1966, 1967), who promulgated a set of ilar to Geiseltaliellus have been referred to that eight informal groupings (here called Corytophan- genus, including three new species from the inae, Crotaphytinae, Hoplocercinae, Iguaninae, Eocene of France and Belgium (Augé 1990a, Oplurinae, Phrynosomatinae, Polychrotinae* 1990b, 2003, 2005). In this contribution I focus and Tropidurinae*, after Schulte et al. 2003). on the osteology of the skull, forelimb and pec- Etheridge and de Queiroz (1988) and Frost and toral girdle of the well-preserved Messel speci- Etheridge (1989) conducted the first quantitative mens of Geiseltaliellus and describe also the phylogenetic analyses of Iguanidae, and many squamation of these. I compare them to closely other workers have examined in detail the rela- related forms from the European Paleogene and tionships within major iguanid clades (e.g., provide data on the ontogeny of species of Eocene Lizards of the Clade Geiseltaliellus • Smith 221

Geiseltaliellus. I then present a new phylogenetic analysis of Iguania, incorporating many new morphological characters and including Messel specimens of Geiseltaliellus, and discuss the implications of the inferred phylogenetic structure of Iguanidae for the historical biogeography and early evolution of the clade.

Abbreviations

The following are the institutional abbreviations used herein: CAS, California Academy of Sci- ences, San Francisco, California, USA; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA; FMNH, Field Museum of Natural History, Chicago, Illinois, USA; GM, Geiseltalmuseum, Martin-Luther-Universität, Halle-Wittenberg, Germany; HLMD-Me, Messel Collection, Hessisches Landesmuseum, Darm- stadt, Germany; MCZ, Museum of Comparative Figure 1. Germany, showing the locations of Messel Zoology, Harvard University, Cambridge, Mass- and Geiseltal. achussetts, USA; SMF, Herpetology Collection, and SMF ME, Messel Collection, Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany; TMM, Texas Memorial from approximately 47 Ma, based on their height Museum, The University of Texas, Austin, Texas above the base of the Messel Formation and aver- USA; UF, Florida Museum of Natural History, age sedimentation rates (Franzen 2005). The Gainesville, Florida, USA; YPM-PU, Princeton Messel locality belongs to, and most recently Collection, YPM-VP, Division of Vertebrate Pale- according to Franzen (2005) defines, European ontology, and YPM-VZ, Division of Vertebrate biochronologic level MP (Mammifères Paléogène) Zoology, Peabody Museum of Natural History, 11. Anoxia at the floor of the lake hindered scav- Yale University, New Haven, Connecticut, USA. enging and resulted in considerable soft-tissue preservation (Franzen 1985; Wuttke 1983). Geologic Settings Also counted among the world’s Lagerstätten is Geiseltal, near Halle an der Saale, in eastern The Messel fossil Lagerstätte (common anglo- Germany (see Figure 1), where middle to late phonic shorthand for Konservat-Lagerstätte, or Eocene lignites were mined for 70 years. Swampy “conservation deposit”) near Frankfurt am Main, conditions are usually poorly suited to the preser- Germany, is a UNESCO World Heritage site vation of vertebrate fossils, because of elevated renowned for its exceptionally preserved speci- concentrations of humic acids, but the ground- mens (Figure 1). These derive from finely lami- water feeding this sump passed through the Tri- nated sediments laid down in a small lake basin, assic limestones of the Muschelkalk, picking up one of several in the area thought to have arisen bicarbonate that buffered the humic acids and by phreatomagmatic eruptions (Harms 2002; allowed biogenic apatite (bone) to be preserved Harms et al. 2003), in which groundwater is (Krumbiegel 1977). Soft-tissue preservation is explosively vaporized by contact with near-sur- also common there, but unlike in Messel, very face magma. The origin of the nearby Eckfelder fine-scale structures (cellular and intracellular) basin has been dated to 44.3 Ϯ 0.4 Ma (Mertz et al. also are preserved (e.g., Voigt 1988). Epidermal 2000). The event at Messel occurred somewhat scales are represented as original keratin, although earlier, 47.8 Ϯ 0.2 Ma (Franzen 2005). The pro- other structures, like muscle tissue, have been ductively fossiliferous layers are estimated to be replaced by silica (Voigt 1937). 222 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Systematic Paleontology Geiseltaliellus maarius sp. nov. Figures 2–11 CLASS Reptilia Linnaeus, 1758 ORDER Squamata Oppel, 1811 Geiseltaliellus longicaudus Rossmann, 2000 [in part]. Geiseltaliellus longicaudus Augé, 2005 [in part]. SUBORDER Iguania Cope, 1864 FAMILY Iguanidae Bell, 1825 Holotype. HLMD-Me 10207 (nearly complete skeleton). GENUS Geiseltaliellus Kuhn, 1944 Topotypical locality. HLMD pit 14, middle Messel Formation, Type species. Geiseltaliellus longicaudus Kuhn, 1944. Germany.

Referred species. Geiseltaliellus grisolli Augé, 2005, G. laman- Paratypes. SMF ME 2, 1769, 2684 and 2938. dini (Filhol, 1877), G. maarius sp. nov. Age and distribution. The species is currently known only from Age and distribution. Early to late Eocene of Europe (MP 7–19: the middle Messel Formation, middle Eocene (MP 11). The Augé 2005). specimens derive from excavation pits that span a stratigraphic range of about 24 m, from the so-called alpha to M horizons Emended diagnosis. An iguanid lizard differing from Lae- (Schaal et al. 1987; Hesse and Habersetzer 1993). At an average manctus, Crotaphytinae, Phrynosomatinae (except Phryno- sedimentation rate in this part of the formation of 0.15 mm/Kyr soma), Tropidurinae* (except Uranoscodon and Plica), (Goth 1990), the maximal temporal separation of the specimens Dipsosaurus and Anolis in having a moderately broad, paris roughly 160 Kyr. Although the hypodigm cannot be seen as allel-sided nasal process of the premaxilla. It differs from all representing a paleopopulation, this temporal span is short other iguanids except Corytophaninae, most Anolis and compared with the inferred age of many living species (e.g., Tor- some members of Iguaninae in having a Y-shaped parietal res-Carvajal and de Queiroz 2009). table; from all other iguanids except Corytophaninae, Cro- taphytinae, Chalarodon and “sand lizards” (Phrynosomati- Etymology. From Maar (German, referring to a lake nestled in nae) in having a highly reduced or absent postfrontal; from the crater formed by a phreatomagnatic eruption), originally all other iguanids except Corytophaninae, Anolis, many from mare (Latin, sea). members of Oplurinae and Leiocephalus in having (variably) a dorsally expanded posterior ramus of postorbital; Diagnosis. Differs from G. longicaudus Kuhn, 1944 in lacking a from Iguaninae, Hoplocercinae, Polychrotinae*, Coryto- strong ventral expansion of the coronoid and in having a clav- phaninae (except some Basiliscus) and Tropidurinae* icle whose ventromedial expansion is most extensive at the level in having a briefly closed but unfused Meckelian groove; of the clavicular fenestra rather than dorsolateral to it. Differs from Polychrotinae*, Iguaninae, Crotaphytus, Oplurinae, from G. grisolli Augé, 2005 in having a broader nasal spine of the Tropidurinae* (except Plica) and Phrynosomatinae in premaxilla and a weaker and more rounded subdental shelf having a ventromedially fenestrated clavicle; from most anteriorly on the dentary. Differs from G. lamandini (Filhol, iguanids except Corytophaninae, Crotaphytinae, Enyalioides 1877) in having a more restricted Meckelian groove, a more del- heterolepis and “sand lizards” in having a ratio of leg length icate anteromedial process of the coronoid, and distinct antero- to snout–vent length greater than 0.67; and from at least lateral and posterolateral processes of the coronoid. some species in all other major clades of Iguanidae in having a ratio of tail length to snout–vent length of more than Comments. Rossmann (2000) previously referred the speci- 2.0. Differs from Corytophaninae in lacking a medial pari- mens from Messel (HLMD-Me 10207 and SMF ME 2, 1769, etal blade and anterior premaxillary foramina. Posterolat- 2684 and 2938) to G. longicaudus. Differences in the coronoid eral process of coronoid extends to level of posterior margin and clavicle, among features that can be compared, distinguish of anterior surangular foramen (autapomorphy). Antero- the genotypic Geiseltal specimens from those at Messel. Varia- lateral process of coronoid at least weakly developed tion observed among the Messel specimens does not extend to (autapomorphy). these features, as discussed in detail below, which leads me to separate the Messel specimens as a different species. Other differences may obtain as well, for significant portions of the skull Comments. All known species are from Europe. Except of specimens of G. longicaudus are obscure. for Geiseltaliellus lamandini (see below), they probably constitute a clade, although showing this by apomorphy is difficult because of the nature of some of the material. Description Rossmann (2000) synonymized Geiseltaliellus louisi Augé, 1990a with G. longicaudus, and also included under this state of the specimens name the Messel specimens. Augé (2005) provisionally Some brief comments on the state of preservation of the speci- accepted this synonymy. Augé (2005) placed the probably mens follow. Letters “a” and “b” refer to part and counterpart, late Eocene Plesiolacerta lamandini (Filhol, 1877) in respectively. Geiseltaliellus. This last species lacks the autapomorphies of Geiseltaliellus given above, and its reference to Geiseltaliel- HLMD-Me 10207. This specimen is a nearly complete, articulated lus is not yet well justified. skeleton. In certain places, pieces of bone (e.g., the anterior portions Eocene Lizards of the Clade Geiseltaliellus • Smith 223 ani; j, jugal; l, lacrimal; m, maxilla; n, nasal; p, parietal; lar foramen; pt, pterygoid; q, quadrate; rapr, retroartic- , Interpretation. Hatching indicates areas where bones are only B , Photograph. A : a, angular; aiaf, anterior inferior alveolar foramen (of maxilla); ar, articular; bs, basisphenoid; Abbreviations sp. nov. (HLMD-Me 10207, holotype). Geiseltaliellus maarius Dorsal view of skull Figure 2. Figure ular process; sa, surangular; snaf, subnarial arterial foramen; sq, squamosal; st, supratemporal. Scale bar is 10 mm. cn, coronoid; crtv, crista transversalis; d, dentary; ec, ectopterygoid; ep, epipterygoid; f, frontal; fct, foramen chorda tymp pf, parietal foramen; pl, palatine; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; psaf, posterior surangu seen as impressions. Dentary tooth impressions are lightly outlined. 224 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 , Interpretation. The B d, dentary; ec, ectopterygoid; eo, exoccipital; j, jugal; ocess; sa, surangular; sot, sphenoöccipital tubercle; sp, m, premaxilla; pmhf, posterior mylohyoid foramen; po, : a, angular; aiaf, anterior inferior alveolar foramen (of mandible); Abbreviations , Photograph (specimen coated in ammonium chloride). A sp. nov. (SMF ME 2938a, paratype). Geiseltaliellus maarius Ventral view of skull Figure 3. Figure specimen was damaged between the drawing of illustration and taking photograph. splenial; sq, squamosal; stpr, supratemporal process; v, vomer; X, foramen for vagus nerve. Scale bar is 10 mm. apr, angular process; bo, basioccipital; bppr, basipterygoid processes; bs, basisphenoid; cb, ceratobranchial I; cn, coronoid; l, lacrimal; m, maxilla; Mgr, Meckelian groove; oc, occipital condyle; or, recess; par, prearticular; pl, palatine; p postorbital; ppr, paroccipital process; pt, pterygoid; pt-qr, quadrate ramus of q, quadrate; rapr, retroarticular pr Eocene Lizards of the Clade Geiseltaliellus • Smith 225

Figure 4. Right dorsolateral view of skull of Geiseltaliellus maarius sp. nov. (SMF ME 1769a+b, paratype). A, Photograph. B, Interpretation. Abbreviations: a, angular; asaf, anterior surangular foramen; cb, ceratobranchial I; cn, coronoid; d; dentary; f, frontal; j, jugal; l, lacrimal; m, maxilla; n, nasal; os, scleral ossicle; p, parietal; pf, parietal foramen; pl, palatine; pm, premaxilla; po, postorbital; prf, prefrontal; q, quadrate; sa, surangular; sq, squamosal; st, supratemporal; stpr, supratemporal process. Scale bar is 10 mm. 226 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

SMF ME 2938. This specimen is a complete but strongly plastically deformed skeleton in ventral view. Most of the skull is readily interpretable, but clarification of the braincase could aid the reader. The left exoccipital portion of the occipital condyle has been separated from the basioccipital portion and rotated, such that the condylar portion faces upward (toward the observer). As with the separation of the parabasisphenoid and basioccipital, it seems likely that the ease of separation of the elements of the condyle indicates that they are not fully fused with one another, pointing to an immature individual.

Privately held specimen. This specimen, the second largest of all referred to Geiseltaliellus maarius, consists of a complete Figure 5. Oblique, right-lateral photograph of the skeleton preserved as part and counterpart. The specimen is parietal of Geiseltaliellus maarius sp. nov. (HLMD-Me currently in a private collection, but plaster casts, which are the 10207, holotype). Note the raised, sinuous anterior basis for observations on the sternum (Figure 8), were kindly portion of the median crest and the broken area imme- made available to me by Dr. Torsten Rossmann (then: Tech- diately posterior to it, which together constitute nische Universität, Darmstadt, Germany). The casts are now approximately one-third the length of the crest. catalogued as SMF ME A332. bones of the skull roof of the dentaries) apparently remained stuck to the counterpart, Premaxilla. The azygous premaxilla consists of a ventral tooth- which was not collected; however, impressions of these parts bearing portion and the posterodorsally directed nasal process. remain (hatched areas in Figure 2B). As in all specimens described The anterior margin of the ventral portion is gently convex and here, the skeleton was almost completely extricated from the its anterior surface is smooth (Figure 2). There are no anterior matrix and embedded in resin. Little soft tissue is preserved. premaxillary foramina in HLMD-Me 10207, which would have passed twigs of the ethmoidal nerve (cf. Oelrich 1956); in SMF SMF ME 2. A nearly complete skeleton preserved in left dorso- ME 1769 the base of the nasal process superficially seems to be lateral view. The tail and much of the abdomen are missing and notched weakly on the right side, but the notch represents a parts of the skeleton are strongly plastically deformed. Scale small fracture that passes through the premaxilla. Thus, there is impressions are preserved along the back, as are portions of the no evidence of variation in the relationship between the nerve inscriptional ribs. and the bone. The nasal process is moderately broad (Figure 2) and rises steeply (Figure 4), although a precise angle to the hor- SMF ME 1769. This specimen is a nearly complete skeleton, izontal is difficult to give because of distortion. The lateral mar- missing only the distal portion of the tail. The body of the skele- gins of the process are parallel so far as they are preserved ton and the head formed part and counterpart, respectively. The (Figure 2), but the process presumably tapered toward its distal head was later separated and glued to the part; because a space end, which is nowhere recognizable, if indeed it is preserved. remains between the head and the surface of the resin, its ven- The depth of the nasal process on the mid-line cannot be deter- tral surface is unobservable. The specimen has suffered consid- mined with available material. erable plastic deformation to certain parts. No specimen presents a view of the palatal surface of the premaxilla, so determining the number of premaxillary tooth SMF ME 2684. This headless specimen is everywhere strongly positions is difficult. Two teeth partially protrude from the pre- plastically deformed and shows little morphology. It is the maxilla of HLMD-Me 10207 (Figure 2) and Rossmann (2000) largest of all the specimens. Most of the specimen is on the part suggested six or eight teeth were present. The two preserved (SMF ME 2684a), a small portion on the counterpart (SMF ME teeth probably represent the first tooth pair and would have 2684b). This specimen is quite unlike the others in attitude. In been separated by a mid-line tooth; the length of the margins lat- the other specimens, the body and tail are relatively straight; the eral to the preserved teeth suggest that two more positions were legs are adducted and the knees somewhat flexed, such that probably present on each side, for a total tooth count of seven. the feet closely approach one another near the tail; the arms are Rossmann (2000) indicated that eight are present in the holo- adducted and directed posteriorly, such that the hands overlie type of Geiseltaliellus longicaudus, which I could not confirm one another (or nearly so) beneath the abdomen. Additionally, because the element is so poorly preserved. the head is present. In SMF ME 2684, in contrast, one leg is On the right side of the lateral process of the premaxilla in adducted, the other abducted; likewise the arms (with both SMF ME 1769, a portion of the dorsolaterally situated articular appendages on each side of the body held in the same position). facet for the premaxillary process of the maxilla is visible. The The vertebral column is multiply bent and the head is missing. medial portion of the maxillary articulation is covered in all These differences clearly indicate a different taphonomic his- specimens. tory for SMF ME 2684 than for the other specimens. It is associated with the other specimens on the basis of general body Nasal. Portions of the nasals are preserved in multiple speci- proportions and the iguanian morphology of the undersurface mens. In all they are smooth externally, without adornment of the vertebrae. or rugosity. In HLMD-Me 10207 the anterior and posterior Eocene Lizards of the Clade Geiseltaliellus • Smith 227 portions of the right nasal are visible (Figure 2). The anteriormost preserved piece bears a longitudinal impression on its medial side, where it would have articulated on the ventrolateral surface of the nasal process of the premaxilla. Posterolat- erally the nasal is in broad contact with the prefrontal, separating the anterolateral spine of the frontal from the maxilla. The nasals seem to have contacted one another on the mid-line posteriorly, but the extent to which their mutual suture was invaded by the distal tip of the nasal process of the Figure 6. Scaled outlines comparing the parietal in premaxilla is unknown. The medial and lateral margins of the different specimens of Geiseltaliellus maarius sp. nov. bone abruptly turn toward one another posteriorly, such that A, SMF ME 2a (paratype). B, HLMD-Me 10207 (holo- the bone tapers to a blunt point, which on the right side in type). C, SMF ME 1769a+b (paratype). During later HLMD-Me 10207 is fully preserved (Figure 2), but whose ontogeny the length of the parietal increases relatively shape is only indicated by the conformation of the nasal facet more rapidly than its width. on the left side of the frontal in SMF ME 1769 (Figure 4).

Septomaxilla. Not discernible. the anterior end of the median scale immediately posterior to it, which is slightly larger and ovoid in outline. Bordering the orbital Prefrontal. HLMD-Me 10207 (Figure 2) and SMF ME 2 and margin on either side is an arcade of impressions for periorbital 1769 (Figure 4) each present the pyramidal prefrontal in roughly scales (i.e., the supraorbital semicircles), which are neither more lateral view. At the apex of the pyramid is a tear-drop shaped nor less pronounced than the median two; the periorbital arcades rugosity adorned with tiny grooves, the prefrontal boss, which are not precisely symmetrical (cf. Rossmann 2000). A small, lon- is drawn out a short distance posteriorly along the lateral edge gitudinal groove behind the last preserved median scale impres- of the frontal process. The boss is distinctly weaker and less sion indicates that the latter was separated from the interparietal rugose in SMF ME 1769 (Figure 4) than in SMF ME 2 or scale by a pair of medially apposed scales. Obvious impressions HLMD-Me 10207 (Figure 2), even though the first specimen is for epidermal scales are not present in SMF ME 1769 (Figure 4), distinctly larger than the other two (Appendix 1). The frontal although they cannot be fully ruled out in SMF ME 2, where the process terminates near midorbit. It has a smooth, nearly flat bone surface is covered by a thin black film. dorsal surface and a sagittally concave, ventrally and slightly lat- The posterior two-thirds of the frontal is undistorted in erally directed orbital surface, both of which taper toward the HLMD-Me 10207 (Figure 2) and SMF ME 2. Longitudinally it distal end of the process. The medial surface of the frontal is gently dorsally convex. In HLMD-Me 10207 the dorsal sur- process is only visible on the left prefrontal in SMF ME 1769 face is weakly tranversely concave, but the concavity is strongly (Figure 4), and only partially at that; it is marked by a longitu- expressed in SMF ME 2. The lateral portion of the frontal is dinal concavity that presumably corresponds to a convexity in upturned because of breakage in SMF ME 1769 and a weak but the prefrontal scar on the frontal, the portion of the medial sur- distinct supraorbital flange (Smith 2009) projects laterally face ventral to the groove being slightly greater than the portion beyond the crista cranii. When viewed as laterally as possible, dorsal to it. The orbital surface arches ventrally anteriorly to the left side of the frontal in SMF ME 2 seems to have an even form the broad antorbital surface, whose medial edge and exact stronger supraorbital flange, also suggested by the great width relations with the palatine cannot be reconstructed because of of the frontal at midorbit relative to its width at the frontopari- deformation and breakage. etal suture in the specimen (0.32; cf. Rossmann 2000). The pari- Ventral to the prefrontal boss, the prefrontal articulates etal foramen is contiguous with the frontoparietal suture, with the lacrimal. The lacrimal seems to articulate flushly on incising both bones in approximately equal measure in SMF the lateral edge of the prefrontal (Figure 4), but crushing and ME 1769, but the frontal more deeply in HLMD-Me 10207. distortion of this region in all specimens render this observa- The lateral margins of the frontal have a slight kink near tion uncertain. Near the junction of the lacrimal and pre- midorbit, which marks the posteriormost extent of the frontal frontal, the orbital rim is deeply impressed, forming a groove process of the prefrontal (Figure 2). The anterior margin of the ventral to the prefrontal boss. More ventrally still, and medial frontal table is W-shaped, with posteriorly extensive facets for to the lacrimal, the edge of the prefrontal is excavated for the the nasals. The median and lateral prongs are about equal in lacrimal ducts. length. The anterolateral face of the prefrontal is covered by the facial process of the maxilla in all specimens, and the paranasal Postfrontal. There is no bone that would correspond to a post- excavation on the medial surface is nowhere visible. frontal in the normal iguanid position on the posterodorsal margin of the orbit (Rossmann 2000). However, there is a small, Frontal. The azygous frontal is constricted between the orbits elongate sliver of bone on the orbital face of the completely (Figures 2 and 4), as in other iguanians. The dorsal surface of the exposed dorsal ramus of the right postorbital in HLMD-Me bone is generally smooth, but in HLMD-Me 10207 the poste- 10207 (Figure 2). The external surface of this sliver is flush with rior half shows a network of shallow grooves that presumably that of the postorbital, which bounds it on most sides except at mark the boundaries of epidermal scales (Figure 2). A median its dorsal extremity, where it may have contacted the frontal. scale with a nearly rounded anterior margin and straight, oblique The sliver is sutured to the postorbital, not the product of a posterolateral margins is found at the transverse level of the dis- break. On topographical grounds the structure in HLMD-Me tal end of the frontal processes of the prefrontals. It barely touches 10207 is best interpreted as a highly reduced postfrontal, as is 228 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

most preserved portion of the facet shows a longitudinal concavity. The anterior margin of the dorsal ramus is only weakly concave. The lateral surface of the bone is adorned with an apically weakly rugose tubercle just posterior to the orbital margin and at a height approximately one-fourth of the way from the basal edge. This tubercle is drawn out dorsally and slightly anteriorly, converging with the orbital margin and contributing to a well-developed orbital face. The tubercle may have been developed more dorsally in SMF ME 2, but the dorsal ramus is covered by the frontoparietal corner in that specimen. The tubercle probably served for the attachment of supraorbital fascia (cf. Oelrich 1956). Although the distal end of the dorsal ramus of the postorbital is fully exposed in SMF ME 2938 (Fig- ure 3), the extent to which it might have underlapped the frontoparietal corner is unclear because of poor preservation. The ventral edge of the bone is very gently concave in lateral view in HLMD-Me 10207 (Figure 2), where it is more noticeable on the left side, and in SMF ME 2. In HLMD-Me 10207 the jugal articulates on this edge for about half its length. This might, however, be an artifactual underestimate caused by the somewhat downward trajectory of the jugal into the slab in that specimen; in SMF ME 2, the jugal remains fully articulated with the ventral edge of the postorbital and extends along it for 78% of its total length (see also SMF ME 1769, Figure 4). The posterior ramus of the bone differs among the specimens. In HLMD-Me 10207 it is dorsoventrally tall, its dorsal margin convex (Figure 2). Only at the anterior junction with the dorsal ramus does this margin become concave. The dorsal margin is also somewhat convex in SMF ME 2, but less strongly. In SMF ME 2938, finally, the posterior ramus is dorsoventrally short and evinces a more or less straight dorsal margin (Figure 3).

Parietal. This azygous bone articulates anteriorly with the frontal at a broad transverse suture along which lies the parietal foramen (Figures 2 and 4). It is roughly rectangular in overall form, but the sides and posterior margin are deeply embayed. The parietal Figure 7. Elements of the pectoral girdle of Geiseltal- table—that portion of the dorsal surface that is not a site of origin iellus maarius sp. nov. (SMF ME 2938a, paratype). A, for the external adductor or pseudotemporalis musculature—is Photograph (specimen coated in ammonium chlo- triangular and only developed anteriorly, forming but a small ride). B, Interpretation. The specimen is highly plasti- fraction of the dorsal surface of the bone in the larger specimens cally deformed, and the cartilaginous sternum is either (Figures 2 and 4). The table is bounded by the sharp adductor not preserved or is not distinguishable as such. Note crests, which project slightly dorsally above the level of the table. the separation of the epiphysis from the diaphysis of These crests converge posteriorly and behind their junction are the humerus. Abbreviations: acf, anterior coracoid fen- continued as a thin median crest, no more than 1 mm in height, estra; cc, coracoid; cl, clavicle; hu, humerus; hu-ep, which continues posteriorly to the end of the bone. The anterior proximal humeral epiphysis; ic, interclavicle; sc, quarter of this median crest in HLMD-Me 10207 is slightly taller scapula; ssc, suprascapula. Scale bar is 5 mm. than the posterior two thirds (Figure 5). This anterior fourth is also finely sinuous in dorsal view (Figure 2). A small portion of the crest (between the posterior edge of the sinuous part of the crest occasionally encountered in other iguanids (see Appendix 3). and the level of the arrow in Figure 2B) seems to have been bro- This region is not exposed in any other specimen, so it is not ken, implying that the sinuous part of the median crest could have clear whether the structure in HLMD-Me 10207 is common or encompassed up to a full third of its length. The parietal crest has a rare variant. a sinuous portion only in HLMD-Me 10207; this could indicate intraspecific variation, because the crest is also very well preserved Postorbital. This triangular bone is best seen in HLMD-Me in SMF ME 1769, which shows no indication of a sinuous, 10207 (Figure 2) and SMF ME 2. The bone articulates expanded anterior portion (Figure 4). anteroventrally on the orbital face of the temporal ramus of the A median parietal blade, as found in living corytopha- jugal. The anterior ramus of the postorbital is relatively short; nines, is not seen in any specimen, regardless of size. Four spec- anteriorly, the facet for the jugal is developed ventrolaterally, imens show the parietal in dorsal view (HLMD-Me 10207, but it rotates posteriorly, coming to face ventrally. The anterior- SMF ME 2 and 1769, and the privately held specimen). Even if Eocene Lizards of the Clade Geiseltaliellus • Smith 229

Figure 8. Sternal apparatus of Geiseltaliellus maarius sp. nov. Drawings of the sternal region of casts (SMF ME A332) of a specimen in a private collection. Cartilaginous areas are stippled. The first vertebra is denoted by an ital- icized x, succeeding vertebrae are numbered x+1, x+2 and so on, through x+6. Similarly, the dorsal rib attaching to vertebra x and its ventral counterpart is denoted by a sans serif x, with succeeding ribs numbered x+1, x+2 and so on, through x+6. I, II, first and second digits of left manus. A, Left dorsolateral view. B, Right ventrolateral view. The narrowness of the distal ends of posterior, left dorsal ribs in A, when compared to B, results from the three-dimensional nature of the preservation: the ribs plunge into the slab and thus progressively less is visible distally. Abbrevi- ations: hu, humerus; ra, radius; stn, sternum; ul, ulna; x, rib of vertebra x; xip, xiphisternum. Scale bar is 5 mm.

a parietal blade were present but absolutely sexually dimorphic adductor crests, but posteriorly they are broadly convex, except (developed to any degree only in males), the probability that all immediately adjacent to the median crest. The supratemporal known specimens are females, given a 1:1 sex ratio, is (0.5)4 ϭ processes seem to be relatively short. The precise connections of 0.0625, which, if not quite significant, is also not very likely. It is parietal to the squamosal, the quadrate and the bones of the nevertheless tempting to view the sinuous, slightly expanded chondrocranium could not be reconstructed. portion of the crest in HLMD-Me 10207 as an incipient blade. What soft tissues might have been associated with the sinuous Supratemporal. The supratemporal is best seen in HLMD-Me portion are unknown. 10207, where it extends along the posterolateral margin of the The shape of the parietal and the relative size of the table supratemporal fossa (Figure 2). Its exterior surface is flush with differs among the specimens. In the smallest of them, SMF ME that of the parietal. It extends as far anteriorly as the level of the 2, the parietal is nearly square (with equal maximum length and midpoint of the median parietal crest. Its medial suture with the width) and the median crest constitutes less than half of the parietal describes a weakly undulating line. mid-line length of the bone (Figure 6A). In the larger specimens HLMD-Me 10207 and SMF ME 1769, in contrast, the parietal Maxilla. The premaxillary process of this element, which forms is much longer than wide and the median crest constitutes the floor of the external naris, articulates on the dorsolateral sur- approximately two-thirds of the mid-line length (Figure 6B and face of the lateral process of the premaxilla (to an artifactually C, respectively). Because these specimens differ in overall size, exaggerated extent in Figure 4). The dorsal surface of the pre- they presumably represent different developmental stages. maxillary process is pierced two foramina; a small groove Clearly, the length of the parietal increases more rapidly than extends anteromedially nearly to the anterior edge of the bone its width during ontogeny, although a component of intraspe- from the more anteriorly situated of the two foramina, the sub- cific variation in the mature size of the parietal table or its narial arterial foramen, whereas the more posteriorly situated growth trajectory cannot be excluded. anterior inferior alveolar foramen opens more dorsally at the The restriction of the parietal table to the anterior portion anterior base of the facial process of the maxilla. There is also an of the bone results in broad supratemporal fossae (Figures 2 and artifactual groove extending anteromedially from the anterior 4), the surfaces where the adductor musculature originates. inferior alveolar foramen in HLMD-Me 10207 (Figure 2). The Anteriorly they are concave in cross section orthogonal to the dorsal surface of the premaxillary process is otherwise smooth 230 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 and flat, and slopes slightly ventrolaterally; it is separated from Jugal. The anterior ramus of this element forms the ventral mar- the dorsomedial surface of the palatal flange of the maxilla by a gin of the orbit. Its long articulation with the maxilla, so far as weak ridge, the crista transversalis (Smith 2006a; see Figure 2). preserved, was described above. The bone extends a short dis- The dorsal surface of the palatal shelf is poorly exposed in all tance posteriorly beyond the posteriormost exposure of the max- specimens. This shelf, seen in medial aspect in SMF ME 2938, illa (Figures 2 and 4). The anterior ramus transitions posteriorly shows at its anterior end a facet, which could be a recess for the into the temporal or ascending ramus, the two rami forming an duct to Jacobson’s organ, but more likely constitutes an articu- obtuse angle (122° in HLMD-Me 10207, 114° in SMF ME 1769). lation surface for the vomer, and at approximately mid-length Although a quadratojugal tubercle is absent in all specimens, the a weak, medial expansion, the flange on which the maxillary strength of the angle of the jugal differss among the specimens, process of the palatine articulates (Figure 3). being stronger (more angular) in SMF ME 1769 (Figure 4) and The broad, smooth facial process of the maxilla rises nearly especially SMF ME 2938 (Figure 3), and very weak in HLMD- vertically at the posterior end of the premaxillary process (Fig- Me 10207 (Figure 2) and SMF ME 2. The posteroventral margin ure 2). Its lateral surface is pierced by one (Figures 3 and 2) or of the temporal ramus is only very weakly concave, whereas the two (Figure 2) rows of labial foramina. In HLMD-Me 10207 the anterodorsal margin has a strong posterior inflection where it first row occurs a little more than 1 mm above the ventral mar- meets the postorbital. The temporal ramus tapers distally and gin of the maxilla and the second row about 1 mm above the terminates in a blunt tip, especially visible in SMF ME 2, which first. Distally, the facial process in this specimen curves slightly inserts for a short distance between the postorbital and the ante- medially to contribute to the dorsal surface of the snout. Con- rior tip of the squamosal. The lateral surface of the bone is tact between the facial process and prefrontal is extensive, but in smooth, but is pierced by a curved row of foramina that parallels all specimens damage or deformation along the suture is exten- the orbital margin. These foramina number six on the left side of sive enough to preclude the examination of details. The suture HLMD-Me 10207, at least five on the right side of SMF ME 1769, line between the jugal and the posterior remnant of the facial and six or seven on the left side of SMF ME 2. In SMF ME 2, the process is distinct in all specimens (Figures 2, 3 and 4). From first five foramina are attended by somewhat anteriorly trending the ventral margin of the jugal the suture line rises first grooves, whereas in SMF ME 1769 and HLMD-Me 10207 the anterodorsally before extending more or less anteriorly. Thus, foramina exit laterally. In SMF ME 1769, the first foramen is the the suture line is dorsally convex, more abruptly so in SMF ME largest, whereas these foramina are of approximately equal size 2 and 1769 (Figure 4) than in SMF ME 2938 (Figure 3) in SMF ME 2 and HLMD-Me 10207. or HLMD-Me 10207 (Figure 2). The suture line continues smoothly onto the lacrimal (Figure 3). Scleral ossicles. Probable scleral elements are preserved in the The left maxilla of SMF ME 2938 has 20 clear teeth or right orbit of SMF ME 1769, lying atop the palatine (Figure 4), tooth spaces; there is additionally room anterior to the first pre- and in the left orbit of SMF ME 2. They are approximately rec- served tooth for about two or three more (Figure 3), for a max- tangular, with rounded corners. The total number of ossicles in illary tooth count of about 22 to 23. The right maxilla of SMF each orbit and their pattern of overlap cannot be determined. ME 1769 has 18 teeth, with additional room for about one posteriorly and two or three anteriorly, for a maxillary tooth count Squamosal. This element is particularly variable among the of 21 or 22. Even an approximate tooth count is unobtainable specimens. Its main rod-like body is dorsally arched and oval in in HLMD-Me 10207. Teeth are moderately high-crowned cross section (Figures 2 and 3). Toward its posterior end, where beneath the anterior base of the facial process, but crown height it articulates with the quadrate and the supratemporal process declines in the posterior one-third of the tooth row (Figures 3 of the parietal, the squamosal expands. A dorsal process, pres- and 4). Anterior teeth are unicuspid. In SMF ME 2938, the 13th ent in all specimens, runs along the supratemporal process. This maxillary tooth (assuming two anteriorly) is the first in the den- process is very strong in HLMD-Me 10207 (Figure 2), moder- tal arcade to show distinct mesial and distal accessory cusps, ate in SMF ME 2938 (about as long as the main rod of the bone and they are weak (Figure 3); the 10th tooth shows a weak dis- is tall; see Figure 3) and weak in SMF ME 1769 (Figure 4). The tal “shoulder” that presages the accessory cusp. In SMF ME thickened posterior margin of the bone is straight in HLMD-Me 1769 the ninth tooth (assuming two anteriorly) evinces weak 10207 (Figure 2), but convex in SMF ME 2938 (Figure 3) and mesial and distal shoulders, and while breakage to more ante- probably so in SMF ME 1769 (Figure 4). A distinct facet for the rior teeth erased crown morphology, it is unlikely that more quadrate could not be found, but the ventral portion of the anterior teeth had accessory cusps; the first distinct accessory squamosal is not well exposed in any specimen. The ventral cusps, separated from the central cusps by labial grooves, are on process seems to be stronger in SMF ME 2 and 2938 than in the 10th tooth. The posterior teeth are tricuspid, with parallel- HLMD-Me 10207 (cf. Figures 2 and 3), but this seeming differ- sided to very weakly tapering crowns. ence could be attributable to poor exposure in the last.

Lacrimal. The lacrimal is at least partially preserved anterior to Quadrate. In most specimens the quadrate remains in articu- the jugal on at least one side in all specimens, but it is best seen lation with the mandible. The hemispherical lateral portion of in SMF ME 1769 (Figure 4). Its posterodorsal edge forms a part the ventral condyle is exposed in SMF ME 1769 (Figure 4). of the orbital rim that is posteriorly continuous with the part The bone texture on the articular surface does not differ on formed by the jugal (see below). The anterodorsal portion of superficial examination from that of the remainder of the the lacrimal is impressed and the indentation continues for a bone. The edge of the articular surface and the synovial joint short distance onto the prefrontal, as discussed above. As indi- capsule is marked by a sharp inward step, dorsal to which the cated by SMF ME 2, the lacrimal formed the lateral margin of lateral concha begins to expand. This posteriorly concave con- the lacrimal foramen, as in most other squamates. cha grows in extent dorsally (Figure 2) and reaches a height Eocene Lizards of the Clade Geiseltaliellus • Smith 231

Figure 9. The right manus of Geiseltaliellus maarius sp. nov. (HLMD-Me 10207, holotype) in dorsal view. A, Photograph. B, Interpretation. Abbreviations: I–V, first through fifth digits; 1–5, first through fifth distal carpals; lc, lateral centrale; pi, pisiform; ra, radius; rae, radiale; ul, ulna; ule, ulnare. Scale bar is 5 mm. approximately equal to that of the main body of the bone (Fig- Palatine. The palatines have been strongly displaced in SMF ME ures 2 and 4). The edge of the lateral concha, the tympanic 2938 and have come to overlap one another partly, the left above crest, is straight in lateral view and slightly thickened (Figure (ventral to) the right (Figure 3). Both palatines are strongly dis- 2). The dorsal margin of the lateral concha is nearly horizon- rupted by what is probably the right prefrontal. The right palatal, as seen in SMF ME 2. It is also slightly swollen and arches tine might remain in articulation with the maxilla, but the bone posteriorly; the swollen portion is separated by a notch from is deformed enough that it is difficult to tell. The left palatine is the hemispherical dorsal (cephalic) condyle of the bone. The forked anteriorly, which probably marks the posterior margin thick posterior crest is nearly straight except near its dorsal of the left fenestra exochoanalis. and ventral ends. In HLMD-Me 10207 there is a small aper- Posteriorly, and only clearly on the left side, the palatine ture at mid-height on the concha just lateral to the posterior contacts the pterygoid (contra Rossmann 2000), although the crest (Figure 2), but it seems to be artifactual; its presence or lateral part of their mutual suture is difficult to follow (Figure 3). absence, however, could not be ascertained in other speci- The palatines do not seem to have borne teeth, but their poor mens. The right quadrate of SMF ME 2938 indicates the pres- state of preservation requires that this observation be treated ence of a medial concha that is more extensive dorsally than with some circumspection. ventrally (Figure 3). The medial concha is presumed to be covered in HLMD-Me 10207. Pterygoid. The broad anterior part of the pterygoid articulated anteriorly with the palatine and laterally with the ectopterygoid, bones of the palate although the latter articulation is not preserved in these specimens. Pterygoid teeth are visible on the ventral surface of the Vomer. The only specimen in which the palate is exposed is bone near its medial edge in SMF ME 2b and 2938 (contra Ross- SMF ME 2938 (Figure 3). Even here, a vomer could not be iden- mann 2000). In SMF ME 2938, the left pterygoid has two dis- tified on the basis of morphology. tinct teeth, the posterior of which is approximately 2 mm 232 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 anterior to the anterior margin of the left basipterygoid process ner of the basioccipital lies the sphenoöccipital tubercle, which (Figure 3). The larger of these teeth, the posterior one, is much seems to be formed primarily of the basioccipital. There are two smaller than any of the visible teeth of the jaws. The absence of main ridges on the tubercle in ventral view: the thicker and teeth on the right pterygoid in this specimen does not seem to apparently longer one extends anteromedially from the apex of be artifactual. The two teeth visible in SMF ME 2b belong to the tubercle, and the less prominent one extends essentially one pterygoid, and the other could not be identified. medially from the apex; a recess is developed between them. At The pterygoid has a mobile medial articulation with the the apex of the tubercle is a small, shallow depression that may basipterygoid process of the basisphenoid at the junction once have been filled by a piece of calcified cartilage (cf. Oelrich between the anterior part and the quadrate ramus. Strongly 1956). The portion of the occipital condyle formed by the medially projecting but anteroposteriorly short flanges under- basioccipital is swollen and projects ventrally below the main lapped the anterior portion of the basipterygoid processes ventral plate of the bone. Because of a very subtle inhomogene- (Figure 3). The quadrate ramus then runs posterolaterally ity of hue and texture, the surface of the condyle does not appear toward the ventral condyle of the quadrate. This ramus is to be as well ossified as the remainder of the bone. The posterior intensely plastically deformed in SMF ME 2938. It is approxi- margin of the condyle is straight in ventral view. mately 1.5 mm tall and mediolaterally thickest along its ventral margin (Figure 3A). It tapers slightly near its distal tip and then Proötic. This portion of the chondrocranium is poorly exposed terminates bluntly (Figure 3). in all specimens. In HLMD-Me 10207 (Figure 2), the piece of bone disrupting the left side of the parietal posterior to the prob- Ectopterygoid. The ectopterygoid is visible in dorsal and slightly able remnants of the basisphenoid may represent portions of it. posterior view in HLMD-Me 10207 (Figure 2). The lateral, triangular articular surface for the jugal is weakly concave and Otoccipital. The exoccipital part of the unit, including the lateral smooth. The posterolateral process is a dorsoventrally tall struc- portion of the occipital condyle, is exposed on the left side of ture and consists of a sharp posterior crest that connects the SMF ME 2938 (Figure 3). The condylar face of the exoccipital dorsal and ventral corners of the process, the ventral one being is exposed to the observer. The condylar portion of this bone is the stronger. The main body of the bone has concave anterior difficult to distinguish on the right side, but large portions of and posterior margins, and part of the posterior articulation the paroccipital process are visible. This process extends pos- facet for the pterygoid. The posterior concavity contributes to terolaterally from the basioccipital toward the cephalic condyle the coronoid recess and has a sharp dorsal edge. The anterolat- of the quadrate and seems to have had a posterior longitudinal eral process of the bone is distinctly longer than the posterolat- concavity. Medially the process is continuous with the remain- eral process (Figure 3). der of the basioccipital. A tendril of bone projecting anteriorly from the base of the paroccipital process marks the lateral bones of the braincase boundary of the occipital recess. Posterior to it is a tiny foramen, probably for the vagus (X) nerve. And posterior, finally, to Orbitosphenoid. Not identified. this foramen is a small, sediment-covered circular depression of uncertain significance; if in fact the sediment masks a hole, Epipterygoid. The distal end of this very slender, rod-like ele- the latter could be a hypoglossal (XI) foramen. ment is preserved in HLMD-Me 10207 (Figure 2). Supraoccipital. Not identified. Basisphenoid. In SMF ME 2938 this element is separated from the basioccipital (Figure 3). The paired basipterygoid processes Stapes. Not identified. extend anterolaterally at an angle of 35° from the mid-line (Fig- ure 3), but the dorsoventral component of their orientation can- bones of the mandible not be reconstructed. The length of each process, from base to tip, is only slightly shorter than that of the ventral mid-line Dentary. This is a robust element attaining considerable height length of the basisphenoid. Each process is expanded distally. posteriorly. The approximate ratio of height at the posterior end In ventral view the basisphenoid is constricted, being narrow- of the tooth row to tooth row length is 0.30 in SMF ME 1769 and est close to the base of the basipterygoid processes. I could not 0.29 in the smaller SMF ME 2938. Anteriorly (from the trans- discern the parasphenoid process. The posterior margin of the verse level of teeth 4 through 13), the exterior surface is divided basisphenoid is concave, with thin posterolateral processes into a dorsal, vertical portion corresponding to the parapet of the extending posterolaterally toward the sphenoöccipital tubercles jaw and an oblique portion that meet at a rather sharp angle. Pos- (Figure 3). The posterior margin is undamaged, so the small, teriorly, this angle decreases and, except for a slight lateral bulge median, semicircular invagination is probably not artifactual. immediately below the tooth row and a medial inflection near the The basisphenoid is not well preserved in any other ventral edge, the exterior surface of the posterior portion of the specimen, but in HLMD-Me 10207 (Figure 2) the left lateral dentary forms a nearly flat, ventrolaterally directed surface. Ante- surface of the parietal, just behind the epipterygoid, was proba- riorly, there is a set of colinear labial foramina just dorsal to the bly disrupted by the processus alaris and crista sellaris of the angulation. There are six in SMF ME 2; they become larger pos- basisphenoid. teriorly, with the ultimate foramen (beneath the 14th tooth) distinctly larger, and the last three attended by posteriorly directed Basioccipital. The ventral surface of the basioccipital is visible grooves. Six labial foramina are also present in SMF ME 2938 in SMF ME 2938 (Figure 3). It is roughly triangular, with a (Figure 3), which are set in weak depressions of small diameter; slightly convex anterior margin. On the right anterolateral cor- grooves are not developed. Five foramina are present in the right Eocene Lizards of the Clade Geiseltaliellus • Smith 233 dentary of SMF ME 1769 (Figure 4), the anterior-most of which transverse level of the apex of the coronoid). Posterodorsally, the is slit-like and scarcely visible; these are gradually larger posteri- splenial is in contact with the prearticular. An anterior mylohy- orly and the last two are accompanied by weak posterior grooves. oid foramen could not be identified and presumably was in the The anterior tip of the dentary curves weakly medially position of one of the artifactual holes that disrupt the splenial. toward the symphsis, which is an anteroposteriorly elongate, weakly convex oval incised ventrally at its sagittal midpoint by Coronoid. The coronally concave medial surface of the tip of the Meckelian groove (Figure 3). The supra-Meckelian lip the coronoid is only visible on the right side of HLMD-Me (Bhullar and Smith 2008) forms a thick, medially convex ridge 10207 (Figure 2). In medial aspect, the tip would have formed beneath the anterior five or six teeth. It tapers in dorsoventral a blunt peak with a nearly vertical posterior margin and an ante- height posteriorly, and its medial face becomes progressively rior margin inclined at roughly 45° to the horizontal. SMF ME flatter, forming a sharp dorsal and ventral edge. A subdental 2938 preserves most of the medial face of the coronoid (Figure shelf is present on the dorsal surface of the supra-Meckelian lip 3). It is arch-shaped, with large anteromedial and posteromedial beneath the first six or seven teeth, but it becomes obliterated processes. The posterior margin of the posteromedial process posteriorly and the lip flattens. Posteriorly from about the eighth seems to be deflected distally toward its tip, but this could be tooth, the height of the medial face of the supra-Meckelian lip artifactual. This process overlaps the prearticular and surangu- remains nearly constant, decreasing only beneath the last three lar. The anteromedial process extends anteroventrally just ven- or four teeth. The Meckelian groove is mediolaterally broad tral to an exposed portion of the supra-Meckelian lip, just anteriorly and faces ventrally. The infra-Meckelian lip grows in reaching the level of the antepenultimate tooth. The ventral prominence posteriorly, extending medially and finally slightly margin of this process seems to overlap the splenial medially. dorsally to completely close the Meckelian groove beneath the Its edge is jagged, possibly an artifact. eighth and ninth teeth. Thereafter it rotates rapidly ventrally to The coronoid sits atop the dorsal margin of the surangu- form a sharp-edged, medially directed flange bracing the sple- lar and overlaps both surangular and dentary laterally (Figures nial from beneath. 2 and 4). As noted above, a short anterolateral process extended Posterolaterally, the lateral surface of the dentary could along the dorsolateral surface of the dentary (Figure 4), but its have been overlapped by a small anterolateral process of the length and morphology are poorly constrained. A tapering pos- coronoid (Figure 4). Unfortunately, this portion of the coro- terolateral process extends posteriorly along the surangular, noid is broken in SMF ME 2, but a weak articulation facet (?) on reaching the transverse level of the posterior margin of the ante- the dentary suggests that this process did not extend as far anterior surangular foramen (Figure 3). The ventrolateral margin riorly as the posterior end of the tooth row. of the coronoid between these two processes is straight to In SMF ME 2938 the right dentary has 20 tooth positions slightly convex. (Figure 3). SMF ME 2 has 24 teeth in the right dentary. There might be weak mesial and distal shoulders on the ninth tooth in Angular. The medial exposure of this element is fusiform (Fig- SMF ME 2938, but the first distinct distal accessory cusp is on ure 3). The anterior point, wedged between the ventral margin the 11th (10th unknown), where the mesial cusp remains a of the splenial and infra-Meckelian lip of the dentary, extends shoulder. Both accessory cusps are well developed on the 13th well anterior to the level of the coronoid apex, reaching the tooth (12th unknown). In SMF ME 2, a weak but distinct mesial transverse level of the antepenultimate dentary tooth. The pos- accessory cusp is present on the eight tooth, but the first clear terior mylohyoid foramen is found on the medial face of the distal cusp is on the ninth (the mesial margin of which is bone near the midpoint of its length, close to the transverse level obscured); both mesial and distal cusps are present on the 10th. of the coronoid apex. Posterior to this foramen the angular is The anteriormost two teeth are distinctly low-crowned, bounded dorsally by the prearticular. but crown height increases rapidly posteriorly until the fifth. The angular attains significant exposure in lateral view The crowns of anterior teeth also curve somewhat lingually, (Figures 2, 3 and 4). Its dorsal border with the surangular forms whereas the crowns of teeth in the posterior two-thirds of the a straight, horizontal line (Figures 3 and 4), curving abruptly tooth row are straight. The teeth are closely spaced. ventrally near its posterior terminus (Figure 2). It is most anteriorly extensive (more so in SMF ME 1769 than in SMF ME 2938), where it occupies a portion of the space bounded by the Splenial. In only one specimen, SMF ME 2938, is the splenial concave posterior margin of the dentary. At the triple junction well exposed. It extends anteriorly to the level of the 11th tooth formed between the angular, surangular and dentary in SMF from anterior (Figure 3). The anterior inferior alveolar foramen ME 1769 and HLMD-Me 10207, the first two bones are slightly is an anteroposteriorly elongate opening located at the transverse separated by a small, triangular projection of the dentary, which level of the boundary between teeth 13 and 14. The foramen does is absent in SMF ME 2938. not seem to be fully enclosed in the splenial, but it is possible that the splenial does enclose it beneath the supra-Meckelian lip (inte- Articular complex. The surangular forms the lateral wall of the rior to the mandible). The splenial increases in height posteri- Meckelian fossa. Anteriorly it maintains sutural contact with the orly until close to the posterior margin of the tooth row. Its dentary, coronoid and angular, as described above. The anterior posterodorsal margin is difficult to follow because of breakage. surangular foramen is on the dorsal margin of the surangular, As it appears, however, it becomes separated dorsally from the medial to the posterolateral process of the coronoid (Figure 4). dentary by the intervening anteromedial process of the coronoid. The posterior surangular foramen is on the dorsolateral face of Its ventral border is much clearer, separated from the dentary the bone, 1 to 2 mm anterior to the glenoid fossa (Figure 2). beneath the last three tooth positions by the anterior process of The prearticular forms the ventromedial margin of the the angular. The splenial continues posteriorly to about the mid- Meckelian fossa. Ventrally it is sutured with the angular (Figure point of the medial exposure of the angular (i.e., close to the 3), and anteriorly and anteroventrally to the splenial. There is in 234 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Figure 10. Body squamation of Geiseltaliellus maarius sp. nov. A, Scales of nape (SMF ME 2a, paratype), with detail in inset. B, Scales of abdomen (SMF ME 2b, paratype). Scale bar is 5 mm. the smallest specimen (SMF ME 2938) a well-developed angular just lateral to this crest and posterior to the crest bounding the process at about the transverse level of the articular fossa (which articular fossa (Figure 2). The posterior end of the process may receives the ventral condyle of the quadrate), indicating that the have been expanded slightly into a tubercle (Figures 2 and 3). process was well developed even early in ontogeny, but its trajec- In dorsal view the articular fossa itself is not obviously tory cannot be reconstructed because of plastic deformation. The divisible into medial and lateral facets (Figure 2), as might be retroarticular process is well exposed in only one specimen, expected if the ventral condyle of the quadrate were so divided. HLMD Me-10207 (Figure 2). Its posterior and medial portions It is a transverse depression confluent with the face of the suran- are obscured by the right squamosal and a mass of bone that gular laterally and bounded anteriorly and posteriorly by raised might represent the quadrate, respectively. It looks like an elon- crests. The posterior crest is thin; the anterior one swells and gate triangle, with the narrow base anterior. It is shallowly concave rises dorsomedially to form a strong protuberance, whose apex in transverse and sagittal cross section, bounded medially by the is marked by a facet. A weak ridge descends this protuberance so-called medial crest. The foramen chorda tympani is developed anteriorly before smoothly joining the surangular. Eocene Lizards of the Clade Geiseltaliellus • Smith 235

Figure 11. Manual squamation of Geiseltaliellus maarius sp. nov. (SMF ME 2684a, paratype), in palmar view. A, Photograph. B, Interpretation. Bone boundaries are difficult to discriminate because of plastic deformation; dashed lines indicate uncertain boundaries. Epidermal scales are stippled. Note the differences in squamation among wrist, palm and subdigital areas. Abbreviations: I–V, first through fifth digits. Scale bar is 5 mm. hyoid elements along its anterior margin, apparently tapering in cross-sectional thickness posteriorly. The right lateral process seems to have Ceratobranchial I. This is the only osseous element in the squa- been artifactually rotated posteriorly, so the angle formed mate hyoid apparatus (Cope 1892) and cartilaginous elements between the left and posterior processes, approximately 63°, is or their impressions could not be distinguished in any speci- taken to be characteristic. men. The elements is best preserved in SMF ME 2938 (Figure Anteriorly, the ventral surface of the posterior process 3). They are slender, rod-shaped bones a little more than 6 mm bears a strong median keel, which tapers posteriorly in medio- long and 0.3 to 0.4 mm in diameter which, as preserved, extend lateral thickness and is flush anteriorly with the thick leading from about the level of the apex of the coronoid and curve pos- margins of the lateral processes. The keel separates a pair of terolaterally. The opposite sense of their curvature (bilateral impressions nestled between the bases of the posterior and lat- symmetry) suggests that it is natural. Expansion at the anterior eral processes. As these impressions shallow posteriorly, the keel tip on the right side may be artifactual. diminishes and finally disappears about 1.5 mm along the length of the posterior process. Beyond the keel, the ventral sur- pectoral girdle and sternal elements face of the process is smoothly convex in cross section. Anteri- orly it is approximately 0.8 mm in width and tapers fairly Interclavicle. The interclavicle is well preserved in ventral view uniformly toward its terminus. The two offsets observed along in SMF ME 2938 (Figure 7; contra Rossmann 2000). This its length are probably due to postmortem distortion. arrow-shaped element is approximately 8 mm long and consists of paired lateral processes and a long posterior process; Clavicle. The elongate clavicle is a comma-shaped element (Fig- anteriorly there is only a weak bump. The ends of lateral ure 7). Its dorsal portion is rod-like, but ventromedially it processes are covered; the right process, of which more is visi- expands in anteroposterior dimensions. The posterior margin ble, was at least 2.0 mm long. The left lateral process is thickest of this expansion is very thin. In the center of the expanded part 236 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Table 1. Lengths of elements of the manus in Geiseltal- Sternum and sternal ribs. The sternum and relations of the ster- iellus maarius sp. nov., based on the right manus of nal ribs are distinct only in the privately held specimen (Figure HLMD-Me 10207 (holotype). Elements numbered I 8A, B). (In SMF ME 2 and 2684 cartilaginous ventral ribs are through V for digits and metacarpals (MC). All lengths visible and, posteriorly, are clearly in contact with the ossified in millimeters. dorsal ribs, but they complexly overlie one another in the region of the sternum such that the details of their connections cannot be elucidated.) The right arm overlies the right dorsal rib series, Element Length which, at least in the cast, cannot be traced; two digits of the left hand also underlie the sternal area (Figure 8B; they were not MC I 2.4 reproduced in Figure 8A). I.1 2.9 Absolute positional information about which ribs attach I.2 1.8 to which vertebrae is unavailable for two reasons. First, the anterior cervical vertebrae are obscure. Second, connections MC II 3.5 between right ventral and dorsal ribs could not be discerned, II.1 2.1 and the left sternal ribs seem to have detached from their dor- II.2 3.0 sal counterparts, presumably during decay (see Figure 8). In II.3 2.3 squamates with four sternal ribs these attach to presacral verte- MC III 4.3 brae 9 to 12 (Hoffstetter and Gasc 1969). The sternum looks like an amorphous plate to which III.1 2.3 several ventral ribs attach posteriorly and posterolaterally (Fig- III.2 2.1 ure 8). There is no indication of a median sternal fontanelle; the III.3 3.7 presence of a small one cannot be excluded if some plastic defor- III.4 1.8 mation is taken into account, but the existence of a large MC IV 3.9 fontanelle, as seen in many tropidurines and phrynosomatines, is unlikely. The interclavicle could not be identified in this spec- IV.1 2.2 imen. The inscriptional ribs in many places look “jointed,” as if IV.2 2.0 broken in many places perpendicular to the shaft. “Jointing” of IV.3 2.0 the cartilaginous ribs commonly occurs during maceration in IV.4 3.1 water (Smith pers. obs.) and is presumably simply caused by the IV.4 1.7 decay process. The right side of the sternum is better preserved than the left. It shows four sternal ribs as well as one rib attach- MC V 2.0 ing to the xiphisternum. The xiphisternum additionally bears a V.1 1.8 short posteromedial process. It is not inconceivable that this V.2 3.6 process represents a severed additional xiphisternal rib, but V.3 1.6 because it tapers distally it could be merely a free process, as seen in certain other iguanids (Etheridge 1965). Thus, on the basis of this single specimen, one may attribute to this specimen a there is a small fenestra with smooth margins (Figures 2A and 5(4+1) rib pattern (see Hoffstetter and Gasc 1969). 7), which in SMF ME 2938 seems to be incomplete medially (Figure 7). This incompleteness, if real, could relate to the rela- axial skeleton tively small size of the specimen or could represent individual Rossmann (2000) has provided a description of the vertebral variation. column. Here, I seek only to quantify the proportions. Proportions. The tail is long in relation to the dimensions of the Scapulocoracoid. Various portions of the scapulocoracoid are body (Rossmann 2000), though this is difficult to quantify for visible in different specimens. In SMF ME 2938 the clavicle body size in some specimens because of the severing of either overlies the anterior portion of this element (Figure 7). The bone the distal tail (SMF ME 2, 1769) or the head (SMF ME 2684). thins in the region of an anterior coracoid fenestra. The ventral The tail length (TL) to snout–vent length (SVL) ratio in speci- border of the coracoid is irregular and is poorly ossified. The mens in which it could be measured was 2.0 (SMF ME 2938) glenoid fossa is not visible because of the articulated proximal and 2.1 (HLMD-Me 10207); in the privately held specimen, humeral epiphysis. There is probably a scapulocoracoid fenes- TL/SVL ϭ 3.2 (see Appendix 1). Clearly, tail length and tra beneath the expanded ventromedial end of the clavicle. More snout–vent length are less precisely measured in a specimen dorsally, at the horizontal level of the scapular foramen, the without soft tissue, but these should be fairly close estimates. scapula also seems to thin anteriorly, but the actual anterior margin of the bone could not be examined in any specimen. appendicular skeleton The posterior margin of the scapula is weakly posteriorly con- Rossmann (2000) provided a general description of the arm of cave. Its dorsal margin is rough. the species, which will not be supplemented here.

Suprascapula. A patch of rough, apparently mineralized tissue Manus. The manus is best preserved on the right side of is found dorsal and posterior to the scapula in SMF ME 2938 HLMD-Me 10207 (Figure 9), where it is seen in plantar view. (Figure 7), which possibly represents part of the suprascapula The radiale and the facet on its dorsomedial surface for the artic- (cf. Rossmann 2000). ulation of the radius are clearly seen, the radius having been Eocene Lizards of the Clade Geiseltaliellus • Smith 237

Table 2. Osteological differences among specimens of Geiseltaliellus maarius sp. nov. Abbreviations: HLMD- Me, Messel Collection, Hessisches Landesmuseum, Darmstadt, Germany; SMF ME, Messel Collection, Senck- enberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany; SVL, snout–vent length.

Osteological characteristic HLMD-Me 10207 SMF ME 2 SMF ME 1769 SMF ME 2938

Development of prefrontal boss Strong Strong Weak ? Degree of frontal concavity Weak Strong Weak–absent ? Presence of frontal sculpture Present Absent? Absent ? Location of postorbital tubercle Ventral Middle? ? ? Convexity of posterior ramus of postorbital Strong Weak ? Absent Sinuousness of median parietal crest Present Absent? Absent ? Length of medial parietal crest Long Middle Long ? Maxillary labial foramina Two rows One row One row One row Curvature of jugal–maxillary suture Gradual Abrupt Abrupt Gradual Angle of jugal Weak Weak Strong Very strong Grooves for jugal lateral foramina Absent Present Absent ? Size of jugal lateral foramina Equal Equal Anterior largest ? Squamosal, dorsal process Strong ? Weak? Moderate Squamosal, posterior margin Straight ? Convex? Convex Posteroventral corner of dentary ? Sharp Sharp Blunt Dentary, projection at Present Present Present Absent angular–surangular junction Angular, anterior extent ? Greater Greater Lesser Tail length:SVL ratio 2.1 ? ? 2.0 Leg length:SVL ratio 0.60 0.75 0.76 0.69 slightly disarticulated. A relatively large, triangular lateral cen- from the proximal end of the humerus to the distal end of trale follows postaxially. The ulnare, however, is difficult to dis- metacarpal III) to leg length (LL, defined here as the distance tinguish clearly. Postaxially is the pisiform. The first distal carpal from the proximal end of the femur to the distal end of is also difficult to distinguish. The small second and third distal metatarsal IV) ranges from 0.47 in SMF ME 1769 to 0.56 in carpals follow it postaxially. The fourth distal carpal—the largest HLMD-Me 10207 (see Appendix 1). Although exact measure- in the series, it has an expanded proximal, preaxial corner—is ments cannot be given for SMF ME 2684, the arm is also quite the most readily identified. It has a convex proximal surface short in that specimen. and the plantar surface is traversed by a transverse groove (Fig- Several authors (e.g., Estes 1983a; Rossmann 2000) have ure 9). Rossmann (2000) mistook the groove for an articulation remarked on relative limb length, particularly leg length (see surface and identified the portion of the bone proximal to the data on leg length in Appendix 1). Measurements on two spec- groove as a distinct ossification (the medial centrale). There is no imens were also included in Rossmann (2000). The LL/SVL evidence for a separate ossification in this position in Geiseltal- ratio in specimens in which it could be measured was 0.60 iellus maarius. The groove on the plantar surface of the fourth (HLMD-Me 10207), 0.75 (SMF ME 2), 0.76 (SMF ME 1769) distal carpal occurs in several different squamate groups (see and 0.69 (SMF ME 2938); in the privately held specimen, Renous-Lécuru 1973). The fifth distal carpal seems to be a small LL/SVL was 0.83. triangular element positioned between the proximal ends of the The ratio of tibial to femoral length seems fairly uniform in fourth and fifth metacarpals. Geiseltaliellus maarius: 0.79 in SMF ME 2938, 0.82 in HLMD- Remaining features of the manus are fairly typical of Me 10207 and SMF ME 2, and 0.84 in SMF ME 2684. limbed squamates, including the phalangeal formula (2–3–4–5–3). The great distal expansion of the fifth metacarpal, squamation however, might be a distinguishing feature. Additionally, the Shallow grooves delineating the edges of epidermal scales were penultimate phalanges are conspicuously elongated relative to described previously on the frontal of the holotype, HLMD-Me the antepenultimate phalanges, by 43%, 76%, 55% and 100% in 10207. In certain other specimens, scales are preserved more digits II to V, respectively (Table 1); in digit III the penultimate directly, either as keratinous material or as “shadows” produced phalanx accounts for 46% of the total length of the nonungual by anoxic bacterial decay of soft tissue (Franzen 1985). Scales phalanges. preserved as shadows are particularly prevalent in SMF ME 2a on the nape and abdomen (Figure 10). On the nape, patches of Proportions. The arms are comparatively short in Geiseltaliellus squamation show individual scales to have been very small (0.15 maarius. The ratio of arm length (AL, defined as the distance mm in edge-to-edge length), uniform, rhomboidal, and arrayed 238 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

in this specimen. Proximally to distally these correspond roughly to the wrist, the palm and the subdigital areas. The scales of the wrist tend to be larger than those of the palm and are sometimes rhomboidal. Especially in the region above the distal ends of the radius and ulna, the scales tend to be thin and poorly preserved. They often overlap slightly. In areas where they can be measured adequately they have dimensions of about 0.75 to 1 mm. More distally, in the area of the palm, the scales are generally smaller (about 0.5 mm) and more rounded. They rarely seem to overlap at all. They are also distinctly thicker than those of the wrist, especially in their centers; thus, they have the form of low rounded bumps. The subdigital scales, finally, are proximodistally short (≤0.5 mm), but greatly widened perpendicular to the axis of the digit. The extremely wide subdigitals that run from the ventral surface of digit I to the ventral surface of digit II are presumably artifactual fusions of subdigitals from both digits. Adjacent to the subdigital scales are smaller ones, best preserved proximally on the postaxial side of metacarpals Figure 12. Comparative skeletal elements of Geiseltal- III and IV. Subdigital scales, but essentially no others, are also iellus longicaudus and Capitolacerta dubia. A, G. long- well preserved adjacent to the digits of the right foot in SMF ME icaudus holotype (GM 4043), left postorbital in lateral 1769, which liken those of the manus described above. There view. B, G. longicaudus holotype (GM 4043), left (top) are no keels on any subdigital scale. That scales of the extremi- and right (bottom) mandibles in lateral and medial ties, especially the subdigital scales, are so well preserved may view, respectively. C, G. longicaudus holotype (GM result from both scale thickness and less intense decomposition 4043), left clavicle in lateral view. D, Capitolacerta than in the more fleshy parts of the body. dubia (GM 4002), parietal in dorsal view. Abbrevia- Rossmann (2001) stated that Geiseltaliellus had a dewlap, tions: a, angular; cn, coronoid; d, dentary; sa, surangu- citing SMF ME 2a. It is not clear what anatomical evidence was lar; sp, splenial. For A–C, the arrow indicates the used to determine this. There is a closely apposed pair of long, dorsal direction; for D, the arrow indicates the ante- curved, rod-like structures preserved beneath the posterior por- rior direction. tion of the mandible, but these seem to be composed of bone (SMF ME 2b) and so are unlikely to be second ceratobranchials. On the other hand, they seem somewhat longer than the first in rows oblique to the long axis of the animal (Figure 10A). ceratobranchials of SMF ME 2938. Whatever the case, support These scales are considerably smaller than those indicated by for the existence of a dewlap is weak. Rossmann (2001) also the grooves on the frontal, which are more than 1 mm in their noted a webbing (Zwischenzehenhaut) between the toes in SMF smallest dimension. On a few scales (e.g., see Figure 10A, right ME 2b. Although there is a patch of scales preserved between side of inset) seem to have keels; these are absent, however, on metatarsal V and the proximal half of metatarsal IV, this could most scales and their occurrence is probably a taphonomic arti- as easily have been caused by the lateral dislocation (during fact. The patches of small scales continue anteriorly onto the decay) of the skin covering metatarsals I to IV. Webbing area adjacent to the right supratemporal fossa, suggesting that is extremely rare in Squamata (so far as I know, it has been this form of squamation extended onto the head and that the noted among living species only in Amblyrhynchus cristatus; see cephalic squamation may have consisted of a patch of enlarged Dawson et al. 1977) and evidence for it in Geiseltaliellus maar- scales above the dermal roofing bones surrounded by a ground- ius is very weak. mass of granular scales. Traces of scales are also seen on the abdominal region of SMF ME 2b, although they are white, not messel specimens compared black (Figure 10B). These scales, slightly larger (up to 0.22 mm I list here the variation among four Messel specimens in 19 fea- in transverse width) than the scales of the nape, are arranged in tures of the cranial skeleton and of proportion (Table 2). In transverse rows. These rows are also separated from one another comparing only HLMD-Me 10207 and SMF ME 2 and 1769 by a space equal to or greater than the anteroposterior length of for those 12 features visible in all three specimens and variable individual scales, perhaps from bloating of abdomen before its among them, (1) HLMD-Me 10207 and SMF ME 2 are more final collapse. similar to one another in four (prefrontal boss size, angle of In SMF ME 2684 scales are preserved on portions of both jugal, size of maxillary labial foramina, dentary projection at feet and, more notably, on the left hand (see Figure 11). The junction of angular and surangular), (2) HLMD-Me 10207 and specimen has suffered extreme plastic deformation and the SMF ME 1769 are more similar in three (frontal concavity, joints between the radius and ulna, the carpals, and the proxi- length of median parietal crest, and grooves for labial foramina mal metacarpals are scarcely distinguishable. The entirety of the of maxilla), and (3) SMF ME 2 and 1769 are more similar in five hand is preserved, however, with the exception of the end of (frontal sculpture, sinuousness to median crest of parietal, num- digit V. The scales are generally larger than those of the body. ber of rows of maxillary foramina, form of jugal–maxillary The keratin of the scales (stippled in Figure 11) has a dark rust suture, and relative leg length). With this nearly uniform distri- color that contrasts with the medium brown of the bone. Three bution of pairwise similarity, no one specimen stands out as dis- more-or-less distinct regions of squamation can be delineated tinctively different from the others, giving no reason for the Eocene Lizards of the Clade Geiseltaliellus • Smith 239 recognition of more than a single species. Sexual dimorphism is taxon has an “anterior, triangular table and expressed prominently in extant Basiliscus (Lang 1989) and prominent posterior sagittal crest.” The skull of Anolis (e.g., Williams 1972), but no feature of Geiseltaliellus maarius (except perhaps the peculiar sinuousness to the median GM 4042 shows a V-shaped, not Y-shaped, pari- parietal crest in HLMD-Me 10207) has obvious implications etal table, which (by definition) lacks a median for mate recognition. Range of variation in tail length in G. crest and whose shape is in any case not so clear maarius is like that seen in some modern long-tailed taxa (e.g., as figured by Kuhn (1944, pl. 20, fig. 3). Estes Brachylophus fasciatus; see below). (1983a) cites no specimen of Geiseltaliellus from Messel, and although one specimen had been Comparisons and Discussion excavated early enough (1976) to have been incorporated in his thought, Estes seems not to Geiseltaliellus longicaudus Kuhn, 1944 have visited the Senckenberg Museum during the This taxon name was introduced by Kuhn (1944) appropriate time (T. Keller, pers. comm. 2009). to encompass several fairly complete, if only mod- How Estes could correctly have inferred a Y- erately well preserved, specimens from the middle shaped parietal table is unknown. Eocene of Geiseltal near Halle an der Saale, Ger- Although the interrelations of the nasal, many. They are similar in size to specimens of frontal, maxilla and prefrontal cannot be deci- Geiseltaliellus maarius (see Appendix 1). Speci- phered in the holotype of Geiseltaliellus longi- mens identified as G. longicaudus derive from three caudus, the dorsal orbital margin of the prefrontal different localities in the Cecilia pit (formerly is fairly well preserved and shows a groove near abbreviated either Cl or Ce) of the former Geiseltal the prefrontal–lacrimal junction. The postorbital strip mine, which was closed officially in 1993 of G. longicaudus, as Kuhn (1944) noted, is a tri- (Hellmund 1997). The holotype comes from Ce III, radiate bone (Figure 12A). It has a longitudinal other reasonably well-preserved specimens from ventral groove for the reception of the posterior Ce II, III and IV (Haubold and Krumbiegel 1984). ramus of the jugal and the squamosal. In the holo- Ce II and III are approximate stratigraphic equiv- type, the dorsal margin of the posterior ramus of alents (Haubold 1990; see also Vetter 1931) in the postorbital is concave in lateral aspect and tapers Oberkohle, or “upper coal,” which has been tied to uniformly toward its posterior end; that is, the European Paleogene reference level MP 14 dorsal margin is not expanded. In GM 4072, how- (Franzen and Haubold 1987; Haubold 1990, 1993). ever, the left postorbital is clearly dorsomedially Considerable work on the depositional environ- expanded, as in SMF ME 2; moreover, the jugal ments and stratigraphic relationships of the facet on anterior ramus of the postorbital in this Geiseltal deposits has been published, including specimen is not exposed laterally, as in HLMD- papers by Weigelt (1931) and Krumbiegel (1955, Me 10207. There is no evidence in GM 4043 of 1977) on paleoenvironment and stratigraphy. a separate postfrontal, as in HLMD-Me 10207. Rossmann (2000) felt that the specimens of A lateral eminence is only weakly developed, Geiseltaliellus discussed here as G. maarius sp. possibly attributable to the smaller size of the nov. could safely be referred to G. longicaudus, specimen. but they were not explicitly compared. The holo- Both mandibles are preserved in GM 4043, type of G. longicaudus, GM 4043, is a moderately the left visible in lateral view, the right in medial crushed but mostly complete skeleton preserved view (Figure 12B). The morphology of the in left ventrolateral view (see Kuhn 1944, pl. 19, mandible, as far as can be seen, is comparable in fig. 1). Important elements of the skull missing in most details to that of Geiseltaliellus maarius. The the holotype and missing or crushed in the dentary has at least 18 teeth (perhaps two more referred specimens include the frontal and pari- each anteriorly and posteriorly). The Meckelian etal. In the original description of G. longicaudus, groove is mostly open but restricted, and it closes Kuhn (1944) did not mention a median parietal below dentary 8 and 9 (assuming two teeth ante- crest, nor did Haubold (1977) in his review of the riorly). The splenial is a long element that Geiseltal squamates. Despite having examined all extends anteriorly as far as the ninth dentary available specimens there that have been identi- tooth. The anterior inferior alveolar foramen is fied as G. longicaudus, I was unable to confirm toward the anterior end of the splenial; it is lon- Estes’ (1983a:33) statement that the parietal in this gitudinally oval and completely enclosed within 240 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Figure 13. Comparison of (A) tail length and (B) leg length, in centimeters, in relation to snout-vent length in Corytophaninae and Geiseltaliellus. the splenial near that bone’s dorsal margin. Ante- but the expansion differs from that of G. maarius riorly the angular is fusiform in medial aspect, (Figure 7B) in being located more dorsolaterally, extending anteriorly well past the apex of the displaced away from the fenestra and possibly giv- coronoid. The posterior mylohyoid foramen ing the clavicle a posterior hook. In G. maarius, penetrates its medial face probably close to the expansion of the clavicle is at the same level as the level of the coronoid apex. The lateral exposure fenestra. of the angular is difficult to gauge, for this region Relative tail length in GM 4146 (TL/SVL) is of the mandible is damaged in the holotype (as estimated to be 3.2 (see Appendix 1) and is simi- well as in referred specimens); it seems more sim- lar in other specimens. ilar to the angular in SMF ME 2938 (Figure 3) in Epidermal scales are not as well preserved in not extending as far anteriorly in the space Geiseltaliellus longicaudus as they are in SMF ME between the posterior processes of the dentary. 2, a paratype of G. maarius, but where they are In lateral view the coronoid has a strong poste- preserved they retain some three-dimensional rior process over the surangular that projects to structure. G. longicaudus had wide, seemingly the end of the anterior surangular foramen (Fig- noncarinate subdigital scales. ure 12B). It also has a broad ventrolateral flange that extends nearly halfway down the body of the Capitolacerta dubia Kuhn, 1944 mandible. (Whether this flange also extended In the same publication Kuhn (1944) also coined anteriorly past the posterior end of the tooth row the name Capitolacerta dubia for the reception cannot be determined, because the end of the of several small lizard specimens from Geiseltal. tooth row is covered by the left maxilla and its The catalog numbers of these specimens and dentition.) the number of individuals they represent have The right pterygoid of a referred specimen of been confused. There are only three specimens: Geiseltaliellus longicaudus, GM 4146 from locality the (missing) holotype, GM 4005, and two Ce II, bears a single row of four teeth close to its referred specimens, GM 4001 (part and counter- medial edge (contra Rossmann 2000). The right part) and GM 4002 (compare Kuhn 1944, quadrate of the holotype is partially preserved; it Krumbiegel 1959, Haubold 1977 and Haubold seemingly differs little from that of G. longi- and Krumbiegel 1984.) caudus, except that the foramen piercing the The type and referred specimens are all quadrate is closer to the ventral condyle. from Ce IV (Haubold and Krumbiegel 1984). The clavicle of Geiseltaliellus longicaudus is This species—placed by Kuhn in Scincidae and fenestrated and ventrally expanded (Figure 12C), later by Haubold (1977) in Lacertidae—was Eocene Lizards of the Clade Geiseltaliellus • Smith 241 synonymized with G. longicaudus by Estes Other Iguanids (1983a), whose arguments were supported and The description of characters used in the phyloge- supplemented by Rossmann (2000). netic analysis (Appendix 3) and the data matrix The specimens of Capitolacerta dubia are not (Appendix 2) provide explicit and implicit osteo- well preserved. Estes (1983a:34) stated that logical comparison with other iguanids. Here I “[s]kull structure in Capitolacerta duplicates that restrict comparison to proportions and squama- of Geiseltaliellus.” Probably he was alluding here tion, particularly in Corytophaninae. In his initial to those features of the skull (e.g., azygous frontal, description Kuhn (1944) noted that Geiseltaliel- open supratemporal fenestra) that removed lus longicaudus is remarkable for its long tail Geiseltaliellus from Lacertidae. As far as it can be (approximately three times body length) and studied, the skull of C. dubia is indeed similar to explicitly compared this to Basiliscus. Rossmann that of Geiseltaliellus longicaudus (and G. maar- (2000) additionally cited the presence of hind ius) in the morphology of the jugal and its rela- limbs approximately twice as long as forelimbs as tion to the maxilla (Smith pers. obs.) and the diagnostic of Corytophaninae in referring morphology of the teeth (Haubold 1977). The Geiseltaliellus to that group. A further discussion frontal in C. dubia, like that of G. maarius, is con- of these body proportions in iguanids more cave in transverse cross section and its sharp generally is warranted, not only for the reasons edges suggest the presence of supraorbital of taxonomy, but also because aspects of body flanges. In one important respect, however, the proportions have been related to the habitat skull of C. dubia departs from that of G. longi- and mode of life of living and fossil iguanids caudus. In the one available specimen of C. dubia (e.g., Losos 1990; de Queiroz et al. 1998; Beuttell in which the structure of the parietal can be eval- and Losos 1999). uated (GM 4001), the parietal table is trapezoidal (Figure 12D). If C. dubia is properly syn- Description onymized with G. longicaudus, and if the latter indeed had a Y-shaped parietal table as an adult Tail length. Tail length is a product of two distinct (Estes 1983a), then the ontogeny of the parietal is properties, individual vertebral length and verte- like that of extant Basiliscus and probably also G. bral number. Etheridge (1967) noted that both maarius. variables play a role in iguanids. Increased tail Tooth count in Capitolacerta dubia is not length is theoretically advantageous for both grossly inconsistent with its synonymy with jumping and sprinting, primarily because the tail Geiseltaliellus longicaudus. GM 4001 has about 18 acts as a counterbalance (e.g., Losos 1990). maxillary teeth count of and approximately 3.0 Tail length and snout–vent length, a proxy for cm SVL. GM 4043, in contrast, has a maxillary body size, were measured in 129 iguanid specimens tooth count of about 22 and approximately 6.0 cm (see Appendix 1) representing Corytophaninae (9 SVL (Kuhn 1944). species), Polychrotinae* (17 species), Iguaninae (2 There is no reason not to retain Capitolac- species), Hoplocercinae (5 species), Tropidurinae* erta dubia in the synonymy of Geiseltaliellus (11 species), Oplurinae (4 species), Crotaphytinae longicaudus. The presumptive difference in pari- (2 species) and Phrynosomatinae (12 species). etal morphology probably reflects ontogenetic Adults from all species of Corytophaninae were transformation. Specimens of C. dubia have the examined, as were juveniles of Corytophanes per- large orbits common to juveniles (Kuhn 1944; carinatus, Basiliscus basiliscus and B. vittatus. The Rossmann 2000), poorly ossified vertebral specimens are all preserved in alcohol, these being condyles, and poorly ossified epiphyses unfused far more numerous and taxonomically diverse to their respective diaphyses (Smith pers. obs.). than available complete skeletal material. Unfortunately, the exhaustion of the known For all corytophanines, TL/SVL ≥ 1.7 (see productive Geiseltal localities (Hellmund 1997) Appendix 1 and Figure 13A). (For this discussion, and the seeming loss of the holotype of C. dubia taxa with TL/SVL ≥ 2.0 are defined as having long mean that there is not likely to be new informa- tails.) Though the ranges overlap, individuals of tion bearing on the problem in the foreseeable Corytophanes generally have a lower TL/SVL future. ratio compared to other corytophanines (1.7 < 242 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

TL/SVL < 2.5, with the lowest values all from sum of upper leg length, lower leg length and foot C. percarinatus), and individuals of Laemanctus length to the end of metatarsal IV) was measured higher TL/SVL (>3.0), while individuals of Basilis- in these same taxa (see Appendix 1). (The speci- cus fall in the middle (2.2 < TL/SVL < 2.9). All mens are alcohol-preserved, so the measurements hoplocercines examined have short tails. The should not be taken to equal exactly the sum of iguanines examined, including the basal (Norell the lengths of the femur, tibia, ankle and and de Queiroz 1991) member Dipsosaurus metatarsal IV; they are approximations. For the dorsalis, have TL/SVL ≥ 2. Among polychrotines, purposes here, species showing LL/SVL > 0.67 are tail length in many species falls well below considered to have “long legs.”) the TL/SVL ϭ 2.0 reference, while many others All corytophanines examined fell above the are above it. Some of the former are members of reference line (Figure 13B), and only 3 of 31 the leiosaur group (Leiosaurus belli, Diplolaemus showed LL/SVL < 0.70. Intraspecific variation bibronii and Pristidactylus torquatus), although (from 0.71 to 0.81, close to that for Geiseltaliellus Anolis (formerly Phenacosaurus) richteri and maarius, excepting the privately held specimen) Urostrophus vautieri are also among them. Exam- was recorded. Most noncorytophanines did not ined species of Polychrus, other Anolis, Enyalius have long legs. However, Anolis was highly vari- and Anisolepis have long tails. Considerable variable; A. richteri showing the lowest ratio and A. ation in Anolis has previously been shown (e.g., cobanensis and A. cristatellus had the highest. Losos 1990). Crotaphytines and examined mem- Crotaphytines, some of which have been docu- bers of Oplurinae were borderline, with a TL/SVL mented to be facultative bipeds (Snyder 1949, ratio just under 2.0. The tropidurines examined 1952), were generally above the reference value. had a TL/SVL ratio well under this value, except The basal iguanine Dipsosaurus dorsalis, also Microlophus occipitalis and Tropidurus torquatus, known to be a facultative biped (Snyder 1952), which were borderline. Most phrynosomatines had short legs (range 0.56 to 0.62); and Brachylo- had considerably shorter tails (TL/SVL < 1.5), phus fasciatus is well below the reference value. with the lowest value shown by Phrynosoma. The hoplocercines examined showed considerable As discussed above, two specimens of interspecific variation, but all except Enyalioides Geiseltaliellus maarius have TL/SVL > 3.0; in the heterolepis were short-legged. Oplurines fell just other two this value is close to 2.0 (cf. Rossmann below the reference line (except one specimen of 2000). Tail length evidently varies considerably in Chalarodon madagascariensis), as did most this taxon, as it does in some extant taxa with long tropidurines and phrynosomatines (except “sand tails (e.g., Brachylophus fasciatus; see Table 1). In lizards,” some of which are known to be cursorial: summary, many living taxa with TL/SVL > 2 are Stebbins 2003). in clades other than Corytophaninae, although In summary, members of Corytophaninae TL/SVL > 3 is rare, confined to Laemanctus, some have longer legs (hind limbs) than members of Polychrus, and some iguanines. most other major clades except Crotaphytinae, although interspecific variation in certain groups, Leg length. Another aspect of the body propor- particularly Polychrotinae*, is high. Geiseltaliel- tions of Geiseltaliellus noted by Kuhn (1944), lus also falls in the long-legged group. Variation in Estes (1983a) and Rossmann (2000) is the rela- Geiseltaliellus maarius, however, is relatively high. tively long legs. All three authors also compared this feature in the fossil taxon with the Coryto- Arm length. Arm length is another morphomet- phaninae, although they did not provide quanti- ric characteristic with ecomorphologic import. tative comparisons. Lang (1989) contended that Increased arm length is expected to have a nega- there was little variation in the hind limb among tive influence on jumping and sprinting ability corytophanines. Functionally, longer legs have (e.g., Losos 1990). Arm length (AL, in specimens been shown to have strong positive effects on per- of extant species, the sum of upper arm length, formance in jumping and sprinting (e.g., Losos lower arm length and hand length to the end of 1990). metacarpal III) was measured in most of the cory- To examine the length of the hind limb in a tophanines, polychrotines, hoplocercines and broader taxonomic context, this length (LL, the iguanines referred to above (see Appendix 1). Eocene Lizards of the Clade Geiseltaliellus • Smith 243

Figure 14. Comparison of manual scales in select iguanids. A, Right manus of Laemanctus serratus (SMF 11016), palmar view. B, Right manus of Corytophanes hernandesii (SMF 11015), palmar view. C, Right manus of Basilis- cus galeritus (SMF 11029). D, Left manus (mirrored) of Polychrus acutirostris (SMF 62421), palmar view. E, Left manus (mirrored) of Enyalius catenatus (SMF 11040), palmar view. F, Left manus (mirrored) of Pristidactylus torquatus (SMF 43945), palmar view; the curious scalation of the palm is not found on the right manus. G, Left manus (mirrored) of Diplolaemus bibronii (SMF 58523 or 58524), palmar view. H, Right manus of Anisolepis undulatus (SMF 11055 or 11056), palmar view. Scale bars are 1 mm.

The ratio of arm length to leg length (AL/LL) from one individual of L. serratus). In species was less than 0.50 in all Basiliscus examined. In where multiple individuals were sampled, there Corytophanes, it ranged from 0.52 to 0.57, and in was no clear ontogenetic trajectory. Species of Laemanctus from 0.52 to 0.63 (the latter value Basiliscus thus have relatively shorter arms than 244 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 other corytophanines and for most corytopha- 53) illustrated subdigital scales in C. percarinatus as nines (except L. serratus) AL/LL ≤ 0.57. For all multicarinate, unlike in the first two species. The examined members of Polychrotinae* had AL/LL palmar scales, as in Laemanctus, are rhomboidal ≥ 0.57, but these species included only one mem- and otherwise do not differ substantially from the ber of Anolis (A. richteri) and none of Anisolepini. subdigital scales. The scales of the wrist are similar I discovered considerable variation in Enyalius in size and morphology to those of the palm but are catenatus and Leiosaurus bellii. Except for more strongly imbricate. Given outgroup condi- Enyalioides heterolepis, which as noted above has tions (see above and below) and the basal position relatively long legs, all hoplocercines had AL/LL ≥ of C. hernandesii (Lang 1989), it is possible that the 0.64. Among iguanines, Brachylophus fasciatus unicarinate condition of that species and of C. had a AL/LL ratio between 0.66 and 0.71, whereas cristatus is primitive for Corytophanes. the facultative biped (see above) Dipsosaurus dor- In Basiliscus galeritus (Figure 14C), B. basilis- salis had relatively shorter arms (AL/LL ratio cus and B. plumifrons the palmar scales of the hand from 0.53 to 0.55), which cannot wholly be attrib- are generally smooth and relatively small (less than uted to an increase in leg length. half the size of the subdigitals). In preaxial–postax- In summary, members of Geiseltaliellus ial cross-section they are wedge-shaped, the tallest have relatively shorter arms than all examined point located postaxially. They are somewhat living Polychrotinae*, all Hoplocerinae except smaller than the scales of the wrist and do not Enyalioides heterolepis, and the faculatative biped overlap so strongly. There is only a slight tendency Dipsosaurus dorsalis. Arm length in Geiseltaliel- in B. plumifrons to develop small bumps on the lus most closely matches the range of Coryto- palmar and subdigital scales. Both juvenile and phanes, being greater than that seen in Basiliscus. adult individuals of B. vittatus, on the other hand, typically have small but distinct bosses on both Squamation of the manus. In Laemanctus palmar and subdigital scales. The palmar scales longipes and L. serratus (Figure 14A), the subdig- remain small, but they are distinctly larger than ital scales are rectangular to hexagonal in outline, the same scales in the other species of Basiliscus. while those of the palm are rhomboidal. Other- The palmar and subdigital scales of B. vittatus wise the scales of the palm are little distinct from approach those of Laemanctus in their morphol- the subdigital scales. Both are relatively large and ogy. The subdigital scales of other species of show comparatively little overlap; additionally, Basiliscus are very wide, that is, strongly expanded both are adorned with large, rounded protuber- perpendicular to the axis of the digit, more so on ances that become worn apically. This adornment the first digits than the last. distinguishes them readily from the scales of the In Lang’s (1989) phylogenetic hypothesis of body, which are similar in size but lack the protu- Corytophaninae, the relationships among Basilis- berances. The scales of the wrist are large, thin, cus galeritus, B. vittatus, and B. basiliscus + B. imbricate and rhomboidal. Because this morphol- plumifrons were not resolved. Members of the lat- ogy is concordant in both extant species, it can be ter clade were united by their common possession taken as primitive for Laemanctus. Although the of greatly elongated neural spines on the thoracic number of keels on the subdigital scales in Lae- vertebrae. B. vittatus is likely to be more closely manctus has been considered to be reduced related to the B. basiliscus + B. plumifrons clade, (Etheridge and de Queiroz 1988; Lang 1989), even because it has somewhat elongate neural spines more distinctive is the morphology of the protu- (Maturana 1962; Smith pers. obs.). B. galeritus, in berance (Boulenger 1885; Smith 1944; Smith and which vertebral spines are not elongated, is in this Laufe 1945). view the most basal member of Basiliscus. This In Corytophanes hernandesii (Figure 14B), conclusion finds support in the phylogenetic C. cristatus and C. percarinatus, the subdigital scales analyses of Schulte et al. (2003), although they did of both manus and pes are nearly rectangular in not include B. basiliscus, and Vieira et al. (2005). aspect. Unlike in Laemanctus each of these scales If B. vittatus is nested within Basiliscus, one can has a strong and distinct keel that usually bears a conclude that palmar scales, much smaller than well-developed projection, sometimes drawn out the subdigital scales, are likely primitive for into a strong spine, at its distal end. Lang (1989, fig. Basiliscus. Notably, Maturana (1962), though he Eocene Lizards of the Clade Geiseltaliellus • Smith 245 did not discuss palmar scales, found B. vittatus to of the more preaxial and distal palmar scales. This be the most divergent of all species of the genus. is taken to be a characteristic of the palmar scales A comprehensive study of the manual squa- themsevles, for (1) there are very few palmar mation of other iguanids is beyond the scope of this scales in comparison with other polychrotines work, but additional comparisons with Polychroti- and (2) the subdigitals are not reduced in size, nae* could be useful for understanding the distri- being equal or nearly equal in width to the digits. bution of certain characters discussed above. Although there are exceptions (e.g., Peterson and Summary of Comparisons Williams 1981), Anolis is well known for the trans- One of the most distinctive features of Geiseltaliel- versely expanded subdigital lamellae (e.g., lus maarius is the median parietal crest, a feature Boulenger 1885; Etheridge and de Queiroz 1988); that cannot be confirmed in G. longicaudus the palmar scales of many species are small relative despite reports of its presence. Certainly G. maar- to the diameter of the digits (excluding lamellae). In ius and G. longicaudus are very similar, for Polychrus acutirostris the polygonal palmar scales instance, in the detailed structure of the mandibu- are thick and moderately imbricate (Figure 14D). lar bones and their relations to one another, in the They are sometimes keeled and always bear one to morphology of the teeth and of the suborbital three small spines. They are small relative to the region, and in their general body proportions. size of the hand and the size of the subdigital scales The largest specimens referred to G. maarius and on digits I and II. Other digits, particularly III and G. longicaudus are similar in size. The presence of IV, have subdigital scales that are narrow with a posterolateral process of the coronoid is an apo- respect to the diameter of the digit, blurring the size morphic feature (Lang 1989) shared by the two disparity between subdigital and palmar scales. species. Many taxa in Leiosaurini have small palmar scales Nevertheless, two features of the type species relative to the subdigitals, although none is exactly of Geiseltaliellus (G. longicaudus) distinguish it like the condition in Basiliscus. In comparison with from G. maarius: a large ventrolateral expansion Basiliscus, the subdigital scales of Enyalius catena- of the coronoid and a clavicle with a more dor- tus are more equidimensional (Figure 14E); the sally located expansion. Known variation in G. decrease in the width of these scales near the mid- maarius (see Table 2) does not extend to these length of the phalanges is more pronounced in this two features. Thus G. longicaudus differs from the species than in many other iguanids, including E. Messel specimens in ways that they do not differ bilineatus. The palmar scales are small in compar- from one another, supporting recognition of the ison with the subdigitals. In Pristidactylus torqua- latter as a distinct species. Expansion of the clav- tus the palmar scales are thick, rounded, smooth icle is limited to the ventromedial end of the bone and essentially nonimbricate (Figure 14F); they are in corytophanines, as in G. maarius, so the dorsal smaller than the subdigital scales, but the distinc- position seen in G. longicaudus is probably the tion is not as strong as in Basiliscus. The subdigitals, derived condition. A strong ventral expansion of in turn, are slightly more transversely expanded the coronoid is known in a few iguanine iguanids than in Enyalius catenatus. In Diplolaemus bibronii (de Queiroz 1987), but is otherwise absent in all ventral scales of the manus, except the distal- Iguanidae. Thus, the expansion in G. longicaudus most subdigital scales, are reduced (Figure 14G). is probably also derived. This character should The palmar scales in particular are very small, prove useful for the rest of the fossil record, which rounded and nonimbricate; they are much smaller is dominated by jaws. than the subdigitals. Thus, in examined members In body proportions Geiseltaliellus is similar of Anolis, Polychrus and Leiosaurini, the palmar to members of Corytophaninae. Both show, scales are generally distinctly smaller than the sub- nearly unexceptionally, long legs (LL/SVL > 0.67), digitals, as in Basiliscus and Geiseltaliellus maarius. short arms (AL/LL ≤ 0.57) and long tails (TL/SVL Not so in Anisolepini. In Anisolepis undulatus > 2.0). Several other clades of Iguanidae also show (Figure 14H) and Urostrophus vautieri (not these features, however. Several polychrotines and shown), the palmar scales are relatively larger the iguanines examined have long tails, and the than in the foregoing taxa, often nearly equaling crotaphytines and oplurines examined have a the size of the subdigitals. This is especially true TL/SVL ratio close to 2.0. Some polychrotines 246 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Figure 15. Strict consensus of 14 equally most-parsimonious cladograms (seven distinct ingroup topologies) of iguanian relationships based on maximum parsimony analysis of morphological characters, assuming all character state changes to be equally weighted (between-state scaling). Numbers above branches are Bremer support values, those below branches are bootstrap percentages (given only for >50% support). Excluding parsimony- uninformative characters, tree length is 638 steps; CI ϭ 0.274, RI ϭ 0.559, RC ϭ 0.153, HI ϭ 0.726. Abbrevia- tions for taxon names (as shown on right): Pol., Polychrotinae*; Cor., Corytophaninae; Igu., Iguaninae; Hop., Hoplocercinae; Cro., Crotaphytinae; Opl., Oplurinae; Tro., Tropidurinae*; Phr., Phrynosomatinae. and (less commonly) phrynosomatines and shows the greatest similarity in relative tail and leg tropidurines have relative leg length approaching length to Crotaphytinae and Corytophaninae; it the condition in Corytophaninae and Geiseltaliel- is also similar to Corytophaninae in having rela- lus, as do most crotaphytines. Thus, Geiseltaliellus tively short arms (which were not measured in Eocene Lizards of the Clade Geiseltaliellus • Smith 247

Crotaphytinae). Relative leg and tail length are mations between a small number of linearly included as characters in phylogenetic analysis to ordered states. It may be useful in the future to further examine their evolution (see below). adopt a gap-weighting scheme (Thiele 1993; For squamation, Geiseltaliellus maarius is Wiens 2001). Phylogenetic structure in the data very similar to Basiliscus in having large scales was probed using the permutation tail probability above the dermal roofing bones, relatively small (PTP) test (Archie 1989; Faith and Cranston scales on most of the body, small, thick and less 1991) in PAUP* v. 4b10 (Swofford 2002) and the imbricate palmar scales, and very wide subdigi- skewness test (g1 statistic; Hillis and Huelsenbeck tals. In Corytophanes and Laemanctus the sub- 1992) based on 10,000 random, equiprobable digitals scales are less distinct from those of the trees produced in MacClade v. 4.08 (Maddison palm and forearm, which is possibly an apomor- and Maddison 2005). phy uniting them given the size disparity Parsimony analyses were conducted with between palmar and subdigital scales in poly- PAUP using the heuristic search algorithm (1000 chrotines (except Anisolepini) and a potential random addition sequence replicates, tree bisec- sister-group relationship between Polychrotinae* tion and reconnection (TBR) branch-swapping; and Corytophaninae (Frost et al. 2001; Conrad other settings default). Outgroups were taken to 2008; see also below). be either the five real species or the hypothetical ancestor. Between-state and between-character Phylogenetic Relationships scaling (Wiens 2001) were adopted in separate of Geiseltaliellus analyses for each outgroup class. Bootstrap support (Felsenstein 1985a) was calculated from 1000 A species-level data matrix for Iguania compris- bootstrap pseudoreplicates, each using a heuristic ing 34 ingroup species and 152 morphological search (as described above) with 100 repetitions. characters (145 parsimony-informative; see Decay indices (“Bremer support”) for individual Appendices 2 and 3) was subjected to phyloge- nodes (Bremer 1994) were computed using netic analysis by maximum parsimony and TreeRot v. 3 (Sorensen and Franzosa 2007). The Bayesian methods. Two different kinds of out- statistical significance of the shortest tree relative group to Iguania were used: (1) a set of real to alternative hypotheses was also assessed by species of living (Sphenodon punctatus) and fossil Wilcoxon signed ranks (WSR) (Templeton 1983; (Diphydontosaurus avonis) members of Rhyn- Felsenstein 1985b) tests, implemented in PAUP. chocephalia and members of each of the three MrBayes v. 3.1.2 (Huelsenbeck and Ronquist major living noniguanian squamate clades 2001; Ronquist and Huelsenbeck 2003) was used to (Eublepharis macularius for Gekkota, Plestiodon conduct Bayesian analysis of the data set. Additive fasciatus for Scincomorpha and Elgaria multicar- characters were treated as linearly ordered states by inata for Anguimorpha) and (2) a hypothetical using the “ordered” argument of the ctype com- ancestor constructed using polarity decisions mand. Diphydontosaurus avonis or the hypothetical based on examination of these and many other ancestor were treated as the sole outgroup in differ- species in outgroups to Iguania (see Appendix 3). ent analyses. One cold and two heated chains were A broad set of outgroup species is advisable given used. Default (generally flat) priors were assumed. conflicting support for the basal compared with The analysis was allowed to run for 2 106 gener- the deeply nested position of Iguania within Squa- ations, after which time the standard deviation of mata in morphological and molecular analyses split frequencies was reduced to about 0.006 and (Estes et al. 1988 and Conrad 2008 vs. Harris et al. convergence was checked. This latter value did not 1999, 2001; Townsend et al. 2004; and Vidal and much vary in dependency on outgroup choice. Hedges 2005). No attempt was made to include Sampling every 100 generations produced 20,000 apomorphies of traditional squamate clades (i.e., trees from the posterior probability distribution, of Scleroglossa, Autarchoglossa) except Iguania. which the first 5,000 were discarded as burn-in. Among species referred to Geiseltaliellus, G. The fossil record of Iguanidae is still poorly maarius alone was included because it is by far the known, despite recent additions of Eocene taxa most completely known. Additive characters were (Smith 2006a, 2006b, 2009; Conrad et al. 2007). treated qualitatively as equally weighted transfor- In the next several years stratigraphic data might 248 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 also reasonably be incorporated (e.g., Fisher 1994; Phrynosoma platyrhinos than to Sceloporus undu- Huelsenbeck 1994; Fox et al. 1999; Pol et al. 2004) latus. Tropidurini and Liolaemini are each mono- in the phylogenetic analysis of iguanian lizards. phyletic, and Tropidurinae* is the sister-taxon to Oplurinae. Relations outside of Clade B are Parsimony Analyses unchanged. The use of a hypothetical ancestor A PTP test (1000 replicates, each based on 100 instead of the five real outgroup species did not heuristic search repetitions) rejects the notion that affect ingroup topology, although the island con- there is no phylogenetic structure in the data (p taining the single ingroup topology was discov- 0.001). This is confirmed by the skewness of the dis- ered in more of the addition sequence replicates tribution of the 10,000 random trees (g1 –0.385, when the hypothetical ancestor was used. p < 0.01); the mean length of these trees is separated Bremer support, like total tree length, was gen- by 19 standard deviations from the most parsimo- erally reduced under between-character scaling. nious solutions computed using traditional However, monophyly of Iguanidae and of Iguania between-state scaling. Using between-state scaling, (without Saichangurvel davidsoni) received higher these encompass seven equally most-parsimonious Bremer support. This presumably reflects the ingroup topologies (14 trees in total) found in a sin- nature of the apomorphies supporting these nodes gle island discovered in more than 90% of the addi- in the most parsimonious topology, which are tion sequence replicates (Figure 15). The Late exclusively transformations of binary characters Cretaceous Mongolian taxon Saichangurvel david- and so receive under between-character scaling rel- soni is found to be a stem-iguanian (cf. Conrad and atively greater weight than transformations between Norell 2007; Conrad 2008). (Crown) Iguania com- adjacent states in multistate characters. Bootstrap prises two sister-clades, Acrodonta (plus its stem) frequencies were similar under both character scal- and Iguanidae (for which no stem taxa were ing schemes. The only clades to receive more than expected or revealed). The Late Cretaceous Mon- 90% bootstrap support were Iguaninae and Anolis. golian taxon Priscagama gobiensis lies on the stem There is only a weak positive relationship between of Acrodonta, as in previous studies (Frost and Bremer support and bootstrap support for individ- Etheridge 1989; Conrad and Norell 2007; Conrad ual clades. The most striking difference is for basal 2008). Seven of the eight major, previously recog- nodes in the tree (Iguanidae, Iguanidae, Clades A nized, suprageneric groupings (Etheridge and de and B), where Bremer support was markedly higher Queiroz 1988; Frost and Etheridge 1989; Schulte et than bootstrap support, compared with more ter- al. 2003) are monophyletic (the monophyly of minal branches. This obtains under both between- Hoplocercinae was not tested). Iguanidae is com- state and between-character scaling schemes. posed of two sister-clades. The first, dubbed Clade Forcing Iguanidae to be nonmonophyletic pro- A, contains Crotaphytinae, Iguaninae, Hoplocerci- duces trees that are not significantly longer than the nae, Corytophaninae, and Polychrotinae*. Crota- most parsimonious topology (WSR test). phytinae is basal in Clade A; Iguaninae + For the sake of brevity, a full listing of unam- Hoplocercinae form the sister-clade to Polychroti- biguous apomorphies supporting nodes is provided nae* + Corytophaninae. The second major iguanid only for the more resolved tree obtained using equal clade, dubbed Clade B, contains Oplurinae, between-character scaling (Appendix 4). This list Phrynosomatinae and Tropidurinae*. Relations of differs from that for the between-state scaling only these three taxa within Clade B are unresolved in in that it is more resolved for Clade B (i.e., apomor- the strict consensus. Additionally, relations within phies for all other clades are the same). Unambigu- Phrynosomatinae are unresolved, and Tropiduri- ous character state changes underwriting some of nae* constitutes an unresolved polytomy among these clades are discussed in detail below (node Liolaemus pictus, Phymaturus palluma, Leio- numbers correspond to those in Figure 16). cephalus personatus and Tropidurini. The use of a hypothetical ancestor instead of the five real out- node 1 group species did not affect ingroup topology. (Saichangurvel davidsoni + Iguania) The use of between-character scaling resulted This clade is supported by the following apomor- in a single most-parsimonious ingroup topology phies: 19(0→1) jugal exposure becomes extensive that resolves the relations in Clade B (Figure 16). dorsal to the posterior extension of the facial Petrosaurus thalassinus is more closely related to process of the maxilla; 22(0→1) quadrojugal Eocene Lizards of the Clade Geiseltaliellus • Smith 249

Figure 16. Single most-parsimonious cladogram of iguanian relationships based on maximum parsimony analysis of morphological characters, assuming all characters to be equally weighted (between-character scaling). Out- groups omitted for clarity. Numbers above branches are Bremer support values, those below branches are bootstrap percentages (given only for >50% support). Excluding parsimony-uninformative characters, tree length is 531.92 steps; CI 0.268, RI 0.556, RC 0.149, HI 0.732. Larger, bold, serif font numbers at nodes are clade identifiers corresponding to the text and Appendix 4. Abbreviations for taxon names (as shown on right): Pol., Polychrotinae*; Cor., Corytophaninae; Igu., Iguaninae; Hop., Hoplocercinae; Cro., Crotaphytinae; Opl., Oplurinae; Tro., Tropidurinae*; Phr, Phrynosomatinae. All taxon names and abbreviations refer to crown clades. process of jugal lost; 43(0→1) frontals fuse early (internal to mandible); and 107(0→1) angular in ontogeny; 44(0→1) frontals constricted process on prearticular developed. Half of these between the orbits; 50(0→1) parietal foramen were previously given as autapomorphies of shifts to frontoparietal suture, but still predomi- crown Iguania (Estes et al. 1988), but with the nantly in the parietal; 90(0→1) teeth become dis- recognition of the recently described Saichangurvel tinctively tricuspid; 97(0→1) coronoid articulates davidsoni as a stem-iguanian, I hypothesize that predominantly lateral to supra-Meckelian lip they apply to a more inclusive clade or clades. 250 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 node 2 Acrodonta (Frost and Etheridge 1989) and no (Iguania) closer stem-taxa than Priscagama gobiensis were Autapomorphies of Iguania relative to discovered or included, which would indicate that Saichangurvel davidsoni are as follows: 1(0→1/2) these features arose outside the crown. Estes et al. premaxillary tooth count reduced to seven or six; (1988) note other autapomorphies of Acrodonta 31(0→1) infraorbital foramen not bounded ven- for living taxa—reduction in the number of scleral trally by palatine; 33(0→1) posterior splint of post- ossicles, loss of caudal autotomy—that have not frontal lost; 35(0→1) postfrontal contribution to yet been discoverable in stem taxa. orbital margin reduced; and 50(1→2) parietal foramen partly invades frontal. The second apo- node 7 morphy—regarding the bounding elements of the (Iguanidae) infraorbital foramen—needs further examination Autapomorphies are as follows: 10(0→1) anterior in Late Cretaceous crown and stem iguanians. inferior alveolar and subnarial arterial foramina separate on premaxillary process of the maxilla; node 3 74(0→1) processus rostralis present on planum (Priscagama gobiensis + Acrodonta) antorbitalis; 76(0→1) commissura vomeronasalis Autapomorphies are as follows: 1(1/2→3) pre- complete; 79(1→2) length of vestibulum to olfac- maxillary tooth count reduced to five or fewer; tory chamber increased; 80(0→1) vestibulum 8(0→1) dorsal projection at anterior end of pre- becomes S-shaped; 95(0→1) Meckelian groove maxillary process of maxilla braces premaxilla; becomes partially closed or fused; and 125(0→1) 14(0→1) maxillae with extensive mid-line con- scapular fenestra. The only previously published tact on palate; 38(0→1) jugal–squamosal overlap morphological evidence for a monophyletic extensive; 45(0→1) dorsal surface of frontal con- Iguanidae is from Conrad and Norell (2007), who cave in transverse cross section; 89(0→1) cheek identified as unambiguous autapomorphies der- tooth shafts become mesiodistally expanded; and mal rugosities present on frontal and parietal, and 93(0→1) tooth implantation at least partly nasal with reduced median contact; neither char- acrodont in cheek tooth series. The state change acter was fully included in this analysis. Conrad for character 14 was previously recognized as an (2008) later argued that Iguanidae is paraphyletic. apomorphy uniting Acrodonta and part of its Two of the character-state changes noted here stem (Frost and Etheridge 1989). Gao and Norell (separation of foramina on premaxillary process (2000) and Conrad and Norell (2007) suggested of maxilla and processus rostralis plani antor- that this state change could apply to all of Iguania bitalis) are virtually unique to Iguanidae and show on the basis of its occurrence in certain Late Cre- almost no reversal (see character discussions). Sev- taceous iguanians [but Conrad (2008) agreed with eral other autapomorphies (those in the nasal cap- Frost and Etheridge’s interpretation]; if these taxa sule) are later reversed in major clades within turned out to be stem acrodontans, there would Iguanidae; their status as autapomorphies of be no need for such a postulate. Conrad (2008) Iguanidae thus hinges primarily on other charac- suggested additional autapomorphies of this clade ters that define iguanid monophyly and ingroup which were not included in this analysis. topology, and they should therefore be interpreted with some caution. Partial closure or fusion of the node 4 Meckelian groove was seen as multiply acquired (Acrodonta) within Iguanidae by Etheridge (1959), a view that Autapomorphies of Acrodonta relative to will probably be confirmed once taxa on the stem Priscagama gobiensis are as follows: 34(0→2) ante- of major clades within Iguanidae become known. rior splint of postfrontal lost; 66(0→1) pterygoid teeth lost; 97(1→0) anteromedial process of coro- node 8 noid articulates significantly on medial surface of (Clade B) supra-Meckelian lip (externally on mandible); Autapomorphies are as follows: 31(1→2) palatine 99(0→1) dentary length increases; and 102(0→1) loses posterolateral process that partially bounds anterior surangular foramen ventrally displaced infraorbital foramen; 37(0→1) postorbital no on mandible. Several of these apomorphies were longer underlaps frontoparietal corner; 103(1→0) previously recognized as autapomorphic of crown anterior extent of angular reduced; 104(0→1) Eocene Lizards of the Clade Geiseltaliellus • Smith 251 posterior extent of splenial increased; 106(0→1) tine process of maxilla reduced in size; 23(0→1) posterior mylohyoid foramen posteriorly displaced; distal tip of temporal ramus of jugal not posteri- 127(0→1) large sternal fontanelle developed; orly deflected; and 135(1→0) mid-dorsal scale 133(0→1) superciliary scales elongate and imbri- row present. A broad nasal process is common to cate; and 134(0→1) enlargement of at least one sub- most members of Clade A, but shows reversal in ocular scale. A potential relationship among Anolis and convergence in Oplurus and Phryno- Oplurinae, Phrynosomatinae and Tropidurinae* soma, among taxa I have examined. Smith (2009) has long been recognized (Frost and Etheridge interpreted this feature as a potential synapomor- 1989) on the basis of the mandibular characters and phy of Polychrotinae* and Corytophaninae features of squamation listed above. To these poten- absent in the basal iguanine Dipsosaurus, but in tial synapomorphies I add the reduction in enclo- this analysis the condition in Dipsosaurus is inter- sure by the palatine of the infraorbital foramen and preted as reversal. To my knowledge, the loss of a the loss of underlap by the postorbital of the fron- deflected tip of the temporal ramus of the jugal toparietal suture (see also Smith 2009); the latter of shows no reversal in Clade A or convergence in these occurs otherwise only in Iguaninae and Clade B and could prove a robust autapomorphy Hoplocercinae, while the former shows only minor of the former, as could reduction in size of the convergence in Clade A. palatine process of the maxilla. node 19 node 22 (Clade A) (Iguaninae + Hoplocercinae) This clade is weakly supported by the following This node is weakly supported by the following autapomorphies: 96(0→1) intramandibular lamella autapomorphies: 37(0→1) postorbital underlap developed; 98(1→0) length of intramandibular of frontoparietal corner lost; 88(1→2) tooth septum reduced; 111(0→1) laterally or ventrolater- crowns flared; 91(0→1) tooth crowns asymmetric; ally facing zygosphenes developed, which are not and 100(0→1) anterolateral process of coronoid separated from the prezygapophyses; 113(1→0) developed. Except for tooth crown asymmetry, four sternal ribs; and 128(0→1) narrow sternal most of these state changes are local, not unique, fontanelle present. Character 96 shows some rever- changes previously recognized as potential sal within Clade A and convergence within Clade synapomorphies of Iguaninae and Hoplocerci- B; these synapomorphies of Clade A members may nae (Frost and Etheridge 1989). Asymmetry of be less robust. Zygosphenes were long recognized the tooth crowns is unique to Iguaninae and as common in the species interpreted here as Clade Hoplocercinae within Iguanidae, but would have A (Etheridge 1959). The presence of a narrow ster- to show reversal in members of the latter (e.g., nal fontanelle largely covered by the posterior Hoplocercus spinosus) and the former clades. If process of the interclavicle is a constant presence H. spinosus is the sister-taxon to the remainder in Clade A outside of Polychrus + Anolis and to my of Hoplocercinae (Torres-Carvajal and de knowledge is otherwise only sometimes encoun- Queiroz 2009), then this autapomorphy of node tered in Phrynosomatinae. It may prove a robust 22 would be ambiguous even if all hoplocercines autapomorphy. Smith (2006a) tentatively inter- except H. spinosus had asymmetric teeth. preted an elongate intramandibular septum as independently synapomorphic of Polychrotinae* node 25 and Phrynosomatinae, pending evaluation in cer- (Corytophaninae + Polychrotinae*) tain other iguanids. This analysis suggests instead This node is supported by the following autapo- that a shortened septum is derived, but that reelon- morphies: 46(0→1) supraorbital flanges present; gation in places is a local apomorphy (Polychroti- 51(0→1) recessus processi ascendentis developed nae*; see below). well anterior of posterior margin of parietal; 76(1→0) commissura vomeronasalis posterior node 21 incomplete; 125(1→0) scapular fenestra lost; and (Clade A exclusive of Crotaphytinae) 139(0→1) femoral pores lost. Some think that Autapomorphies are as follows: 3(0→1) nasal these two groups could be closely related (e.g., process of premaxilla broad and parallel-sided for Etheridge 1959; Frost and Etheridge 1989; Frost a significant portion of its length; 16(0→1) pala- et al. 2001; Conrad and Norell 2007; Conrad 2008). 252 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Smith (2009) suggested supraorbital flanges and recovered a sister-group relationship between relatively anterior placement of the recessus pro- Geiseltaliellus and Corytophaninae. cessi ascendentis as potential synapomorphies of Corytophaninae and Polychrotinae*. The loss of node 28 femoral pores here requires that they be regained (Corytophaninae) in Polychrus (cf. Etheridge and de Queiroz 1988). Autapomorphies are as follows: 1(1→0) premaxillary tooth count increases from seven to eight or NODE 26 more; 18(0→1) gutter developed on palatal shelf (Corytophaninae + (Geiseltaliellus maarius + of maxilla; 36(0→1) posterior ramus of postor- early Eocene iguanid)) bital dorsally convex; 50(0→1) parietal foramen Autapomorphies are as follows: 27(0→1) groove anterior displaced; 55(0→1) parietal blade devel- in orbital rim at junction of prefrontal and oped; and 62(0→1) ridge on posterior face of lat- lacrimal; 30(0→1) lacrimal facet of prefrontal eral conch of quadrate. formed by semicircular projection; 52(0→2) parietal table Y-shaped; 119(0→1) clavicle ventrome- node 30 dially expanded and fenestrate; and 153(0→1) leg (Polychrotinae*) length increases to two-thirds or more of Autapomorphies are as follows: 47(0→1) rugosi- snout–vent length. Several of these state changes ties developed on dermal roofing elements other have previously been recognized as autapomor- than the frontal and parietal; 82(0→1) calcified phies of Corytophaninae (Etheridge and de endolymphatic sacs extend into nuchal muscula- Queiroz 1988; Frost and Etheridge 1989; Lang ture; 98(0→1) length of intramandibular septum 1989). Relative leg length, though not previously increases; and 147(0→2) hemipenis strongly included in a phylogenetic analysis, has also been bicapitate. The first, second and fourth character- argued to be an autapomorphy of the clade (Estes state changes have long been recognized as poten- 1983a). Extreme reduction or loss of the post- tial synapomorphies of polychrotines (Frost and frontal could also eventually be determined to Etheridge 1989); strong hemipenial capitation have arisen on the stem of Corytophaninae, with was first recognized by Frost and Etheridge reenlargement of the postfrontal in Laemanctus. (1989). Increased length of the intramandibular That this is not presently so depends on the opti- septum is a potential autapomorphy (suggested mization of a merely closed Meckelian groove as by Smith 2006a), but the necessity of disarticu- a synapomorphy of Geiseltaliellus and the new, lated specimens or CT scans has limited its eval- early Eocene iguanid. The latter has a fully devel- uation in Anisolepini and many Leiosaurini. oped postfrontal (Smith 2009), and Geiseltaliellus Etheridge (1959) recognized a convergence in only a highly reduced one. However, it is highly skull form between Anolis and Polychrus on the likely that fusion of the Meckelian groove pro- one hand and corytophanines on the other. One ceeded independently within Polychrotinae* and such aspect of skull form, snout elongation, is even within Corytophaninae (Smith 2009); for included here, and Etheridge’s (1959) hypothesis instance, the groove is merely closed in Basiliscus of convergence is confirmed. There is some galeritus and B. vittatus, which are basal to B. molecular support for mutually conflicting kinds basiliscus (Lang 1989; Vieira et al. 2005). The early of polychrotine nonmonophyly (Frost et al. 2001; Eocene iguanid incorporated in this analysis has Schulte et al. 2003), which has not been discov- no autapomorphies. Reoptimization of the Meck- ered in morphological analyses (Frost et al. 2001; elian groove on the basis of denser taxon sam- Conrad et al. 2007; this study). pling in Polychrotinae* + Corytophaninae might then result in extreme reduction of the postfrontal node 31 as a synapomorphy of Geiseltaliellus and Coryto- (Polychrus acutirostris + Anolis) phaninae; 34(0→1). Several of these apomorphies Autapomorphies are as follows: 1(1→0) premax- were previously recognized as pertaining to the illary tooth count increased to eight or more; corytophanine crown, but clearly arose along the 29(0→1) canthal crest well developed on pre- stem. Augé (2005), in a phylogenetic analysis frontal; 87(0→1) posterolateral processes of using Frost and Etheridge’s (1989) matrix, also basisphenoid lost; 95(1→2) Meckelian sulcus Eocene Lizards of the Clade Geiseltaliellus • Smith 253

Figure 17. Fifty-percent majority-rule consensus of 15,000 trees drawn from the posterior probability distribution in Bayesian analysis. Numbers above branches are posterior probabilities. Abbreviations for taxon names (as shown on right): Pol., Polychrotinae*; Cor., Corytophaninae; Hop., Hoplocercinae; Cro., Crotaphytinae; Opl., Oplurinae; Tro., Tropidurinae*; Phr., Phrynosomatinae. Branch lengths are scaled. closed or fused for more than 50% of tooth row and 138(0→1) gular fold lost. Denser sampling of length; 108(0→1) ceratobranchial 2 longer than living and fossil taxa in Polychrotinae* may show ceratobranchial 1; 113(0→1) number of sternal premaxillary tooth count to have increased in par- ribs reduced; 121(0→1) clavicular insertion on allel in Polychrus and Anolis. In particular, Pristi- suprascapula; 132(0→1) mental scale divided; dactylus torquatus is unique among examined 254 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Leiosaurini in having seven teeth (the others have The posterior probability of this clade is 0.28 five or six) and stem members of Polychrus also when the hypothetical ancestor is used, and no have fewer than seven (Smith unpublised data). other clade formed of Acrodonta and an iguanid On the other hand, a well-developed canthal crest sister-group had a higher posterior probability. on the prefrontal might be more broadly distrib- The only ways in which the Bayesian tree was uted in Polychrotinae*. Lang (1989) stated that an positively incongruent with the parsimony tree elongate ceratobranchial 2 is present in all poly- using between-character scaling are that in the chrotines, but this is not the case at least in Pristi- former, Leiolepis and Brookesia are sister-taxa dactylus torquatus (see scans of FMNH 206964 and Microlophus is the sister-taxon to Tropidu- [DigiMorph.org. 2002–2005]). Further study of rus + Plica. Relationships within Acrodonta are the loss of the posterolateral processes of the not yet well resolved, so there is little background basisphenoid might benefit from frequency cod- against which to evaluate these results. However, ing, as this feature was found to be intraspecifi- the recovery in Bayesian analysis of a mono- cally variable in some Polychrus (Frost et al. 2001). phyletic “Tropidurus group” is consistent with A close relationship between Polychrus and Ano- previous morphological studies (Frost 1992). lis was first hypothesized by Etheridge (1959); this Both nodes were weakly supported under maxi- hypothesis was further developed by Frost et al. mum parsimony. (2001). Etheridge and de Queiroz (1988) found conflicting evidence on the position of Polychrus, Summary and Discussion with some characters suggesting it is basal in a The clearest result this analysis shares with previ- monophyletic Polychrotinae*, others suggesting ous ones (Etheridge and de Queiroz 1988; Frost a more nested position. and Etheridge 1989; Macey et al. 1997; Schulte et al. 1998, 2003) is stronger support for the primary Bayesian Analysis groups of Iguanidae than for the relationships A 50% majority-rule consensus tree from samples among them. Insofar as many of the characters of the posterior probability distribution is shown used by Etheridge and de Queiroz (1988) and in Figure 17. Monophyly of Iguania to the exclu- Frost and Etheridge (1989) also appear in this sion of Saichangurvel davidsoni receives strong analysis, and new conflicting characters were not support (p > 0.99). Monophyly of Iguanidae, added, monophyly of these groups was expected. however, is not strongly supported (p 0.25). Both parsimony and Bayesian analyses reject the Support for the monophyly of Acrodonta and of position of Saichangurvel within crown Iguania. Agama + Physignathus, and for the position of Between-character compared with between-state Priscagama gobiensis on the stem of Acrodonta is weighting schemes did not result in positive strong. There is also moderately strong support incongruence in tree topology, although between- for the monophyly of Corytophaninae and for the charater weighting produced a more resolved position of Geiseltaliellus maarius and the early topology. The Bayesian topology shows only Eocene iguanid on the stem of that clade. Support minor incongruence with the parsimony trees. for a sister-group relationship between Polychrus Parsimony suggests Iguanidae is monophyletic and Anolis is strong (p > 0.99), but monophyly of and unique autapomorphies are recovered in sup- Polychrotinae* and the sister-group relationship port of it. Iguanidae receives higher support from between Corytophaninae and Polychrotinae* is decay indices than in previous studies (Conrad notably weaker. Other clades with high posterior and Norell 2007), but bootstrap support remains probabilities are Phrynosomatinae, Oplurinae weak. Iguanid monophyly is not contradicted by and Iguaninae. When Diphydontosaurus avonis the Bayesian analysis. Both analyses suggest, with is used as the outgroup, there is some support at least moderate statistical support, that Geiseltal- (p 0.7) for a sister-group relationship between iellus is on the stem of Corytophaninae and that the total clade of Acrodonta and Crotaphytinae, Corytophaninae and a monophyletic Polychroti- which is presumably related to similarities such nae* are sister-taxa. as the loss of the postfrontal, mesiodistal expan- These results are in large measure consistent sion of the tooth shafts (in Crotaphytus), and with recent reanalysis (Schulte et al. 2003) of presence of a quadratojugal tubercle on the jugal. an old data set (Frost and Etheridge 1989). Eocene Lizards of the Clade Geiseltaliellus • Smith 255

Although iguanid monophyly was not supported and Tropidurinae* form a clade with Oplurinae (Acrodonta was sister-taxon to Corytophaninae), as their sister-taxon, which would make the par- two major clades were also recovered with nearly allel more striking.) identical topology; the only difference was the position of Crotaphytinae, which in the tree of Comments on “Messelosaurinae” Schulte et al. (2003, fig. 1) was basal to what is here Messelosaurinae was erected (Rossmann 2000) called Clade B. as a subfamily of Corytophanidae (sensu Frost There is little agreement between the most and Etheridge 1989) and was intended to encom- parsimonious topologies given here and maxi- pass Aciprion formosum Cope, 1873, Geiseltaliel- mum likelihood topologies based on molecu- lus longicaudus Kuhn, 1944, Cadurciguana lar–genetic data. Monophyly of Tropidurinae* hoffstetteri Augé, 1987, Cypressaurus hypsodontus and Polychrotinae* in particular find little sup- Holman, 1972, and ?Crotaphytus oligocoenicus port in molecular studies. Anolis has been related Holman, 1972. The name Messelsaurinae was to the tropidurine genus Leiocephalus with high first published in an abstract (Rossmann 1993) posterior probability (Schulte et al. 2003). Opluri- and then in two later papers (Rossmann 1999a, nae has been related to South American anoloids 1999b) without establishment of a type genus. A (Schulte et al. 2003; Noonan and Chippindale nominotypical generic name that would yield 2006). Polychrus has been related to Basiliscus “Messelosaurinae” for the subfamily does not (Noonan and Chippindale 2006) and to exist. Rossmann (1999a) placed ?Crotaphytus Tropidurini (Schulte et al. 2003). Crotaphytus has oligocoenicus in the genus Holmanisaurus, agree- been related to Phrynosoma (Noonan and Chip- ing with Estes (1983a, who included Gambelia in pindale 2006) or placed at the base of Iguanidae the synonymy of Crotaphytus) that reference of (Schulte et al. 2003), with weak support. the species to Crotaphytus was poorly supported. Etheridge (1959) undertook the first serious The generic name Holmanisaurus, however, was treatment of relationships within Iguanidae in his published earlier (Rossmann 1993) without diag- quest to understand the relationships of the nosis, a type species or a list of included species, anoles. He recognized three “groups” of iguanids. and is a nomen nudum (Sullivan and Holman Group 1 corresponded, more or less, to what is 1996). Diagnostic features of the subfamily were now recognized as Tropidurinae* and Phryno- given as follows: parietal foramen at frontopari- somatinae; Group 2 to Crotaphytinae, Iguaninae, etal suture; “trapezoidal parietal” with a median Hoplocercinae, Oplurinae, Corytophaninae, sagittal crest (but without an occipital extension); Leiosaurini and part of Anisolepini; and Group 3 Meckelian sulcus open along its entire length; to Anolis and Polychrus. In this phylogeny, Ano- coronoid with a large, posteriorly directed labial lis and Polychrus (Group 3) are embedded at the process; and medial centrale present in carpus. tip of Clade A, and Phrynosomatinae and Cadurciguana and Geiseltaliellus were also linked Tropidurinae* (Group 1) constitute the bulk of by “the similar morphology of the dorsal verte- clade B. There is a logical relationship between brae, the frontal, the tooth insertion on a Lamina Etheridge’s scheme and the phylogenetic hypoth- subdentalis and a distinct heterodonty (mono- esis presented here (see Figure 16). In particular, and tricuspid teeth)” (Rossmann 1999b:586). Groups 1 and 3 were characterized not only by 1. Only one of the proposed diagnostic features plesiomorphic states of characters transformed of “Messelosaurinae,” the state of the Meckelian in the other group, but by uniquely derived and groove, can be evaluated in all included taxa. explicitly recognized apomorphies. In contrast, 2. The location of the parietal foramen at Group 2 was characterized principally by ple- the frontoparietal suture distinguishes “mes- siomorphies for Groups 1 and 3, or by apomor- selosaurines” from extant corytophanines (except phic features he considered to be multiply Laemanctus), in which the foramen is located in derived or to be shared with one or the other the frontal. However, because the sutural location group. This character distribution is comprehen- of the foramen is a synapomorphy of iguanians sible insofar as most Group 2 taxa are basal to (e.g., Estes et al. 1988) and thus primitive for the Group 3 taxa in the phylogeny presented here. (It lineage leading to Corytophaninae, this feature is not out of the question that Phrynosomatinae offers no evidence of particular relationship 256 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 within Iguania. It is merely consistent with, rather plateau” (Rossmann 1999a). Rossmann than indicative of, Geiseltaliellus and Aciprion (2000:128) reported that the neural spines are (but not other “messelosaurines,” where it is broadened in Geiseltaliellus from Messel, but unknown) being on the stem of Corytophaninae. remarked that no zygosphene– zygantral articula- 3. Within Iguania a median parietal crest is tions could be shown. Accessory vertebral articu- present at least in Geiseltaliellus as well as in Cory- lations are known in several iguanian groups (see tophaninae, some iguanines (de Queiroz 1987) Appendix 3). The small “plateau” is common in and many Anolis (Etheridge 1959). It is not pres- Iguania (Smith pers. obs.) and provides little evi- ent in the only other member of “Messelosauri- dence of relationship. nae” in which the feature can be evaluated, 8. The “similar morphology” of the frontal in Aciprion formosum (as Rossmann noted). Thus, Geiseltaliellus and Cadurciguana hoffstetteri so far as has been shown, a median parietal crest might refer to the following features mentioned is unique to Geiseltaliellus among proposed mem- by Augé (1987) and/or Rossmann (1999b): bers of “Messelosaurinae.” The parietal table (that absence of sculpturing on the dorsal surface of the is, not simply the parietal excluding the supratem- frontal, an hourglass shape, and the presence of a poral processes) in Geiseltaliellus is not trape- “frontal shelf” supporting the nasal. All are prim- zoidal, but in Aciprion it is. A trapezoidal parietal itive features common to many or all iguanians. table is plesiomorphic for Iguania (Etheridge and 9. A “lamina subdentalis ( subdental shelf)” de Queiroz 1988). (Rossmann 1999a:189) is potentially useful, but it 4. An open Meckelian groove has been con- is present in at least some living corytophanines sidered primitive for Squamata and offers no evi- and certain other iguanids and acrodontans dence of particular relationship within the group. (Smith 2006a, 2006b). It has not been carefully It is present in several other iguanian taxa (see examined in other proposed members of “Mes- Appendix 3). Additionally, the Meckelian groove selsaurinae.” is fused in Cadurciguana hoffstetteri (Augé 1987; 10. Heterodonty, as defined above, is common Rossmann 1999a). Finally, the Meckelian sulcus to virtually all iguanians (see, e.g., Hotton 1955). of Geiseltaliellus maarius and G. longicaudus is In summary, no clear feature has been (briefly) closed in at least some specimens, adduced to support the particular relationship of whereas that of ?Crotaphytus oligocoenicus is any included taxon (except Geiseltaliellus) to open, if restricted (Holman 1972). Corytophaninae, much less show the monophyly 5. A posterolateral process of the coronoid is of “Messelosaurinae.” “Messelosaurines” share no present in Geiseltaliellus maarius and G. longi- similarities of presence or absence with one caudus. It is a derived feature in Iguania (Lang another that are not found in other iguanids; 1989). The feature is not present in the only other moreover, in cases in which characters can be member of “Messelosaurinae” in which it can evaluated in more than one taxon, they conflict. presently be evaluated, Aciprion formosum Thus, it is recommended that the name “Mes- (YPM-PU 10015, Smith pers. obs.). Thus, so far as selosaurinae” be avoided. is known, a posterolateral process is unique to Geiseltaliellus among proposed members of Mode of Life of Geiseltaliellus “Messelosaurinae.” 6. A medial centrale, as distinct from a first dis- Living corytophanines spend considerable time off tal carpal, is absent in squamates (Renous-Lécuru the ground. Members of Corytophanes (Köhler 1973). The hand is only reasonably well known in 2000) and Laemanctus (Martin 1958) are often Geiseltaliellus maarius among proposed members found in trees, or flee to them when encountered of “Messelosaurinae,” and in this species there is on the ground; members of Basiliscus perch in no separate medial centrale (see above). vegetation and on objects lying on the ground 7. The only remarkable features of the pre- (Hirth 1963; Vaughan et al. 2007). (Juveniles of sacral vertebrae attributed to Cadurciguana hoff- Basiliscus, however, are more terrestrial; Hirth stetteri are zygosphene–zygantral accessory 1963.) However, terrestrial bipedal locomotion articulations and a broadening of the dorsal ter- has been documented in each genus (Coryto- mini of the neural spines, “forming a small phanes and Basiliscus, Snyder 1952; Laemanctus, Eocene Lizards of the Clade Geiseltaliellus • Smith 257

McCoy 1968). As Losos (1990) emphasized, short The two propensities, as in Corytophaninae, are arms, long legs and a long tail are expected to be not mutually exclusive. of benefit to both arboreal hoppers and terrestrial bipeds. Thus, the presence of these characteristics Iguanid Biogeography in Geiseltaliellus and its living relatives does not offer conclusive evidence on their habitat The early and middle Eocene were times of great preferences. warmth at high latitude (e.g., Markwick 1998). Relative phalangeal length provides addi- The diminution of latitudinal temperature gradi- tional evidence bearing on the mode of life of ents (Greenwood and Wing 1995) seems to be members of Geiseltaliellus (thanks to J. A. Gau- responsible, in part, for the intercontinental dis- thier for pointing this out, pers. comm.). Hop- persal of several terrestrial taxa over many mil- son (2001) found that within living theropods lions of years (e.g., Krishtalka et al. 1987; Beard (birds), arboreal taxa (and others adapted for 1998). Among these taxa are glyptosaurine grasping) show elongated penultimate pha- anguids (Meszoely et al. 1978) and the heloder- langes, which constitute 40% to 60% of the total, matid Eurheloderma (Hoffstetter 1957) of nonungual phalangeal length. Geiseltaliellus Europe, and the acrodont iguanian Tinosaurus of maarius also has long penultimate phalanges (see both North America (Estes 1983b; Smith 2006b) Table 1). Lateral digits (III, IV and V) especially and Europe (Augé 1990a, 2003, 2005). Early show a greater disparity in length between Eocene migration between North America and penultimate and ultimate or antepenultimate Europe could have taken place through Green- phalanges. The simplest explanation is that the land, when the ocean basins separating it from the manus was used in locomotion for grasping, and two continents were narrower (McKenna 1983). the forested nature of the middle Eocene land- Oplurinae, as noted in the introduction, is one scape around Messel (Wilde 1989) in turn sug- of the only clades of Iguanidae outside the New gests locomotion in vegetation. I therefore World and the one whose occurrence is least sat- conclude that Geiseltaliellus, like its inferred isfactorily explained. Because neither Geiseltaliel- closest living relatives (corytophanines, most lus nor Corytophaninae seems closely related to Anolis, and Polychrus), spent considerable time Oplurinae, the presence of the first in Europe has in vegetation. no bearing on the presence of the last in Madagas- There is no evidence to suggest that the habit car. Corytophaninae seems to be primitively of the of Basiliscus of running across water (e.g., Laerm New World, where all its modern representatives 1973; Snyder 1949) is not autapomorphic of that (Maturana 1962) and its living sister-taxon (Poly- clade. Nevertheless, Rossmann (2001) stated that chrotinae*), according to this analysis, reside. The the locomotory repertoire of Geiseltaliellus prob- stem of Corytophaninae is first known in North ably included supra-aqueous running or swim- America (Smith 2009). The dispersal of a member ming. As discussed above, the evidence for of the stem of Corytophaninae from North Amer- webbing in Geiseltaliellus is inconclusive. Further- ica to Europe is a reasonable interpretation of these more, Basiliscus lack webbing between the toes, biogeographic data. Geiseltaliellus constitutes yet relying instead on rectangular fringes (Luke 1986) another example of intercontinental dispersal, and on the postaxial margin of each toe for increased an evolutionary dead-end, as Estes (1983b), with- surface area (Laerm 1973). Thus, there is no evi- out an explicit phylogeny, previously supposed and dence to support aquatic locomotion of any kind Augé (2005) further maintained. As for the timing in Geiseltaliellus. of migration, the Geiseltaliellus lineage, though best While facultative bipedalism in Geiseltaliellus known from the middle Eocene of Germany, cannot be excluded, the elongated antepenulti- might be represented by jaw fragments from the mate phalanges, in combination with the inferred very early Eocene of Belgium and France (Augé Eocene environment of the area around Messel, 1990a, 1990b) and is certainly present by standard point least ambiguously to arboreal habits, as is level MP 10 (Augé 2003). Geiseltaliellus conceiv- probably primitive for Polychrotinae* and Cory- ably dispersed to Europe in association with rapid tophaninae based on the habitat preferences of global warming at the Paleocene–Eocene bound- living members of these groups (Etheridge 1959). ary (see review in Gingerich 2006). 258 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

The sister-group relationship between (with P. gutturosus found also in Central America: Tropidurinae* and Oplurinae inferred in this Köhler 2000). study is consistent with the hypothesis of an Even if the results of the phylogenetic analy- ancient vicariance event, as previously suggested ses presented here are accurate, they do not much (e.g., Estes 1983b; Noonan and Chippindale help constrain the geographic origin of Iguanidae 2006). However, if this event is related to the (Estes and Price 1973) beyond being New World. break-up of Africa and South America, this But they do allow hypotheses to be drawn on the divergence should be very old (Early Creta- subsequent biogeographic history of the group. ceous), and Hugnall et al.’s (2007) divergence In particular, it is possible that Clade A, if it exists, time estimates suggest that this divergence can- is an ancestrally North American one. Fossil cro- not be earlier than Late Cretaceous. If, on the taphytines dating from the latest Eocene are other hand, Oplurinae represents dispersal from North American (Holman 1972; Estes [1983a], the south (Blanc 1982) or vicariance related to however, doubted their attribution). The wide- the separation of Antarctica and Indo-Madagas- spread presence of Iguaninae in tropical South car, a more recent (perhaps Late Cretaceous) America is due exclusively to the broad distribu- divergence time would be predicted (cf. Hay et tion of one species, Iguana iguana (de Avila Pires al. 1999; de Wit 2003). Unfortunately, the Creta- 1995), which could have entered that continent ceous fossil records of Madagascar and Antarc- during the faunal exchange that began in the tica are impoverished for squamates. The one middle Miocene and culminated in the Pliocene– Cretaceous lizard known from Madagascar Pleistocene Great American Biotic Interchange seems to be a cordyliform, not an iguanian (see review in Woodburne 2004). The iguanines (Krause et al. 2003). In any case, published diver- Amblyrhynchus and Conolophus are found on the gence time estimates, which are congruent with equator west of Ecuador, but the Galápagos the latter hypothesis (Noonan and Chippindale Islands lie in the intertropical convergence zone, 2006), await better fossil calibration points from where currents of the North Pacific Gyre sweep within Iguanidae. south and west from Central America. The earli- The long-standing problem of the origin of est fossil record of Iguaninae is in North America Oplurinae (Boulenger 1918) is only one of the (Norell and de Queiroz 1991). The earliest record many difficulties in understanding iguanid bio- of the corytophanine stem is from Wyoming geography. Crotaphytinae is exclusively found in (Smith 2009). Polychrus may be an essentially the temperate zones of North America. Phryno- South American clade today, but its earliest rep- somatinae is also entirely North American, but resentatives are from the Eocene of North Amer- some clades (e.g., Sceloporus) have representatives ica (Smith 2006a; manuscript in preparation); in tropical North America. Two clades, Iguaninae thus, its (near-)restriction to South America may and Corytophaninae, are primarily North Amer- be a post-Eocene phenomenon. Stem relatives of ican, but have members with partly South Amer- Anolis might also be present in the early Eocene of ican and, in the case of Iguaninae Antillean, Wyoming (Smith 2009; manuscript in review), distributions. Tropidurinae* is predominantly which is not to say that the genus is not long- South American, but the genus Leiocephalus is established in South America. Conrad et al. (naturally) restricted to the Antilles. Hoplocerci- (2007) additionally suggested that another early nae is primarily South American, but species in Eocene species is related to Anisolepini. (Notably, two genera are found north of the Isthmus of however, this latter species shows the Anolis-type Panama (Köhler 2000). Polychrotinae* has per- relationship between autotomy septa and tran- haps the most complicated distribution (conceiv- verse processes in the caudal vertebrae [Conrad ably partly reflecting nonmonophyly): Anisolepini et al. 2007].) If this portrait is accurate, then the and Leiosaurini are exclusively South American; polychrotine clades Anisolepini and Leiosaurini, Anolis is widespread in tropical North America, which are found only in the southern hemisphere the Antilles, and South America, and probably dis- of South America, possibly indicate an early dispersed more than once between the Caribbean persal from North America, and Hoplocercinae islands and the mainland (Poe 2004); and Poly- might show the same post-Eocene tropical chrus today is nearly confined to South America restriction as Polychrus. Eocene Lizards of the Clade Geiseltaliellus • Smith 259

The history of Clade B is not especially well prove particularly useful in the identification of constrained by the results of the phylogenetic fossil remains, which have thus far consisted pri- analysis, although the sister-group relationship marily of dentigerous elements. (On the other between Oplurinae and Tropidurinae* inferred in hand, the latter will be useless.) There is also mor- this and previous analyses (Titus and Frost 1996; phological evidence for the division of Iguanidae Conrad 2008) is consistent with an ancient diver- into two clades. One of these contains Crotaphyti- gence between South American and Malagasy nae, Iguaninae + Hoplocercinae, and Coryto- taxa (Noonan and Chippindale 2006) and finds phaninae + Polychrotinae*. It may be ancestrally further support in the possible occurrence of North American, where it had a broad distribu- (stem) tropidurines in the Paleocene of Brazil tion in the Eocene. Yet persistent conflict between (Estes 1983b). Clearly, new fossils from the New morphological and molecular data, which yield World, preferably from the Paleogene and Creta- wildly different pictures of iguanian phylogeny, ceous, will be necessary to test this biogeographic must be worked out. portrait. A recently reported iguanian frontal from the medial Cretaceous of Argentina Acknowledgments (Apesteguía et al. 2005) is auspicious and indicates that we have more to await from South I am deeply indebted to Eberhard “Dino” Frey for America. the opportunity to study with him in Karlsruhe. I thank H. Haubold, H. Heinisch, M. Hellmund, N. Conclusions Micklich and S. Schaal for access to fossil specimens. C. J. Bell, J. A. Gauthier, M. Kearney, G. The Eocene lizard taxon Geiseltaliellus represents Köhler, K. Krysko, J. Losos, S. Rogers, J. Vindum, an incursion of Iguanidae into the Old World that H. Voris and G. Watkins-Colwell provided access probably occurred during the greenhouse climate to modern specimens. I am grateful to S. of the earliest Eocene. Although specimens from Marschall and C. Kurz for their hospitality dur- the early Eocene of France and Belgium have been ing my stays in Frankfurt. T. Rossmann kindly referred to Geiseltaliellus (Augé 1990a, 1990b, made available casts of the private specimen of 2003), the taxon is best known from the middle Geiseltaliellus maarius. I am appreciative of C. J. Eocene of Germany. The two known middle Bell, B.-A. Bhullar, M.-C. Buchy, E. Frey, J. A. Eocene species form a clade united by the com- Gauthier, J. Habersetzer, N. Micklich, M. Rück- mon presence of a posterolateral process of the lin, S. Schaal, D. Schreiber and T. Rowe for dis- coronoid. Geiseltaliellus is attributable to the stem cussion; J. A. Gauthier in particular recognized of Corytophaninae, sharing several derived fea- the significance of the elongated penultimate pha- tures with that clade, some of which are modified langes in G. maarius. M.-C. Buchy read a primi- within the crown (e.g., invariant dorsal expansion tive draft and made helpful suggestions. K. de of the postorbital and a groove at the prefrontal– Queiroz, R. M. Sullivan, G. Watkins-Colwell and lacrimal junction). Geiseltaliellus is most similar R. Etheridge provided constructive critiques of to Basiliscus in squamation and the ontogeny of the final version. A. Hebs took the photographs the parietal, suggesting that these features are in Figures 3A, 4A, 7A and 10; W. Fuhrmannek primitive for Corytophaninae. All specimens of took the photographs in Figures 2A and 9A; and Geiseltaliellus lack the parietal blade characteristic V. Griener took the photograph in Figure 11A. T. of the crown clade. Colantonio drew Figures 20C, D and 22. Finan- Morphological characters support the mono- cial support came successively from the Baden- phyly of Iguanidae. Chief among these are (1) the Württemberg Scholar Exchange Program, a Yale presence of separate subnarial arterial and ante- University Fellowship, a National Science Foun- rior inferior alveolar foramina on the premaxil- dation Doctoral Dissertation Improvement lary process of the maxilla and (2) the presence of Grant, and the Texas Memorial Museum at the an anterior projection of the ventral margin of the University of Texas at Austin. cartilaginous planum antorbitale (processus rostralis plani antorbitalis). These show little reversal Received 18 November 2008; revised and accepted and little or no convergence. The former should 25 March 2009. 260 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Appendix 1: Morphometric Data on Body Proportions in Selected Iguanid Lizards Parentheses indicate lot-catalogued specimens. Institutions: HLMD-Me, Messel Collection, Hessisches Landesmu- seum, Darmstadt, Germany; SMF, Herpetology Collection, and SMF ME, Messel Collection, Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany; TMM, Texas Memorial Museum, The Uni- versity of Texas, Austin, Texas USA; YPM-VP, Division of Vertebrate Paleontology, and YPM-VZ, Division of Vertebrate Zoology, Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA. Abbre- viations: AA, arm length, measured from axil to the end of metacarpal III in wet specimens; LL, leg length, measured from groin to the end of metatarsal IV in wet specimens; SVL, snout–vent length; TL, tail length. All measurements in centimeters. Species name Institution Specimen SVL TL LL TL/SVL LL/SVL AL AL/LL

Corytophaninae Laemanctus longipes SMF a 13.9 52.8 9.7 3.80 0.70 5.0 0.52 L. serratus SMF 11016 11.5 38.6 7.9 3.36 0.69 5.0 0.63 Basiliscus basiliscus SMF 50919 19.4 — 13.3 — 0.69 6.2 0.47 B. basiliscus SMF 50920 15.7 — 11.3 — 0.72 5.1 0.45 B. basiliscus SMF 50921 14.0 — 9.9 — 0.71 4.8 0.48 B. basiliscus SMF 50922 11.2 — 8.1 — 0.72 3.7 0.46 B. basiliscus SMF 50923 8.5 22.6 6.9 2.66 0.81 3.0 0.43 B. basiliscus SMF 50924 8.4 20.8 6.1 2.48 0.73 2.8 0.46 B. basiliscus SMF 50925 7.7 20.3 6.2 2.64 0.81 2.6 0.42 B. galeritus SMF 11029 18.8 47.9 13.6 2.55 0.72 6.3 0.46 B. plumifrons SMF 24882 18.6 40.8 13.1 2.19 0.70 6.1 0.47 B. vittatus SMF 11019 6.1 14.2 — 2.33 — — — B. vittatus SMF 53574 8.4 22.1 6.2 2.63 0.74 2.8 0.45 B. vittatus SMF 42112 11.4 27.1 8.8 2.38 0.77 3.8 0.43 B. vittatus SMF 42108 12.1 26.9 9.2 2.22 0.76 4.2 0.46 B. vittatus SMF 42119 12.9 37.1 10.5 2.88 0.81 4.3 0.41 B. vittatus SMF 53684 13.7 35.5 10.5 2.59 0.77 4.3 0.41 B. vittatus SMF 52060 15.9 42.7 12.0 2.69 0.75 4.8 0.40 B. vittatus SMF 47207 8.7 22.7 6.7 2.61 0.77 2.8 0.42 B. vittatus SMF 42113 10.0 27.0 7.6 2.70 0.76 3.1 0.41 B. vittatus SMF 52055 11.7 29.2 8.7 2.50 0.74 3.5 0.40 B. vittatus SMF 42109 12.1 31.9 8.7 2.64 0.72 3.6 0.41 B. vittatus SMF 48080 13.1 34.5 9.8 2.63 0.75 3.9 0.40 Corytophanes cristatus SMF 81504 8.5 18.2 6.5 2.14 0.76 3.2 0.49 C. hernandesii SMF 11015 8.5 21.6 6.3 2.54 0.74 3.4 0.54 C. percarinatus SMF 80733 9.6 21.1 7.2 2.2 0.75 3.9 0.54 C. percarinatus SMF 80734 10.6 20.1 7.2 1.9 0.68 4.1 0.57 C. percarinatus SMF 80735 9.6 18.1 6.7 1.89 0.70 3.8 0.57 C. percarinatus SMF 80736 9.0 19.1 6.8 2.12 0.76 3.9 0.57 C. percarinatus SMF 80737 3.5 6.1 2.7 1.74 0.77 1.5 0.56 C. percarinatus SMF 80738 3.2 5.7 2.7 1.78 0.84 1.4 0.52

Hoplocercinae Enyalioides heterolepis SMF 11038 9.1 12.8 6.3 1.41 0.69 3.2 0.51 “E. laticeps” SMF 72624 11.1 16.9 6.2 1.52 0.56 4.3 0.69 “E. laticeps” SMF 72625 10.4 14.7 6.2 1.41 0.60 4.0 0.65 “E. laticeps” SMF 72626 7.6 10.4 4.4 1.39 0.58 2.8 0.64 E. oshaughnessyi SMF 75804 13.9 19.6 7.7 1.41 0.55 5.0 0.65 Continued Eocene Lizards of the Clade Geiseltaliellus • Smith 261

Appendix 1 continued

Species name Institution Specimen SVL TL LL TL/SVL LL/SVL AL AL/LL

Morunasaurus annularis SMF 78053 13.6 — 5.7 — 0.42 3.7 0.65 M. annularis SMF 78054 10.6 — 4.9 — 0.46 3.2 0.65 M. annularis SMF 78055 8.7 — 4.2 — 0.48 2.7 0.64 M. annularis SMF 78056 6.9 8.9 3.3 1.29 0.48 2.2 0.67 Hoplocercus spinosus SMF 62417 10.9 4.3 4.1 0.39 0.38 3.1 0.76

Iguaninae Brachylophus fasciatus SMF 73361 15.7 — 9.0 — 0.57 5.9 0.66 B. fasciatus SMF 73362 16.1 34.1 8.0 2.18 0.50 5.7 0.71 B. fasciatus SMF 73363 16.9 59.6 8.4 3.53 0.50 — — B. fasciatus SMF 73364 16.0 40.7 8.7 2.54 0.54 5.8 0.67 Dipsosaurus dorsalis SMF 65652 11.8 — 6.7 — 0.57 3.6 0.54 D. dorsalis SMF 65653 10.3 20.6 6.4 2.00 0.62 3.4 0.53 D. dorsalis SMF 65654 9.5 19.9 5.3 2.09 0.56 2.9 0.55

Polychrotinae* Anisolepis undulatus SMF b 5.5 13.8 2.7 2.51 0.49 — — A. undulatus SMF b 6.9 16.8 — 2.43 — — — Anolis cobanensis SMF 82688 5.0 11.5 3.8 2.3 0.76 — — A. cristatellus YPM-VZ R12074 6.4 11.2 4.3 1.75 0.67 — — A. cuvieri YPM-VZ R12066 13.5 — 8.7 — 0.64 — — A. distichus YPM-VZ 3993 5.1 7.4 3.3 1.45 0.65 — — A. polylepis YPM-VZ 1866 3.8 8.7 2.2 2.29 0.58 — — A. richteri SMF 60817 6.2 8.4 3.3 1.35 0.53 2.0 0.61 A. richteri SMF 60820 7.3 10.9 — 1.49 — — — A. ricordii SMF 24859 13.4 24.2 7.4 1.82 0.55 — — A. wetmorei YPM-VZ 1866 3.8 8.7 2.2 2.29 0.58 — — Diplolaemus bibronii SMF 58523 8.3 7.8 3.9 0.94 0.47 2.4 0.62 D. bibronii SMF 58524 9.4 8.5 4.4 0.9 0.47 2.7 0.61 Enyalius bilineatusc SMF 11054 8.5 25.3 5.4 2.94 0.63 3.1 0.57 E. catenatus SMF 11040 8.8 19.2 6.5 2.18 0.74 3.7 0.57 E. catenatus SMF 11041 9.9 23.2 5.8 2.34 0.59 4.1 0.71 Leiosaurus bellii SMF 11066 9.8 10.3 4.5 1.05 0.46 3.4 0.76 L. bellii SMF 11067 8.2 8.6 3.9 1.05 0.48 2.3 0.59 Polychrus acutirostris SMF 62421 11.0 21.2 4.0 1.93 0.36 3.1 0.78 P. marmoratus SMF d 8.4 25.9 4.3 3.08 0.51 — — P. marmoratus SMF d 10.1 29.6 4.4 2.93 0.44 2.8 0.64 Pristidactylus torquatus SMF 43945 10.4 15.6 5.3 1.5 0.51 — — Urostrophus vautieri SMF 11063 6.1 9.3 — 1.52 — — — U. vautieri SMF e 7.5 11.0 — 1.47 — — — U. vautieri SMF e 6.7 8.9 2.8 1.33 0.42 — —

Tropidurinae* Leiocephalus cubensis YPM-VZ 6652 5.6 9.4 3.4 1.68 0.61 — — L. cubensis YPM-VZ 1161 3.6 5.9 2.1 1.64 0.58 — — L. schreibersii YPM-VZ 3233 7.1 — 3.9 — 0.55 — —

Continued 262 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Appendix 1 continued

Species name Institution Specimen SVL TL LL TL/SVL LL/SVL AL AL/LL

L. stictigaster YPM-VZ 642 5.9 10.5 4.2 1.78 0.71 — — Liolaemus pictus SMF (30284) 3.2 6.3 1.5 1.95 0.46 — — L. pictus SMF (30284) 4.8 — 2.1 — 0.44 — — L. pictus SMF (30284) 4.8 7.8 2.0 1.62 0.42 — — L. pictus SMF (30284) 6.7 9.4 2.5 1.40 0.37 — — Microlophus albemarlensis YPM-VZ 2005 10.6 — 6.3 — 0.59 — — M. occipitalis SMF 70762 5.2 — 2.9 — 0.56 — — M. occipitalis SMF 70763 6.5 12.8 3.8 1.97 0.58 — — Phymaturus palluma SMF 83678 4.9 4.5 2.1 0.92 0.43 — — P. palluma SMF 83677 4.5 4.5 2.0 1.0 0.44 — — P. palluma SMF 83675 8.2 8.8 3.8 1.1 0.46 — — P. palluma SMF 83676 9.1 8.4 3.6 0.92 0.39 — — Plica plica SMF 76307 5.5 — 3.6 — 0.65 — — P. plica SMF 76306 9.8 — 6.4 — 0.65 — — P. plica SMF 76305 10.5 — 7.3 — 0.70 — — P. plica SMF 76304 12.4 20.6 8.6 1.66 0.69 — — Tropidurus peruvianus YPM-VZ 7122 9.7 — 5.9 — 0.61 — — T. torquatus YPM-VZ R13186 10.5 — 7.4 — 0.70 — — Uranoscodon superciliosus YPM-VZ R11661 11.8 — 6.7 — 0.57 — —

Oplurinae Chalarodon madagascariensis YPM-VZ R14576 5.6 — 4.1 — 0.73 — — C. madagascariensis YPM-VZ R14577 6.9 — 4.7 — 0.68 — — Oplurus fierinensis YPM-VZ R13412 9.6 — 6.1 — 0.64 — — O. grandidieri YPM-VZ R13414 9.0 — 5.8 — 0.64 — — O. quadrimaculatus YPM-VZ R13415 11.9 23.4 7.8 1.97 0.66 — — O. quadrimaculatus YPM-VZ R13416 9.1 18.1 5.9 1.99 0.65 — —

Crotaphytinae Crotaphytus collaris YPM-VZ 1147 7.4 15.2 5.8 2.05 0.78 — — C. collaris YPM-VZ 1220 6.4 12.5 4.9 1.95 0.77 — — C. collaris YPM-VZ 1144 9.3 18.4 7.1 1.98 0.76 — — Gambelia wislizenii YPM-VZ 7135 11.4 — 7.4 — 0.65 — — G. wislizenii YPM-VZ 7818 9.2 19 6.4 2.07 0.70 — —

Phrynosomatinae Callisaurus draconoides YPM-VZ 646 7.8 — 5.6 — 0.72 — — C. draconoides YPM-VZ 2832 3.0 4.2 2.1 1.40 0.70 — — Cophosaurus texanus YPM-VZ 911 5.8 6.5 4.0 1.12 0.69 — — C. texanus YPM-VZ 1007 6.8 7.9 4.9 1.16 0.72 — — C. texanus YPM-VZ 697 5.5 6.3 3.9 1.15 0.71 — — Holbrookia maculata YPM-VZ 1014 5.3 — 3.0 — 0.57 — — H. maculata YPM-VZ 1018 5.6 6.9 3.4 1.23 0.61 — — Petrosaurus thalassinus YPM-VZ R14664 8.9 16.4 5.3 1.84 0.60 — — Phrynosoma cornutum YPM-VZ R13395 9.4 4.4 5.1 0.47 0.54 — — Sceloporus magister YPM-VZ 919 9.3 12.2 4.6 1.31 0.49 — —

Continued Eocene Lizards of the Clade Geiseltaliellus • Smith 263

Appendix 1 continued

Species name Institution Specimen SVL TL LL TL/SVL LL/SVL AL AL/LL

S. malachiticus YPM-VZ R13003 9.8 — 4.4 — 0.45 — — S. malachiticus YPM-VZ R13004 8.1 8.7 4.1 1.07 0.51 — — S. malachiticus YPM-VZ R13005 6.4 8.1 3.3 1.27 0.52 — — S. undulatus SMF (41657) 4.5 5.9 2.4 1.3 0.54 — — S. undulatus SMF (41657) 5.0 6.7 2.6 1.3 0.52 — — S. undulatus SMF (41657) 5.7 7.9 3.0 1.4 0.52 — — S. variabilis YPM-VZ R14005 3.5 — 1.7 — 0.49 — — S. variabilis YPM-VZ R14006 5.1 7.5 2.8 1.47 0.55 — — S. variabilis YPM-VZ R14008 7.4 9.2 4.2 1.24 0.57 — — Uma notata YPM-VZ 649 6.4 8.8 4.6 1.38 0.72 — — Urosaurus microscutatus YPM-VZ R6692 3.8 5.7 1.6 1.50 0.42 — — Uta stansburiana YPM-VZ R2851 3.6 6.8 2.1 1.89 0.58 — — U. stansburiana YPM-VZ R2846 4.5 — 2.7 — 0.60 — —

Geiseltaliellusf G. maarius HLMD-Me 10207 8.6 18.4 5.2 2.14 0.60 2.9 0.56 G. maarius SMF ME 2 7.3 — 5.5 — 0.75 2.8 0.51 G. maarius SMF ME 1769 9.8 — 7.4 — 0.76 3.5 0.47 G. maarius SMF ME 2938 6.8 13.4 4.7 1.97 0.69 2.4 0.51 G. maarius SMF ME A332 8.8 27.8 7.3 3.16 0.83 — — G. longicaudus GM 4146 6.8 21.5 — 3.16 — — — a Either 11017 or 11018 (specimen with jaws closed). b One specimen is 11055, the other 11056. c Catalogued as Enyalius fitzingeri (see Etheridge 1969). d One specimen is 54351, the other 54352. e One specimen is 11064, the other 11065. f All measurements are obviously of the respective bony elements. 264 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Appendix 2: Data Matrix Used in Examination of Iguanian Relationships The “early Eocene iguanid” is described by Smith (2009) as Suzanniwana patriciana, who argues that it is a stem- corytophanine.

#NEXUS BEGIN DATA; DIMENSIONS NTAX = 39 NCHAR = 152; FORMAT SYMBOLS = “ 0 1 2 3 “ MISSING = ? GAP = - INTERLEAVE ; MATRIX [ 10 20 30 40 50] [.....] Polychrus_acutirostris 0011011011 10?0000000 0110000110 1110100011 0011111002 Pristidactylus_torquatus 1010000001 00?0000000 0010000000 2010100010 00110110(01)2 Anolis_ricordi 0011000001 1100000000 0010000011 10101100?? 0011111001 Anolis_cristatellus 0000000000 1100000000 0010000010 201011000? 0011110001 Basiliscus_basiliscus 0111000001 10?0001100 0110001001 1?1?110011 0?11110003 Corytophanes_cristatus (01)011000001 00?0001100 1110000011 1?12110021 0111110003 Laemanctus_longipes 0001000001 00?0001100 0010001001 1010110011 00111110(01)(23) Geiseltaliellus_maarius 1?10000001 10??00??00 01100?100? ?0111(01)?010 0?11110002 early_Eocene_iguanid 1010?00001 ?0??00(01)000 011?0?1?01 ?010?00?10 0?111(01)0002 Crotaphytus_collaris 1000000001 00?0011110 0000000000 1?12100110 0011001002 Gambelia_wislizenii 1000000001 00?0010110 0000000000 1?12100110 0011000002 Dipsosaurus_dorsalis 1101000001 00?0000110 0111000000 2010101111 0011000003 Brachylophus_fasciatus 1110000001 00?000?110 011100000? 0010101011 0011000002 Iguana_iguana 1111000001 10?0000110 0111000000 1010101011 0011000002 Enyalioides_oshaughnessyi 0011010011 00?000?000 011001000? 1010101011 0111101002 Oplurus_quadrimaculatus ??1?00?00? 0??001??00 010000000? 2?1?11100? 0011000002 Chalarodon_madagascariensis 2101000001 00?001?110 010000100? 2?12101010 0011000002 Stenocercus_scapularis 1000000000 011001??00 010000000? 2010100010 001111010? Microlophus_occipitalis 2000000001 ?110010010 010?000000 201010100? 0?11010001 Tropidurus_torquatus 3000001001 ?110010010 010?00?000 201010100? 0011000001 Plica_umbra 2011001000 0110010010 010?00?000 201010100? 0111000001 Phymaturus_palluma 3101000001 ?0?0000010 010???0?00 2???101?0? 0?11000002 Liolaemus_pictus 2000000001 ?0?0010010 010?????00 2??0111?0? 0?11000002 Leiocephalus_personatus 2000101001 00?0000110 0100000000 201010100? 001100000(23) Phrynosoma_platyrhinos 200100?001 00?0110000 1100100??? 2210101021 0111011002 Petrosaurus_thalassinus 1000001001 011001?010 010000000? 201010100? 00110000?2 Sceloporus_undulatus 2000000001 0110010000 0100?00?00 2?1?10100? 00110000?? Physignathus_cocincinus 3000000100 00?1000101 0000010000 1?1210010? 001111000? Agama_agama 3001000100 0??1010101 000001000? 1?1210010? 0011100012 Priscagama_gobiensis 300?00?10? 00?101??10 000000000? 11?0100110 0?11101002 Leiolepis_belliana 30010001?0 00?111?110 010000000? ??12100110 0011(01)0011? Brookesia_superciliaris 300010?110 00?1110100 0110110?1? 2????0002? 0011111003 Saichangurvel_davidsoni 00000000?0 00??????10 010000?00? 000000?0?? 1?110?00?1 Sphenodon_punctatus 201000?000 00?0000000 0010100?0? 000010?00? 00001000?0 Diphydontosaurus_avonis 000000?000 ?0??00?00? 000?????0? 000000??0? ??00?000?0 Eublepharis_macularius 0000000000 00?0010000 0?1000000? 1000?????? 101101010? Plestiodon_fasciatus 1000001000 00?0010001 0010001001 000000?00? 1000001000 Elgaria_multicarinata 0000001000 00?0000000 0010001?00 000000?010 1000011000 Hypothetical_ancestor 0000000000 00?00?0000 0000000000 000000?00? ?0????0000

Continued Eocene Lizards of the Clade Geiseltaliellus • Smith 265

Appendix 2 continued

[ 60 70 80 90 100] [..... ] Polychrus_acutirostris 10100?0011 1100011100 00210001?? 0120011101 01022?1?00 Pristidactylus_torquatus 10000?0000 100100???? ?????????? 0120000101 0002111110 Anolis_ricordi 12100?0001 110020???? ?????????? 01?0101111 0002201111 Anolis_cristatellus 12000?0010 000020?100 00211011?? 0120101111 0002201111 Basiliscus_basiliscus 1200100000 1100101000 02100000?? 0020000201 0002111000 Corytophanes_cristatus 1210111011 1100001000 02110000?? 1020110201 ?002210000 Laemanctus_longipes 1210110011 110000???? ?????????? 10?0000101 0002111000 Geiseltaliellus_maarius ?2000??000 ?00000???? ?????????? ???0??0101 00011?1?01 early_Eocene_iguanid 12000????? ?????0???? ?????????? ???????101 000111???0 Crotaphytus_collaris 00000?0000 0001002100 0111010021 00000??111 0100011000 Gambelia_wislizenii 00000?0000 100100???? ?????????? 00?0000101 0001111000 Dipsosaurus_dorsalis 00000?0100 000010???? ????????21 00?0000202 1002201001 Brachylophus_fasciatus 00000?0100 1000102100 0101010011 0010000202 10022?1?01 Iguana_iguana 12000?0100 1000101100 0001010011 00?00??202 1002201001 Enyalioides_oshaughnessyi 00000?0000 011000???? ?????????? 00?001020(12) 1000011001 Oplurus_quadrimaculatus ?0000?0000 0101001100 0111010121 00000??101 00011???10 Chalarodon_madagascariensis 00000?0000 000000???? ?????????? 00000??101 0002111100 Stenocercus_scapularis ?0000?0000 000000???? ?????????? 00?0000101 00022?1?10 Microlophus_occipitalis 00000?0?00 000000???? ?????????? 00?0010101 0002201010 Tropidurus_torquatus 00000?0?00 000000???? ?????????? 00?0000201 0002201010 Plica_umbra 00000?0?00 000001???? ????????10 00?0000101 0002201110 Phymaturus_palluma 20000?0010 00?0002000 0111010021 0020010201 0000001001 Liolaemus_pictus 20000?0000 00?000???? ?????????? 00?0001101 0002201111 Leiocephalus_personatus 00000?0000 0000012000 0101010021 0010000201 0002201001 Phrynosoma_platyrhinos 20100?0010 0100?10?00 1021010022 00200??000 0000001110 Petrosaurus_thalassinus 00100?0000 0000011100 1021010022 0010010101 00011?1?00 Sceloporus_undulatus 00000?0000 000021010? ?021000022 00?00??101 0001101100 Physignathus_cocincinus 00000?0000 1100010011 0100000000 0121000?10 ?010001010 Agama_agama 00000?0000 1000010111 0010000?22 012100??10 0010000110 Priscagama_gobiensis ?0100?0000 1?0000???? ?????????? ???00?0?10 0?100?1?00 Leiolepis_belliana 00000?0100 1?1001???? ????????10 00?0001?11 0010000?10 Brookesia_superciliaris ?0100?0??? 1??0?1???? ?????????? 1?200?0?11 0010000110 Saichangurvel_davidsoni ?0000?0??? ?????????? ?????????? ???00??101 ??0???1??? Sphenodon_punctatus ?2000?00?? 1??1010000 ?210001000 00000?0??0 ?010000110 Diphydontosaurus_avonis ?0000????? ??0100???? ?????????? ?????????0 0?01100110 Eublepharis_macularius 20110?0010 0000?1???? ?????????? 01?000?000 1102200001 Plestiodon_fasciatus 00110?0010 0000?010?? ?200000000 00?000?100 11000??000 Elgaria_multicarinata 10110?0000 000000???? 121????100 00?0000100 1100000101 Hypothetical_ancestor 00?00?00?0 ?00000??00 0??0000000 0000000?01 0?00000?0?

Continued 266 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Appendix 2 continued

[ 110 120 130 140 150] [..... ] Polychrus_acutirostris 0000101120 21221?1100 1111000000 0100?00100 0000102000 Pristidactylus_torquatus 0010101010 0100000101 0010010110 0001000010 1010102000 Anolis_ricordi 10?0011120 1111021100 1010000110 0100?00110 1010102000 Anolis_cristatellus 00??011120 011102?100 1010000010 0100100110 1010102000 Basiliscus_basiliscus 00??1?10?0 2200001010 001000??10 0011000010 0100100010 Corytophanes_cristatus 001?001120 2?011?1010 10?00?0110 0000000010 0000100010 Laemanctus_longipes 00?0101020 21?11?1010 01?0000110 0000000?10 0000100010 Geiseltaliellus_maarius 100?101?2? ?10?1?1010 0010000?1? ?????0???? ?????????? early_Eocene_iguanid 00??1?1??? ?????????? ?????????? ?????????? ?????????? Crotaphytus_collaris 0010101010 11001?0000 00?0110110 0000100000 0000100000 Gambelia_wislizenii 0010011010 1100001010 0010110110 0000100000 0000100000 Dipsosaurus_dorsalis 0000001020 1100010?01 0000100110 0011000000 0000000001 Brachylophus_fasciatus 0000101120 2100110100 ?0??10??10 0000000000 0000000001 Iguana_iguana 0010101120 2100110100 0010110110 0000000000 0000000001 Enyalioides_oshaughnessyi 0110101?10 ??200?1??? ?0?0????10 0000010000 0000100000 Oplurus_quadrimaculatus 000?111020 ?1?0???200 00?0101110 1011101010 0011100000 Chalarodon_madagascariensis 0001111010 0110001200 0110100010 1011000010 0011100000 Stenocercus_scapularis 0001011010 ?1101?1?01 0110111?1? 001100?110 0000001000 Microlophus_occipitalis 00?1011010 0110000001 001011??1? 0011000110 0000?02000 Tropidurus_torquatus 0???0?1010 0120000000 000?11??11 0011000110 0000002000 Plica_umbra 00?1111010 01?0000010 000011??11 00???00110 0000002000 Phymaturus_palluma 0???0?1000 0000000000 000100??10 00??100111 0000100000 Liolaemus_pictus 0???0?10?0 01?0000001 011001??10 0011100111 0000100000 Leiocephalus_personatus 0001011110 011000?001 0110101?10 0011000110 0000000000 Phrynosoma_platyrhinos 0001011010 01111?0000 0001001010 0011110000 0000010100 Petrosaurus_thalassinus 0000011000 0100000000 0000001011 0011100000 0000010100 Sceloporus_undulatus 0???1?1010 0110000001 0010101?11 0011100100 0000110100 Physignathus_cocincinus 0110011020 02121?1010 1000002010 0000000000 0000100010 Agama_agama 011?001?20 01121?1?00 1001002010 ?0??100??? ?0???????? Priscagama_gobiensis 00?01?1??? ?????????? ?????????? 0????????? ?????????? Leiolepis_belliana 0110001020 01101??010 0110002?10 0000100000 0000100110 Brookesia_superciliaris 0110000??1 01211?01?? ????00??00 0000000110 ?0?1100010 Saichangurvel_davidsoni ??????1??? ?1?0000??? ?????0???? ?????????? ?????????? Sphenodon_punctatus 011?000010 ?110020000 ?001000000 0000010010 100??????0 Diphydontosaurus_avonis 0?????0??? ?????????? ?????????? ?????????? ?????????? Eublepharis_macularius 0001010010 0111000?10 0100110000 00??110??0 ?00?0??0?? Plestiodon_fasciatus 0000010010 0110010?11 0100001010 00??1000?0 ?00?0??0?? Elgaria_multicarinata 000000?0?0 0110020?01 0110000000 00??1000?0 ?00?0??0?? Hypothetical_ancestor 00??000010 01100000?0 00000?00?0 00????0000 000??????0

Continued Eocene Lizards of the Clade Geiseltaliellus • Smith 267

Appendix 2 continued

[] [] Polychrus_acutirostris 00 Pristidactylus_torquatus 00 Anolis_ricordi ?? Anolis_cristatellus 01 Basiliscus_basiliscus 11 Corytophanes_cristatus 11 Laemanctus_longipes 11 Geiseltaliellus_maarius 11 early_Eocene_iguanid ?? Crotaphytus_collaris 11 Gambelia_wislizenii 11 Dipsosaurus_dorsalis 10 Brachylophus_fasciatus 10 Iguana_iguana ?? Enyalioides_oshaughnessyi 00 Oplurus_quadrimaculatus 00 Chalarodon_madagascariensis 00 Stenocercus_scapularis ?? Microlophus_occipitalis ?? Tropidurus_torquatus ?? Plica_umbra ?? Phymaturus_palluma ?? Liolaemus_pictus ?? Leiocephalus_personatus ?? Phrynosoma_platyrhinos 00 Petrosaurus_thalassinus 00 Sceloporus_undulatus 00 Physignathus_cocincinus ?? Agama_agama ?? Priscagama_gobiensis ?? Leiolepis_belliana 00 Brookesia_superciliaris 00 Saichangurvel_davidsoni ?? Sphenodon_punctatus 00 Diphydontosaurus_avonis ?? Eublepharis_macularius 00 Plestiodon_fasciatus 00 Elgaria_multicarinata 00 Hypothetical_ancestor 00 ; END; BEGIN ASSUMPTIONS; OPTIONS DEFTYPE = unord PolyTcount = MINSTEPS ; TYPESET * UNTITLED = unord: 2-10 12-17 19-30 32-33 35-38 40-49 51 53-57 59-66 68-71 74-78 80-87 89 91-94 96-98 100-108 110 114-146 148-152, ord: 1 11 18 31 34 39 50 52 58 67 72-73 79 88 90 95 99 109 111-113 147; END; 268 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Appendix 3: liana come from two paragraphs in Gabe and Saint Characters Used in the Phylogenetic Analysis Girons (1976) on the nasal passages. The nasal cavities For a character that is new or one not parsed according of several chamaeleons have been examined, but to previous authors, or if there are significant new obser- unfortunately not one of the earliest diverging lineage vations, a rather longer character description is neces- (Brookesia; Rieppel and Crumly 1997). Present obser- sary. I have made every attempt to cite properly the vations are therefore insufficient to bracket phyloge- original authors for comparative observations, includ- netically B. superciliaris, the representative of ing character number in previous cladistic analyses. Chamaeleonidae included here. Clearly, further study Citations with character numbers include the following: of features of the nasal capsule in this and other Arnold (1984), Conrad and Norell (2007); de Queiroz species in Iguania is desirable. (1987); Etheridge and de Queiroz (1988); Estes et al. Outgroup species were chosen to represent the (1988); Frost and Etheridge (1989); Frost et al. (2001); following major clades of Squamata: Gekkota (Euble- Gauthier et al. (1988); Hallermann (1994); Lang (1989); pharis macularius), Scincomorpha (Plestiodon fascia- Moody (1980); McGuire (1996); Poe (2004); Wiens and tus) and Anguimorpha (Elgaria multicarinata). Etheridge (2003). Certain taxa were unavailable to me. Additionally, the living rhynchocephalian Sphenodon Scoring of Liolaemus and Phymaturus follows Etheridge punctatus and one fossil species, Diphydontosaurus and de Queiroz (1988) and Frost and Etheridge (1989) avonis Whiteside, 1986, were included. Few data are for a few characters that could not directly be evaluated. available on the nasal capsule of E. macularius, except Coding for Priscagama gobiensis is from the literature what can be inferred by the phylogenetic bracketing of (Borsuk-Bial⁄ynicka and Moody 1984). gross anatomical observations on the eublepharid This study includes many characters observable on Coleonyx variegatus (Stebbins 1948) and other isolated cranial elements, especially the jaws, which are gekkotans (Born 1879; Malan 1946). Rice (1920) gave the most commonly identified elements in the fossil a careful study of the chondrocranium of Plestiodon record. Some of these were identified in recent works fasciatus. Observations on the nasal capsule in on fossil material (Smith 2006a, 2006b, 2009). Their Anguimorpha have not been taxonomically diverse inclusion in an explicit phylogenetic analysis clarifies (but see Bernstein 1999 for a recent expansion) and their phylogenetic history and so constrains the inter- most studies have focused on the common European pretation of other fossil forms. forms in Anguinae. Stebbins (1948) provides direct observations on the gross morphology of the nasal pas- All information on the nasal capsule derives from sages in E. multicarinata; without other evidence, the literature. Hallermann’s (1994) work is the most observations made by Malan (1946) on Diploglossus recent and most exhaustive comparative treatment of pleii and Ophisaurus ventralis, when the same, are used the nasal capsule in lizards (particularly in Iguania); to infer certain aspects of the morphology in Elgaria he expanded greatly on observations previously made multicarinata (this bracketing holds regardless of by Born (1879), Malan (1946), Gabe and Saint Girons which current molecular hypothesis of anguid phy- (1976), Bellairs and Kamal (1981), and others. (My logeny one accepts: Macey et al. 1999; Wiens and translation into English of Hallermann (1994) is avail- Slingluff 2001). able on request.) Among Anolis, Hallermann (1994) studied A. lineatopus and A. marmoratus, which fall in the beta anole grahami series and alpha anole 1. Premaxillary I. de Queiroz (1987) no. 43-44, Smith bimaculatus series, respectively (Poe 2004). From the (2009). Modal number of teeth: (0) eight or more, (1) phylogenetic hypothesis of Poe (2004), we may rea- seven, (2) six, (3) five or fewer. sonably suppose, in the absence of evidence to the Both Eublepharis macularius (with 13) and Elgaria contrary, that any feature shared by these two species multicarinata (with 9) show state 0, as does Diphydon- will also be present in A. cristatellus, which is included tosaurus avonis (Whiteside 1986). Plestiodon fasciatus, in this analysis (as it happens, the two species did not like many scincomorphs (Smith 2009), has seven pre- differ for any character); when Malan’s (1946) obser- maxillary teeth. Young individuals of Sphenodon punc- vations on the beta anole A. sagrei are compared, their tatus have three distinct teeth on each premaxilla agreement with those of Hallermann strengthen this (Schauinsland 1900; Werner 1962; CT scans of CM phylogenetic inference. The relationships of the 30660 [DigiMorph.org 2002–2005]; cf. Robinson 1976). anoles to one another are not being tested in this Six teeth are present in each premaxilla of the probable work; thus, no circularity—merely uncertainty—is stem-squamate Huehuecuetzpalli mixtecus (Reynoso introduced with this phylogenetic bracket. Malan’s 1998). These data suggest that a premaxillary tooth (1946) work was used to score Sceloporus undulatus count greater than seven could be plesiomorphic for and Iguana iguana and to check scoring in other Squamata, despite the smaller number (6) in S. puncta- species for potential incidences in which selected tus. State 0 is considered primitive. The character is species are not representative of the larger clades to ordered. which they belong. Born (1879) described the ethmoidal region of Liolaemus pictus and Sceloporus 2. Premaxilla II. de Queiroz (1987) no. 45. Teeth (0) undulatus. Slaby´’s (1981, 1982a, 1982b) studies sup- unicuspid, (1) with accessory cusps. plement these works. The only data on Leiolepis bel- Polarization follows de Queiroz (1987). Eocene Lizards of the Clade Geiseltaliellus • Smith 269

Figure 18. Skulls of iguanians in dorsal view. A, Basiliscus galeritus (UF 61491). B, Anolis trinitatis (TMM M8943). C, Acanthocercus atricollis (TMM M8439). Abbreviations: aiaf, anterior inferior alveolar foramen (of maxilla); ec, ectopterygoid; f, frontal; j, jugal; m, maxilla; n, nasal; p, parietal; pm, premaxilla; po, postorbital; prf, prefrontal; snaf, subnarial arterial foramen; sq, squamosal; st, supratemporal. Scale bars are 3 mm.

3. Premaxilla III. Frost (1992), Smith (2009). Nasal 4. Premaxilla IV. Smith (2009). Anterior foramina are process is (0) narrow to broad basally, tapering slowly (0) absent, (1) present. or rapidly distally, (1) moderately to very broad and Anterior premaxillary foramina transmit the medial parallel-sided for a significant portion of its length. ethmoidal nerves onto the snout (Oelrich 1956). In many Smith (2009) noted that in Basiliscus (Figure 18A), iguanian species these foramina are absent (Figure 18B, Corytophanes, Polychrus, Plica, Uranoscodon and some C).Without these foramina, presumably a branch of the Phrynosoma, among the taxa surveyed, the nasal process nerve passes just lateral to the nasal spine (Smith 2009), of the premaxilla is moderately to very broad basally, but dissections are desirable. These foramina were pres- and its lateral margins remain parallel for a consider- ent unilaterally in the one specimen each of Plica umbra able distance posteriorly; this is also true of many South and Basiliscus galeritus (Figure 18A). American anoloids. Other taxa known to have broad Smith (2009) surveyed the distribution of these and parallel-sided nasal spines include many iguanines foramina in Scleroglossa, noting their absence in (de Queiroz 1987) and hoplocercines. In contrast, the Gekkota, Scincomorpha (except for Amphisbaenia [if nasal process in most other taxa is broad basally, but that clade belongs here: Camp 1923; Schwenk 1988; tapers distally (Figure 18B); if it tapers rapidly near the Townsend et al. 2004] and Cordyliformes), most base it could be narrow along most of its length, as in Anguidae (except some derived Glyptosaurinae and most Anolis. The apparent width of the nasal process gerrhonotines), and some Varanidae. They are absent relative to the width of the bone as a whole could be as well in known rhynchocephalian outgroups, and related in some taxa (e.g., many acrodontans) to reduc- their presence is considered derived. tion of the lateral processes of the premaxilla and therefore decreased width of the rostral body of the bone (Figure 18C). Stenocercus scapularis shows the anguid 5. Premaxilla V. Etheridge (1966), de Queiroz (1987) condition (see below). no. 5, Etheridge and de Queiroz (1988) no. 1, Frost and Although Sphenodon punctatus has a wide nasal Etheridge (1989) no. 1. Nasal process of premaxilla (0) process, it is narrow in the rhynchocephalians Diphy- distally exposed between nasals, (1) substantially or dontosaurus avonis (Whiteside 1986), Clevosaurus hud- completely covered by nasals. soni (Fraser 1988) and Gephyrosaurus bridensis (Evans My specimen of Phymaturus palluma shows state 0, 1980), and in the other living outgroup species selected. which is uncommon in Liolaemini but consistent with The primitive state for Anguidae seems to be a nasal Frost and Etheridge’s (1989) observation of interspecific process that is expanded at mid-height (Smith 2006b). variability in Phymaturus. State 0 is considered primitive for Iguania on the basis Polarization follows Etheridge and de Queiroz of these outgroup comparisons. (1988). 270 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

rior end of the maxilla (Fraser 1988), is the only known exception. State 0 is therefore considered primitive.

7. Maxilla II. Smith (2006a). Dorsal surface of premaxillary process (0) is flat, (1) has deep concavity at the anterior base of the facial process. In many iguanids the dorsal surface of the premaxillary process is mostly flat (Figure 18A, B). In some taxa (several tropidurines, Polychrus), this surface evinces a strong transverse concavity at the anterior base of the facial process because of the development of a strong lateral crest; the concavity is developed between cristae transversalis and lateralis. Outgroup ambiguity requires that the transformation be treated as unpolarized.

8. Maxilla III. Evans et al. (2002). Anterior end of premaxillary process (0) overlaps only the lateral process of the premaxilla, (1) curves dorsally at its anterior end to brace the nasal spine on either side. Acrodontans are unusual among squamates in that in them the anterior end of the premaxillary process of the maxilla is strongly upturned, rising along the lateral margin of the nasal process of the premaxilla and bracing it (Figures 19D and 20B). In most other squamates, the premaxillary process simply overlies the lateral process of the premaxilla, presenting, if anything, only a very weak dorsal projection (Figure 19A–C). As state 1 occurs only in Acrodonta among examined taxa, it is considered derived. Figure 19. Skulls of iguanians in left lateral view. A, Sceloporus orcutti. The frontal of this specimen is frac- 9. Maxilla IV. Smith (2006a). Lateral rim of premaxil- tured. B, Enyalioides oshaughnessyi, anterior left max- lary process (0) does not project, or does so only dor- illa only (SMF 67590). C, Basiliscus galeritus (UF sally, (1) projects laterally, overhanging lateral margin 61491). D, Acanthocerus atricollis (TMM M8439). of bone. Abbreviations: apmf, anterior premaxillary foramen; The overhanging rim of the premaxillary process ec, ectopterygoid; ep, epipterygoid; f, frontal; j, jugal; l, is found in Polychrus and Enyalioides oshaughnessyi lacrimal; m, maxilla; n, nasal; p, parietal; pm, premax- (Figure 19B). The entire maxilla shows a lateral rim in illa; po, postorbital; prf, prefrontal; pt, pterygoid; q, Brookesia superciliaris, which also is scored 1. quadrate; re, reentrant of maxilla on jugal. Scale bar for A, C and D is 3 mm; for B, 1 mm. The absence of an overhanging rim of the premaxillary process in Sphenodon punctatus and surveyed scleroglossans indicates that state 1 is derived. 6. Maxilla I. Smith (2006a). Premaxillary process (0) overrides lateral process of premaxilla, (1) fits in slot on 10. Maxilla V. Smith (2009). On the premaxillary lateral surface of premaxilla and so is overridden by it. process (0) there is only a single foramen, (1) there are The usual condition in Lepidosauria is for the max- two foramina (for subnarial artery and anterior inferior illa to articulate on the dorsolateral surface of the lateral alveolar nerve). process of the premaxilla (Figure 19A, C, D). In Poly- In Iguanidae, uniquely in Lepidosauria, there are chrus and Enyalioides oshaughnessyi (Figure 19B), the usually two distinct foramina on the dorsal surface of premaxillary process of the maxilla inserts in a slot in the maxilla, a posterior one at the base of the facial the lateral portion of the premaxilla and so is overridden process (the anterior inferior alveolar foramen) and a by the latter (Smith 2006a). There may be a slight ten- more anterior egress for the subnarial artery (the contin- dency for this to occur in certain Phrynosoma. uation of the main trunk of the maxillary artery: Oel- All examined living outgroups, as well as many fos- rich 1956), which is commonly attended by an anterior sil rhynchocephalians (Evans 1980, 1991), show state 0, groove (Figure 18A). Smith (2009) noted that in a few which is considered primitive. Clevosaurus hudsoni, in iguanids, including Anolis (Figure 18B), there is but a which the lateral processes of the premaxilla are greatly single foramen. It is possible that in these species, and in expanded and articulate on the dorsal surface of the ante- other lepidosaurs, the anterior inferior alveolar nerve Eocene Lizards of the Clade Geiseltaliellus • Smith 271 exits the body of the maxilla through this same foramen alongside the subnarial artery, in which case these structures could have reacquired a common aperture. Fur- ther anatomical study is needed. Apart from Anolis, there are very few iguanids that do not show this morphology. In many iguanines, however, it is obscured by confluence of the two foramina, forming an (sometimes only unilaterally) an elongate opening. In Sphenodon punctatus, there is but a single foramen on the dorsal surface of the premaxillary process of the maxilla, just medial to the narial rim (Figure 20A). This foramen seems to transmit the maxillary branch of the temporal artery from the body of the maxilla toward the premaxilla (O’Donoghue 1921) and perhaps also the anterior branch of the superior alveolar nerve. This single foramen is present in all scleroglossans and is generally located just anterior to the base of the facial process of the maxilla, although in certain anguimorphs (Xenosauridae, Varanoidea and extinct outgroups to the latter) it is shifted to lie posteromedial to the crista transversalis (Smith 2006b). Also in Acrodonta there is but a single foramen at the base of the facial process (Figure 18C). The anterior end of the premaxilla, which is expanded dorsally along the base of the nasal process of the premaxilla (see above), is often deeply grooved, or even pierced by a foramen that transmits a vessel or structure passing anteriorly along the dorsal surface of the maxilla, because there is no separate opening from the maxillary body. Late Cretaceous iguanians from Mongolia likewise show state 0 (see figures in Gao and Norell 2000; Conrad and Norell 2007). Marmoretta oxoniensis seems similar to Sphen- odon punctatus in having a single anterior foramen on the medial surface of the premaxillary process of the maxilla (see Evans 1991). The apparently single foramen in Gephyrosaurus bridensis (see Evans 1980) is possibly located more dorsally and laterally. If identified correctly, the same would apply to Diphydon- tosaurus avonis (see Whiteside 1986, fig. 6a), but the canal exposed by breakage on the medial surface of the maxilla in this species (Whiteside 1986, fig. 6b) suggests to me that the canal opened medially in this species and Figure 20. Maxillae of lepidosaurs. A, Sphenodon that the foramen was misidentified. The distribution of punctatus (YPM 5436), left maxilla in medial view. B, these foramina in Lepidosauria suggests that state 0 is Agama agama (TMM M8448), left maxilla in dorsal primitive. view. C, Basiliscus vittatus (YPM R11132), left maxilla in dorsal view. D, Anolis cristatellus (YPM R12048), 11. Maxilla VI. Etheridge (1959), Conrad and Norell right maxilla in dorsal view. Abbreviations: aiaf, ante- (2007) no. 1. Snout-to-skull length ratio (0) <0.33, rior inferior alveolar foramen (of maxilla); crtv, crista (1) 0.33. transversalis; ccr, canthal crest; jb, jugal buttress; nf, nasal facet; plpr, palatine process of maxilla; saf, supe- The length of the snout (measured from tip of pre- rior alveolar foramen; snaf, subnarial arterial foramen. maxilla to anterior border of orbit) relative to the length Scale bars are 1 mm. of the skull (measured from tip of premaxilla to occipital condyle) is >0.4 in Polychrus and Anolis, greater than in other ingroup and outgroup taxa, where the ratio is less than 1/3 in nearly every case. However, corytopha- 12. Maxilla VII. Smith (2006a, 2006b): Facial process nines generally also have a relatively elongated snout (0) vertical or weakly to moderately arching medially, (ratio is 0.33 but <0.4); Basiliscus galeritus was excep- (1) folded medially around an oblique, anteroventrally tional in showing state 0. trending axis, creating a strong canthal crest and Outgroup comparisons suggest 0 is the primitive anterodorsally facing surface continuous with the pre- state. maxillary process of the maxilla. 272 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Figure 21. Palates of iguanians in ventral view. A, Basiliscus galeritus (UF 61491). B, Uma scoparia (CAS 42135). C, Acanthocercus atricollis (TMM M8944); the dotted line indicates the posterior extent of the palatine process of the maxilla in this species (it is underlapped by the palatine in this and several other agamids). D, Brookesia superciliaris (TMM M8676). Abbreviations: m, maxilla; pl, palatine; plpr, palatine process of maxilla; pm, premaxilla; pt, pterygoid; v, vomer. Scale bars are 3 mm.

As defined, state 1 is present in most phrynoso- The character is treated as unpolarized, because matines, the Tropidurus group and Anolis (Figure 20D). outgroup comparison is impossible. The angle of the canthal crest is much lower in Anolis than in the other taxa, probably because of snout elon- 14. Maxilla IX. Cope (1864:225), Moody (1980) no. 40, gation in the former. There is no clear basis for discrim- Estes et al. (1988), Frost and Etheridge (1989) no. 2. ination of the states found in the different groups. Maxillae (0) not in contact medially on palate, (1) meet However, it is notable that seemingly intermediate states on mid-line behind palatal flange of premaxilla of two kinds are found in Iguanidae (Smith 2006a). In Moody (1980), Rieppel (1987) and Rieppel and crotaphytines, oplurines and many tropidurines Crumly (1997) observed variation in Acrodonta. In the (including Leiocephalus and liolaemins), the facial specimens included here, the maxillae are well-exposed process is weakly folded medially, creating a narrow on the palate in ventral view and touch on the mid-line. anterodorsally facing surface that is mediolaterally wider Polarization follows Frost and Etheridge (1989). than in, say, Basiliscus (Figure 20C). In Corytophaninae and Polychrus, in contrast, the facial process is strongly 15. Maxilla X. Smith (2006b). Palatine process is (0) bent medially, but only towards its distal end; this dis- symmetrical, or nearly so, (1) strongly asymmetrical, tinct kind of bending creates a small, dorsally facing sur- decaying abruptly behind the palatine articulation into face. These different intermediates suggest that state 1 a narrow posterior shelf. may have been achieved independently in different groups of Iguanidae. Acrodontans are characterized by a broader (and dorsomedially directed) palatal flange of the maxilla The facial process of the maxilla is not folded in anterior to the palatine articulation, whereas in iguanids Sphenodon punctatus, Gephyrosaurus bridensis (Evans and other squamates generally the palatal flange is no 1980, fig. 31), Clevosaurus hudsoni (Fraser 1988, fig. 22), wider anteriorly than posteriorly. Yet in most agamids Diphydontosaurus avonis (Whiteside 1986, fig. 6f) or the palatine process is still symmetrical in that the lead- scleroglossan outgroups. State 0 is therefore considered ing and trailing edges of the process maintain about the primitive. same angle to the axis of the jaw (Figure 21C; here the palatine underlaps the palatine process somewhat). In 13. Maxilla VIII. Novum. If a strong medial fold is pres- contrast, in Leiolepis, Uromastyx and Chamaeleonidae ent in the facial process of the maxilla, the angle it forms (Figure 21D), the palatal shelf decays strongly and with the horizontal plane is (0) <35° or (1) >35°. abruptly in width just posterior to the palatine process, The angle is perhaps related to snout elongation. at an angle of approximately 90° to the long axis of the Species lacking a strong medial fold are necessarily jaw (Smith 2006b). This condition also obtains in rare scored as unknown. An early Eocene species from the iguanids (e.g., some Phrynosoma). Bighorn Basin of Wyoming (Smith 2006b) has an angle A palatal flange is effectively absent in Sphenodon of 29°. punctatus. In Scleroglossa it can be broad, but the Eocene Lizards of the Clade Geiseltaliellus • Smith 273 palatine process is not usually asymmetrically devel- taxa with an enlarged superior alveolar foramen (e.g., oped. State 0 is judged to be primitive. Hoplocercinae and Tropidurinae*), it is usually closer to state 0 than to state 1. 16. Maxilla XI. Smith (2006a). In adult, palatine process Smith (2006b) disputed his earlier conclusion is (0) weak to moderate in size, rounded, (1) large and (Smith 2006a) that a gutter on the dorsal surface of the subtriangular. palatal shelf is the primitive condition. In particular, he The palatine process of the maxilla, on which the found that a well-developed gutter is not present in Scle- palatine articulates, is in many iguanids (Corytophani- roglossa. A well-developed gutter for the superior alve- nae, Polychrotinae, Iguaninae and Hoplocercinae) lit- olar nerve is present neither in Sphenodon punctatus tle more than a weak, medial swelling of the palatal (Figure 20A) nor in fossil rhynchocephalians (Diphy- flange (Figure 21A). In others (Phrynosomatinae, dontosaurus avonis: Whiteside 1986, fig. 7b; Gephy- Tropidurinae, Oplurinae and Crotaphytinae) it is a rosaurus bridensis: Evans 1980, fig. 32B). The presence strong projection, subtriangular to triangular, which of a gutter is therefore considered derived. provides an extensive surface for the palatine articulation (Figure 21B). A strong process is commonly, but 19. Jugal I. Novum. (0) Lateral wall of maxilla covers not universally, present in acrodontans (Figure 21C, D; jugal (and lacrimal, when present) extensively below see Moody 1980). orbit, forming an at least weakly convex upward suture; In rhynchocephalian outgroups, a palatal flange is (1) jugal (and lacrimal, when present) exposed broadly effectively absent on the maxilla and the palatine artic- at orbital rim, generally forming a straight or concave ulates more or less directly with the maxillary body. The upward suture with the maxilla. palatine process is strong in Eublepharis macularius and In some iguanians, the jugal attains broad posterior Plestiodon fasciatus, but weak in Elgaria multicarinata exposure dorsal to the maxillary wall that overlaps it and many other anguimorphs, so state 0 is considered (Figure 19A). The suture line can parallel the orbital primitive. margin, exposing a more or less invariant amount of the jugal; in some taxa, the jugal exposure is progressively 17. Maxilla XII. Smith (2006b). (0) Dorsal surface of reduced anteriorly, but the suture line is concave- palatal shelf flat posteriorly or with only a small and upward, and the jugal broadly exposed posteriorly (e.g., weak longitudinal swelling anteriorly, medial to the many phrynosomatines). In many other taxa, however, jugal articulation; (1) medial to the jugal groove, palatal the suture line is concave-downward; the lateral expo- surface projects dorsally as a strong ridge, buttressing sure of the jugal is strongly and rapidly restricted ante- the articulation. riorly. This latter state is found in hoplocercines, In Corytophaninae, the anterior end of the jugal is corytophanines (Figure 19C), Geiseltaliellus (see Figures buttressed medially by a thin, tall, longitudinal flange of 2 and 3), and many polychrotines; it is also found in bone that projects dorsally from the palatal surface of Oplurus quadrimaculatus (but not other examined the maxilla and is commonly accompanied by an espe- oplurines) and Phrynosoma platyrhinos. In agamids cially deep and narrow jugal groove (Figure 20C). A but- exclusive of Leiolepis the jugal-maxillary suture takes on tress is moderately developed in Crotaphytus collaris as a peculiar shape becausse of the maxillary reëntrant well, which also shows a deep and narrow jugal groove, (Smith 2006b; Figure 19D: see below); these taxa are also and it is variably present in an early Eocene species from scored 0, although their condition might not be homol- Wyoming (Smith 2006b). ogous with that in iguanids identically scored. Leiolepis The absence of a jugal buttress in most iguanians and Priscagama gobiensis show state 1. and in all examined outgroups indicates that state 1 is Although some scleroglossans show state 1, all derived. included outgroups except Plestiodon fasciatus show state 0, as does Diphydontosaurus avonis (Whiteside 1986, fig. 18. Maxilla XIII. Smith (2006a, 2006b). Dorsal surface 9k). State 0 is consequently treated as primitive. of palatal shelf (0) flat, or nearly so, with a relatively small superior alveolar foramen, (1) with an elongate 20. Jugal II. Smith (2006b). (0) The suture line between depression (gutter), at whose floor the superior alveolar the maxilla and jugal is a line or a simple curve in lateral nerve and maxillary artery penetrate the body of the view, (1) maxilla shows a posterior reëntrant on the maxilla. jugal. Smith (2006a) described the presence of a well- In most squamates the suture line between the developed gutter on the dorsal surface of the palatal maxilla and jugal forms a straight line or a simple curve flange of the maxilla in several Holocene and fossil (i.e., only one inflection point) in lateral view (Figure iguanids; the superior alveolar nerve or maxillary artery 19A, C). In many acrodontans, the maxilla has a distinct penetrate the maxilla in multiple places at the base of reëntrant on the jugal; from its posterior end, the this gutter. In some iguanids, however, the superior maxillary–jugal suture line extends first anterodorsally, alveolar foramen is restricted to a relatively small open- then abruptly curves posterodorsally, then extends ing near the transverse level of the posterior end of the at last anteriorly (Figure 19D). This reëntrant is found facial process, and the dorsal surface of the palatal flange in agamines and in Uromastyx, but not Leiolepis or is otherwise flat. Quantitative variation occurs, but in Chamaeleonidae. 274 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

The rhynchocephalians Sphenodon punctatus in Autarchoglossa (see above) and Rhynchocephalia (Smith pers. obs.) and Gephyrosaurus bridensis Evans, (Gauthier et al. 1988). Thus, state 0 is probably primitive. 1980, the stem-squamate Huehuecuetzpalli mixtecus Reynoso, 1998, most scleroglossans, chamaeleonids and 23. Jugal V. Novum. Distal end of temporal ramus of all iguanids show state 0, which is deemed ancestral. jugal (0) posteriorly deflected, (1) colinear with main State 1 seems to have been acquired independently in trend of temporal ramus. Scincoidea (Smith 2006b). It is unclear in Diphydon- In many iguanids (Phrynosomatinae, Tropidurinae, tosaurus avonis (see Whiteside 1986). Oplurinae and Crotaphytinae), as well as most acrodontans, the tip of the temporal ramus of the jugal is poste- 21. Jugal III. Novum. Temporal ramus (0) slender or riorly deflected. The deflection is particularly marked on subequal in size to suborbital ramus, (1) posteriorly the posteroventral margin of the tip and gives the tempo- expanded. ral ramus a sinusoidal appearance (Figure 19A, D). In In most iguanian taxa the temporal (ascending) Polychrotinae*, Corytophaninae (Figure 19C), Iguani- ramus of the jugal is subequal in size to the suborbital nae and Hoplocercinae, however, the tip follows the ramus, but in a few it is posteriorly expanded. The same trend as the more proximal portion of the tempo- expanded condition is confined to Phrynosoma ral ramus; there can be a slight deflection of the dorsal platyrhinos and Corytophanes cristatus among ingroup margin where it meets the postorbital, but the posterior taxa. margin of the temporal ramus is nearly straight, espe- Outgroup comparison suggests state 1 is derived. cially in Corytophaninae and Polychrotinae*. The posterior margin of the temporal ramus is 22. Jugal IV. Gauthier et al. (1988) no. 33, McGuire nearly straight in Sphenodon punctatus, but the Jurassic (1996), Smith (2006b). Angle of jugal (0) is drawn out and Triassic rhynchocephalians Pleurosaurus (Cocude- into a sharp corner, with a moderate to strong posterior Michel 1963) and Diphydontosaurus avonis (Whiteside (quadratojugal) process, (1) is rounded, generally 1986, pl. 2i), respectively, show a deflected tip, as does broadly so. the stem-squamate Huehuequetzpalli mixtecus (Reynoso Gauthier et al. (1988:49) argued that a strong 1998, fig. 4). The tip of the jugal is broken in illustrated quadratojugal process of the jugal (Figure 19D) is prim- specimens of Marmoretta oxoniensis (Evans 1991) and itive for Lepidosauromorpha, the clade including many Gephyrosaurus bridensis (Evans 1980). In most scle- fossil outgroups to Lepidosauria. They also stated that roglossans, the temporal ramus of the jugal is straight, this process is not present in Squamata, by which they but the relation of this bone to the postorbital is consid- presumably meant that it is reduced in comparison with erably altered in that clade (J. A. Gauthier, pers. comm. Rhynchocephalia; Smith (2006b) observed a quadrato- 2006). Many (stem) taxa from the Late Cretaceous of jugal process (state 0) in nearly all autarchoglossans. Mongolia, including Saichangurvel davidsoni (Conrad However, Eublepharis macularius, like other gekkotans, and Norell 2007), priscagamids (Borsuk-Bialynicka and cannot be scored because the temporal ramus of the Moody 1984; Gao and Hou 1995; Alifanov 1996) and jugal—needed to define a homologous position for the Anchaurosaurus gilmorei (Gao and Hou 1995), show base of a quadratojugal process—is so highly reduced the deflected condition. Available data thus suggest that (Estes et al. 1988). In most iguanids the quadratojugal state 0 could have been primitive for Squamata and process is lacking, and the ventral margin of the jugal Iguania. near the junction of the suborbital and temporal rami forms a smooth curve (Figure 19A, C), more rarely a 24. Ectopterygoid I. Novum. Ventral corner of postero- corner approaching 90°. A quadratojugal process is also lateral process of ectopterygoid (0) achieves weak lateral lacking in many iguanians from the Late Cretaceous of exposure or none at all below angle of jugal, (1) is Mongolia (Gao and Norell 2000; Conrad and Norell exposed laterally as a wedge inserting between the angle 2007). of the jugal and the posterior tip of the maxilla. The development of a quadratojugal process is pre- In iguanians, as in lepidosaurs primitively (Smith sumably related to the quadratojugal ligament. Beddard 2009), the posterolateral process of the ectopterygoid is (1905a) dissected several squamates and found a dis- well developed, contributing to the coronoid recess. The tinct ligament absent in Iguana iguana, which lacks a process consists of two corners, a dorsal one extending posterior process, and present in Physignathus, which along in the medial surface of the jugal and a ventral one has one (ligament confirmed in FMNH 51708). How- extending toward the angle of the jugal. In many ever, I find the ligament in phrynosomatines (Scelo- iguanids (Crotaphytinae, Oplurinae, Tropidurinae, porus, Uta and Urosaurus), which lack the process, Phrynosomatinae and Pristidactylus torquatus), the ven- implying that loss of the ligament is not necessary for tral process is particularly strong and is exposed laterally loss of the process. Possibly the origin of the ligament on the skull between the posterior end of the maxilla differs. and the angle of the jugal. In Iguaninae, however, the Some iguanians, mainly acrodontans, but also Cro- lateral exposure of the ventral corner is a laterally flat- taphytinae, Leiosaurini and many Anolis, show state 0; tened, wedge-shaped structure inserting between the McGuire (1996:18) suggested this feature as an autapo- angle of the jugal and the posterior tip of the maxilla. In morphy of Crotaphytinae. State 0 is furthermore present Acrodonta, Corytophaninae, Anolis, Polychrus and Eocene Lizards of the Clade Geiseltaliellus • Smith 275

Hoplocercinae the posterolateral process achieves no lateral exposure, possibly because of posterior extension of the jugal. The ectopterygoid is not exposed laterally in Sphen- odon punctatus or most scleroglossans (except Xenosauridae), probably in large part because of the loss of the posterolateral process of that bone (Smith 2009).

25. Lacrimal I. de Queiroz (1987) no. 17, Etheridge and de Queiroz (1988) no. 4, Frost and Etheridge (1989) no. 5. Lacrimal is (0) present or (1) absent as a distinct ele- Figure 22. Right prefrontal of Basiliscus vittatus (YPM ment. R11132) in lateral view. Note the teardrop-shaped pre- Polarization follows Etheridge and de Queiroz orbital tuberosity and the semicircular lacrimal facet. (1988). Abbreviation: lf, lacrimal facet. Scale bar is 1 mm.

26. Lacrimal II. Moody (1980) no. 35, Etheridge and de Queiroz (1988) no. 5, Frost and Etheridge (1989) no. 6, Hallermann (1994) no. 27. Lacrimal foramen (0) not infraorbital margin by apposition of jugal and pre- much larger than palatine foramen, (1) much larger frontal. than palatine foramen State 1 is confined to Polychrus among living taxa Polarization follows Etheridge and de Queiroz and is therefore taken to be derived. (1988). 29. Prefrontal I. Smith (2009). Orbital corner of pre- 27. Lacrimal III. Lang (1989) no. 7. Inferior orbital rim frontal (0) smooth, or with teardrop-shaped boss, or (1) (0) continuous across lacrimal–prefrontal junction, not with strong, straight, projecting canthal crest continuing impressed, or (1) impressed briefly and abruptly by onto maxilla. indentation at lacrimal–prefrontal junction, obliterat- As discussed in Smith (2009), a strong, straight can- ing the rim. thal ridge on the prefrontal is primarily confined to cer- In most iguanians the orbital rim is more or tain polychrotines, especially Anolis and Polychrus. All less continuous from jugal (to lacrimal) to prefrontal species with a supraorbital spine on the prefrontal have (Figure 19A, D). In some taxa, the lateral surfaces of the a similar ridge (e.g., Gonocephalus, Phrynosoma), but lacrimal and prefrontal are not smoothly continuous, these species are only given state 1 if the ridge is introducing a step, but this condition is not judged to be expressed on the main body of the prefrontal. homologous with the distinct lacrimal–prefrontal groove The presence of state 0 in rhynchocephalian and in Corytophaninae (Figure 19C) and in Geiseltaliellus scleroglossans outgroups, as well as most iguanians, maarius (Figure 4). Most important in distinguishing implies that that state is primitive. between a groove as seen in corytophanines and similar structures in other species (e.g., Oplurus cuvieri) is fol- 30. Prefrontal II. Smith (2009). (0) Lacrimal is but- lowing the inferior orbital margin. In corytophanines, tressed only dorsally, posteriorly, or both, by a weak lap- the orbital margin is distinctly impressed ventral to the pet on the prefrontal; (1) a small, semicircular facet is prefrontal boss and the orbital rim is completely obliter- completed by the addition of a ventral projection ated. In other species the orbital margin is uninterrupted around the lacrimal articulation surface. (e.g., Gambelia wislizenii). This indicates that in other In most iguanians, the lacrimal is only weakly species, the development of a prefrontal boss is primary braced by the prefrontal, from which a small lappet may in the appearance of a step, whereas in corytophanines project dorsal or posterior to the lacrimal articulation. A the groove would exist regardless of whether the boss did small, semicircular articulation facet is developed in (cf. SMF ME 1769). Dissection suggests that the groove Corytophaninae (Figure 22). The state of this character functions to create a more protrusive prefrontal boss, to is unknown in Geiseltaliellus maarius, in which the pre- which connective tissue attaches. frontal and lacrimal are still in articulation, but a semi- In some species examined, a groove simply exag- circular facet is seen in an early Eocene taxon thought to gerate the prefrontal boss to which skin and connective be primitive for both Corytophaninae and Geiseltaliel- tissue are attached, much like the postorbital boss lus (Smith 2009). Leiocephalus also shows state 1. The described by Oelrich (1956) in Ctenosaura pectinata. I facet clearly cannot be scored in species lacking a do not find state 1 to be present in Gambelia wislizenii. lacrimal. I do find it, however, in some Leiocephalus. Sphenodon punctatus and most other sphenodon- Polarization follows Lang (1989). tidans lack a lacrimal (e.g., Whiteside 1986; Gauthier et al. 1988). Gephyrosaurus bridensis, however, has one; 28. Lacrimal IV. Frost et al. (2001) no. 51. Lacrimal (0) it fits into a vertically elongate facet on the lateral mar- forms part of infraorbital margin, (1) excluded from gin of the prefrontal (Evans 1980) and so displays a 276 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

A posterolateral process of the palatine character- izes most Acrodonta (except chamaeleons and Uro- mastyx) and many iguanids (Figure 23A), but is reduced in many other ingroup taxa. The posterolateral palatine process is present but very weak in certain taxa (e.g., Pristidactylus torquatus and Stenocercus guentheri: Torres-Carvajal 2003) and is coded as absent in these. The process is present in most Polychrus, but happens to be absent in P. acutirostris. In most phrynosomatines and tropidurines, the process is absent (Figure 23B). It is also absent in most Dipsosaurus dorsalis, but many other iguanines show state 0 (de Queiroz 1987). The process is considered absent in Chalarodon madagascariensis, but present in Oplurus cuvieri. Conrad and Norell (2007) describe the infraorbital foramen in Saichangurvel davidsoni as partly bounded by the maxilla but illustrate it (2007, fig. 6) as completely enclosed within the palatine; for the present, I code it 0. Temu- jinia ellisoni also shows state 0 (Gao and Norell 2000). State 0 obtains in Anguidae, Xenosauridae (sensu Estes et al. 1988), Varanidae, Helodermatidae, nearly all Scincidae and Cordyliformes, Lacertidae and the xan- tusiid Klauberina riversiana among Scleroglossa, including the species-level outgroups used here. State 1 is common in Gekkota (especially Eublepharidae) and Xantusia, whereas Teiidae and Gymnophthalmidae show state 2. State 0 is also seen in Sphenodon punctatus, Gephyrosaurus bridensis (Evans 1980) and Diphydon- tosaurus avonis (Whiteside 1986). On the basis of these outgroup comparisons, state 0 is considered primitive. As the character describes degree of enclosure of the infraorbital foramen by the palatine, it is ordered.

32. Postfrontal I. Estes et al. (1988) no. 12, Etheridge and de Queiroz (1988) no. 6, Frost and Etheridge (1989) no. 9, Lang (1989) no. 12, Poe (2004) no. 62. Postfrontal (0) separate, (1) fused to postorbital or (2) fused to Figure 23. Left antorbital region of iguanians in posfrontal. terodorsal view. A, Basiliscus galeritus (UF 61491). B, In living iguanians in which the postfrontal is dis- Sceloporus orcutti. Abbreviations: f, frontal; j, jugal; l, tinct, it is always confined to the orbital margin (Estes lacrimal; m, maxilla; mpf, maxillopalatine foramen; p, et al. 1988; Figure 24A, C). In no iguanian with a dis- parietal; pl, palatine; pl-lpr, lateral process of palatine; tinct postfrontal does the postorbital extend along the prf, prefrontal. Scale bar is 1 mm. orbital margin of the frontal. These observations are useful in considering the fate of the element in species in which the postfrontal is not distinct. If a distinct postfrontal is absent but the postorbital seems to extend a long anterior process along the orbital margin of the morphology close to state 0, as described above. frontal (e.g., Leiosaurus belli), I regard the postfrontal to Plestiodon fasciatus has a lacrimal facet likening the be fused to the postorbital, a view that is strengthened in corytophanine condition, but it is absent in Elgaria mul- some cases by the presence in close relatives (for ticarinata. State 0 is considered primitive. Leiosaurus, for instance, in Aperopristis paronae and A. catamarcensis, following the taxonomy of Frost et al. 31. Palatine I. de Queiroz (1987) no. 23. (0) Infraorbital 2001) of a distinct postfrontal in precisely the position of foramen wholly enclosed in palatine, or nearly so, by the supposed anterior process of the postorbital. On the upgrowth of posterolateral process of palatine; (1) pos- other hand, in taxa in which there is no distinct post- terolateral process of palatine with strong posterolateral frontal and the postorbital articulates solely with the process, enclosing the infraorbital foramen ventrally, frontoparietal corner and does not seem to extend ante- posteriorly, or both ; (2) posterolateral process weak or riorly along the orbital margin of the frontal (Crota- absent, infraorbital foramen bordered only by maxilla phytinae and Agamidae), I regard the postfrontal as posteriorly. absent; in such cases, this character is scored?. Eocene Lizards of the Clade Geiseltaliellus • Smith 277

Figure 24. Postorbital complexes in iguanians. A, Right postorbital complex of Enyalioides oshaughnessyi (SMF 67590) in posterodorsal view. B, Right postorbital complex of Basiliscus galeritus (UF 69491) in anterior view. C, Right postorbital complex of Sceloporus occidentalis (CAS 178414) in posterodorsal view. D, Left postorbital complex of Phrynosoma platyrhinos (TMM M8952) in lateral view. Abbreviations: f, frontal; p, parietal; po, postorbital; pof, postfrontal. Scale bars are 1 mm.

The postfrontal is reinterpreted here in several ing entirely in most “sand lizards.” Weiner and Smith iguanian taxa. Geiseltaliellus maarius has a small post- (1965) regarded the postfrontal as fused to the postor- frontal (contra Rossmann 2000) that has lost most con- bital in crotaphytines; I disagree for reasons articulated tact with the frontal. There is also in Basiliscus galeritus above. Beddard (1905b) reported a tiny postfrontal in a tiny and separate ossification along the orbital surface Uromastyx spinipes; I have found such a postfrontal in at the junction between the postorbital and frontal (Fig- several Uromastyx, among them U. acanthinurus. There ure 24B), which I interpret as a small postfrontal; this has been debate about the status of the postfrontal in structure could not be confirmed in other Basiliscus. chamaeleons: some have treated it as absent (Frank Phrynosoma and the “sand lizards” Uma, Callisaurus, 1951; Frost and Etheridge 1989), whereas others (Camp Holbrookia and Cophosaurus (Smith 1946), thought col- 1923; Rieppel 1987, 1993; Rieppel and Crumly 1997) lectively to form the sister clade to Phrynosoma regard the postorbital element to be ambiguous. In (Etheridge and de Queiroz 1988; Frost and Etheridge Brookesia superciliaris, the postorbital extends a consid- 1989), generally lack a distinct postfrontal (e.g., erable distance anteriorly along the lateral margin of the Etheridge 1964; Presch 1969). A juvenile (SVL “ 38 mm) frontal, suggesting that this portion is the postfrontal P. platyrhinos shows an element on the ventrolateral that has fused to the postorbital. For the present, I code margin of the posterior supraorbital spine that is dis- the species as unknown. tinct laterally (Figure 24D), but apparently fused medi- The element that clasps the frontoparietal suture in ally. This element may represent as an incompletely Eublepharis macularius is regarded here, on the basis of fused postfrontal, though evaluation in cleared-and- topographical relations and size, to represent a post- stained juveniles of this and other Phrynosoma is desir- frontal (Camp 1923), not a fusion of the postorbital and able. Among “sand lizards,” one Uma scoparia (CAS postfrontal (cf. Estes et al. 1988). A distinct postfrontal 42135) showed a tiny postfrontal on the right side only, is primitive for Lepidosauromorpha (Gauthier et al. inconspicuous at the junction of the frontal and postor- 1988) and is found in most scleroglossans (except some bital along their orbital face, much as in B. galeritus; it is anguimorphs; Estes et al. 1988). State 0 is therefore con- possible that the postfrontal is extremely small or lack- sidered primitive. 278 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

33. Postfrontal II. Novum (but cf. Estes et al. (1988) no. The postorbital makes a stronger contribution to 13). Posterior splint of postfrontal (0) present, extend- the orbital margin than the postfrontal in Sphenodon ing onto posterior side of anterolateral corner of parietal punctatus (Smith pers. obs.), but in Diphydontosaurus (and forming a clasp if the anterior splint is present), or avonis the opposite is true (Whiteside 1986). It is not (1) absent, leaving postfrontal (if present at all) confined clear, on the basis of available elements, whether this is to orbital margin. true in Clevosaurus hudsoni (Fraser 1988) or Jurassic Estes et al. (1988) recognized that a postfrontal that rhynchocephalians (Cocude-Michel 1963). It is possi- clasps the frontoparietal suture is characteristic of Scle- ble that state 0 is primitive for Squamata. roglossa, whereas the postfrontal in iguanians, when present, is confined to the orbital margin. Recent dis- 36. Postorbital I. Etheridge and de Queiroz (1988), coveries in the Late Cretaceous of Mongolia have shown Lang (1989). Posterodorsal margin (0) concave to that a clasping morphology is found in some (stem-) straight, posterior ramus of postorbital tapers smoothly iguanians as well (Gao and Norell 2000; Conrad and posteriorly, or (1) convex, expanded dorsomedially over Norell 2007) and so could be primitive for Squamata, a supratemporal fossa. conclusion that also finds support in the similar mor- State 1 is found in corytophanines (Lang 1989; phology of the postfrontal in Rhynchocephalia. The Figure 19C) and many Anolis (Smith 2006b), as well as clasping postfrontal of Estes et al. (1988), who described Chalarodon madagascariensis, some Oplurus, Leio- the postfrontal in Scleroglossa as having a “semilunate” cephalus and some agamids. The transformed nature of form (which Sphenodon does not have), should not be the chamaeleon skull makes this character difficult to taken as identical to state 0 here. I treat the anterior and evaluate there. It is variably present in Geiseltaliellus posterior rami of the postfrontal like independent fea- maarius and G. longicaudus. In other taxa, the posterior tures. Iguanians are characterized by the loss of the pos- ramus of the postorbital tapers smoothly posteriorly terior ramus (this character); some iguanians are further (Figure 19A, D). characterized by the diminution, loss or fusion of the The postorbital in Sphenodon punctatus is dorsally anterior ramus (characters 30 and 32). It happens that convex because the whole bone is arched, not because it no iguanian has only a posterior ramus. is dorsomedially expanded per se; this species is thus 34. Postfrontal III. Novum (but cf. Estes et al. (1988) scored 0. A tapering posterior ramus of the postorbital no. 12). If postfrontal is present, its anterior splint is (0) is also seen in Diphydontosaurus avonis (Whiteside relatively large, visible in dorsal view, or (1) small, 1986), Gephyrosaurus bridensis (Evans 1980) and Mar- obscured from dorsal view by the frontal and postor- moretta oxoniensis (Evans 1991). There is the appear- bital; (2) anterior splint of postfrontal absent. ance of dorsomedial expansion in Clevosaurus hudsoni The postfrontal in iguanians is a relatively small (Fraser 1988, fig. 9), but it seems to be caused by ventral bone that contacts only the frontal and postorbital. Yet, expansion instead, for the squamosal and jugal articula- if the iguanian postfrontal represents the anterior splint tions are massively broadened in this species. The pres- of the primitive lepidosaurian postfrontal (see above), ence of state 0 in (most) rhynchocephalians and most this splint cannot be considered especially small itself scleroglossans that have a well-developed postorbital (Figure 24A, C). Basiliscus has been considered to lack implies that this is ancestral. a postfrontal (Etheridge and de Queiroz 1988; Lang 1989). As noted above, however, a tiny postfrontal (state 37. Postorbital II. McGuire (1996) no. 6, Smith (2009). 1) is present in B. galeritus (Figure 24B) and I have seen Postorbital (0) broadly underlaps frontoparietal corner, one as well in B. plumifrons. Geiseltaliellus maarius or (1) does not underlap corner, articulates on lateral shows a morphology not too unlike that in B. galeritus corner of frontal, parietal, or both. and is coded the same. In Corytophaninae, Polychrotinae* and Crota- State 0 is taken to be ancestral for Squamata on the phytinae, among iguanids, the postorbital broadly basis of its presence in Sphenodon punctatus and other underlaps the frontoparietal corner (Smith 2009; Figure rhynchocephalians and in Scleroglossa. The character 24B) and also may articulate to a greater or lesser extent is ordered. on the posterior surface of the anterolateral process of the parietal. Underlap means that a mediolaterally broad 35. Postfrontal IV. cf. Estes et al. (1988) no. 17. Contri- orbital face of the postorbital is usually developed, and bution of postfrontal to posterior orbital margin (0) could have a sharp lateral boundary. In most other strong, greater than that of postorbital, or (1) weak. iguanids, significant underlap of the frontoparietal cor- A strong contribution of the postfrontal to the ner is absent, and the orbital face of the postorbital is orbital margin was regarded as a autapomorphy of Scle- minimal. In most Iguaninae, Hoplocercinae and a few roglossa among squamates (Estes et al. 1988). However, tropidurines, the postfrontal is comparatively large and the lateral portion of the postfrontal in at least some articulation of the postfrontal is with the posterior face newly described (stem-)iguanians from the Late Creta- of the parietal corner and the posterior surface of ceous of Mongolia (Gao and Norell 2000; Conrad and the postfrontal (Figure 24A). In Oplurinae, most Norell 2007) is well developed and restricts the contri- tropidurines, and Phrynosomatinae the postorbital bution of the postorbital to the posterior orbital margin, articulates only laterally on the parietal (Figure 24C) raising the prospect that this is primitive for Squamata. because of the comparatively small postfrontal. On the Eocene Lizards of the Clade Geiseltaliellus • Smith 279 basis of articulation surfaces, state 1 seems to obtain in tive, heritable positional development (position, as Phymaturus palluma, although an articulated specimen opposed to size, is coded separately below). State 1 is was not available. State 0 obtains in one examined commonplace in Corytophaninae, Crotaphytinae and tropidurine, Stenocercus scapularis. In agamids, the pos- Polychrotinae* (most clearly in Leiosaurini). When torbital strongly underlaps the frontoparietal corner and most strongly developed (state 2), the tubercle forms a also frequently invades the mutual suture of these bones, spine that projects anteriorly over the orbit and may becoming visible between them in dorsal view; this mor- meet midway a counterpart developed on the prefrontal phology is viewed as an expression of state 0. to form a complete supraorbital bar. Phrynosoma (Fig- This iguanian scheme is difficult to apply to other ure 24D), Corytophanes and some Gonocephalus have lepidosaurs, in which the postfrontal is completely inter- the best-developed supraorbital spines. It seems that the posed between the postorbital and the frontoparietal massively expanded supraorbital shields in Brookesia corner. Note, however, that in Sphenodon punctatus superciliaris and other chamaeleons represent a combi- (Smith pers. obs.) and Gephyrosaurus bridensis (Evans nation of supraorbital flanges (see below) and supraor- 1980), the postorbital underlaps the postfrontal. I for- bital spines, for in at least some species the casque over mally regard the polarity to be ambiguous, but suspect the orbits is incompletely developed in young ontoge- that state 0 is primitive for crown Iguania, in which the netic stages, permitting the observation of both. posterior splint of the postfrontal has been lost and the Etheridge (1959) describes a similar phenomenon in lateral extent of the bone reduced. McGuire (1996) ten- certain large Anolis. tatively viewed state 1 as an autapomorphy of Crota- Supraorbital spines (state 2) are absent in most phytinae, while noting its appearance in certain other iguanians and in all outgroups examined. The lateral iguanians. exposure of the postorbital is highly restricted in Elgaria multicarinata, as in other autarchoglossans (Estes et al. 38. Postorbital III. cf. Estes et al. (1988) no. 18. (0) Jugal 1988), yet a distinct boss is present in this species. In and squamosal show little or no contact below postor- Plestiodon fasciatus, the postorbital is entirely excluded bital, even in adult specimens; (1) squamosal broadly from the orbit; yet, there is no tubercle on the posfrontal underlaps jugal beneath postorbital in adults. in the same position, so it is scored 0. The absence of In many iguanians, the squamosal and jugal remain such an eminence in Sphenodon punctatus and its separate (Figure 19A) or briefly overlap (Figure 19C). apparent absence in Diphydontosaurus avonis suggest In all agamids examined, however, the squamosal that state 1 also is derived. The character is ordered. broadly underlaps the jugal beneath the postorbital (Fig- ure 19D), as it does in Crotaphytinae. These species are 40. Postorbital V. Novum. Tubercle on posterior mar- scored 1. Note that in Iguana iguana and Dipsosaurus gin of orbit, when present, is located (0) at mid-height dorsalis, at least, squamosal–jugal contact seems to on the postorbital, or (1) near the top of the postorbital increase during ontogeny. D. dorsalis shows some vari- or on the postfrontal. ation, even bilateral (CAS 200863), in the amount of See discussion above. In crown Corytophaninae, underlap, but generally it is significant (de Queiroz Iguaninae and some Hoplocercinae, a tubercle is pres- 1987). ent but is developed at the orbital corner (state 1). The presence of state 0 in Sphenodon punctatus and Polarity is considered equivocal. most scleroglossans (in which the relevant bones are in contact at all) suggests that this state is primitive. In 41. Squamosal I. Robinson (1967), Estes et al. (1988) fossil rhynchocephalians, the postorbital, jugal and no. 34, Conrad and Norell (2007) no. 69. Dorsal process squamosal are not preserved in articulation; the facets (0) present or (1) absent. on the postorbital suggest the last two bones were in contact, but the extent of this contact is unknown. Estes et al. (1988) considered the dorsal process to be primitive for Squamata and the acquisition of a 39. Postorbital IV. Lang (1989) no. 6, 16, McGuire “hockey-stick-shaped” squamosal to be derived in Scle- (1996) no. 7, Smith (2009). Posterior margin of orbit (0) roglossa where it occurs. A dorsal process is present in smooth, (1) with small tubercle, or (2) with strong, ante- Huehuequetzpalli mixtecus (Reynoso 1998). Conrad riorly projecting supraorbital spine. and Norell (2007) considered the absence of a dorsal process to be a synapomorphy of several Late Creta- In many iguanids the lateral surface of the postor- ceous Mongolian taxa within Iguanidae. bital curves smoothly from its dorsal terminus near the frontoparietal suture to its ventral margin where it is Polarization follows Estes et al. (1988). sutured with the jugal. In some taxa there is a distinctly raised and often rugose area of bone—a tubercle—near 42. Frontal I. Moody (1980) no. 31, de Queiroz (1987) mid-height on the lateral surface. This tubercle is prob- no. 9, Lang (1989) no. 1, Poe (2004) no. 64. Frontonasal ably an attachment site for skin and connective tissue fontanelle (0) absent in adult specimens or (1) present in (Oelrich 1956). In other taxa, a tubercle is developed, adult specimens. but it is found higher up on the postorbital or is even There is generally no gap between the frontal and developed on the postfrontal (e.g., Enyalioides oshaugh- nasals in adult squamates. In a few taxa, however, there nessyi: Figure 24A). Thus, the tubercle shows alterna- is a persistent, median frontonasal fontanelle in adults. 280 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Dipsosaurus dorsalis has a pair of a foramina larger than 30660 [DigiMorph.org 2002–2005]) it seems concave normal near the frontal–nasal suture (de Queiroz 1987) up, but the edges are marked by thick, rounded, perior- that are not homologous. A fontanelle occurs in Hoplo- bital ridges. It is tentatively scored 1. The dorsal surface cercinae (de Queiroz 1987; Lang 1989), Plica (Frost of the frontal of the lepidosauromorph Marmoretta 1992), Corytophanes (Lang 1989), some small Scelo- oxoniensis, described by Evans (1991:396), is “flat...with porus, and a few agamids (e.g., Gonocephalus abbotti); upturned lateral edges.” Published descriptions and fig- see Poe (2004) on Anolis. Many chamaeleonids show a ures of other fossil rhychocephalians (Cocude-Michel large pair of frontonasal fontanelles (e.g., Rieppel 1987; 1963; Evans 1980; Whiteside 1986; Fraser 1988) leave Rieppel and Crumly 1997); these structures are presum- doubt as to the expression of this character, and the orig- ably not homologous, and in any case Brookesia super- inal specimens could not be examined. The frontals are ciliaris happily lacks them. generally flat in Autarchoglossa, but transverse concav- The absence of a frontonasal fontanelle in most ity is variably expressed in Gekkota (Smith 2009). This scleroglossans and in Sphenodon punctatus implies that character is left unpolarized. state 0 is ancestral. 46. Frontal V. Smith (2009). Supraorbital flanges are 43. Frontal II. Estes et al. (1988) no. 6. Frontals in late (0) absent or very poorly developed or (1) moderately to ontogeny (0) paired or (1) azygous. well developed. Polarization follows Estes et al. (1988). Smith (2009) discusses the distribution of supraorbital flanges in Iguania and other taxa, but he did not 44. Frontal III. Estes et al. (1988) no. 7. Frontals (0) with characterize them as precisely as is done here. It is nearly straight orbital margins or (1) constricted important to make a distinction between those species between orbits, giving them an hourglass shape. with broad cristae cranii and those with true supraorbital flanges. Flanges project beyond the margin of the Constriction is noticeable even in taxa with broad cristae and are distinguished from broad cristae by a supraorbital flanges, because the frontal expands in change in orientation; in transverse cross section, the width anterior to midorbit. ventral surface of the lateral edge of the frontal is con- Polarization follows Estes et al. (1988). cave (Figure 25). In contrast, iguanines (to choose one example) have broad cristae cranii that superficially look 45. Frontal IV. Smith (2009). (0) Interorbital portion like flanges, but whose ventrolateral surfaces lack this of frontal flat in transverse section in adult specimens; concavity and instead are straight or convex in trans- (1) frontal margins turned upward, conferring on the verse cross section. As described here, supraorbital frontal a concave-up shape in transverse section. flanges are restricted to Polychrotinae*, Corytophani- In some species there is an ontogenetic transforma- nae, Microlophus occipitalis (borderline) and Phryno- tion from state 1 to 0 (e.g., Iguana iguana), so it is wise soma platyrhinos (but not P. asio) among included to restrict consideration to ontogenetically mature spec- Iguanidae, as well as certain acrodontans (Brookesia imens. But the transformation is far from universal, superciliaris and Physignathus cocincinus, but not even in closely related forms (the frontal is flat in Priscagama gobiensis). Ctenosaurus similis in specimens ranging in SVL from Supraorbital flanges are absent in Sphenodon punc- 99 to 263 mm). In some cases periorbital ridges are tatus (Smith pers. obs.) and Diphydontosaurus avonis developed on the frontal (e.g., Leiocephalus), which give (Whiteside 1986, fig. 9d). They are present in Elgaria the dorsal surface a superficially concave appearance; multicarinata, but absent in many other anguids, when significant rugosities or other eminences are pres- including the gerrhonotine Mesaspis morelettii. They ent, then, the foundational surface of the frontal alone are present in Eublepharis macularius and several other must be considered. State 1 is found in Corytophaninae gekkotans. For the present I regard the polarity to be (Figure 18A), Anolis (Figure 18B) and Polychrus, but is ambiguous. absent in Leiosaurini. It is difficult to see in Laemanctus longipes, because rugosities on the dorsal surface of the 47. Dermal skull rugosities. Etheridge and de Queiroz frontal, which are developed centrally and not specifi- (1988) no. 7, Frost and Etheridge (1989) no. 7, Pregill cally periorbitally, tend to make the surface of the frontal (1992, no. 11), McGuire (1996) no. 37. Rugosities (0) seem taller centrally than it really is; even so, the degree absent or restricted to skull roof, or (1) extensive and of concavity seems to be low in this species. State 1 is found on other dermal skull bones besides the frontal also found in Priscagama gobiensis (Borsuk-Bialynicka and parietal. and Moody 1984; Smith pers. obs.) and in some Frost and Etheridge (1989) coded Leiocephalus ? agamids (notably Australian taxa and Physignathus), because of interspecific variability. According to Pregill’s and variably in Leiolepis belliana, but Uromastyx and (1992) phylogenetic analysis of the taxon, extensive other agamids evince state 0. Chamaeleonids, including rugosities are not primitive, but rather evolved three Brookesia superciliaris (Smith pers. obs.), frequently times independently within Leiocephalus. have upturned orbital margins (e.g., Frank 1951). If these kinds of dermal rugosities arise from The frontal of Sphenodon punctatus is difficult to the interaction of the bone with overlying dermis, a rea- evaluate, for in adults (but scarcely in juveniles: see CM sonable case could be made for the presence of these Eocene Lizards of the Clade Geiseltaliellus • Smith 281 rugosities in Elgaria multicarinata and Plestiodon fasciatus, in which osteoderms underlie the broad cranial scutes. I tentatively score these species 1. Polarization follows Etheridge and de Queiroz (1988).

48. Parietal foramen I. Etheridge and de Queiroz (1988) no. 2. (0) Present or (1) absent. The parietal foramen is absent in some Polychrus (Etheridge 1959), but is present in P. acutirostris. A parietal foramen is found in Brookesia superciliaris (Sieben- rock 1893). Polarization follows Etheridge and de Queiroz (1989).

49. Parietal I. Novum. Parietal fontanelle (0) filled predominantly by parietal during ontogeny, or (1) partially filled by frontal, forming a median posterior projection on the frontal. A anteromedian parietal fontanelle is found in juvenile specimens (Edinger 1955). During ontogeny this space is generally filled by the parietal, with the exception of a small anterior notch for the parietal organ (Fig- ure 18B). However, in some taxa the frontal grows posteriorly to fill a significant portion of the fontanelle (Figure 18C) and sometimes to surround the parietal organ (see also next character). Agamids (including Leiolepis and Uromastyx) most clearly show the derived state, in which the frontal is seen to have a strong posterior projection on the mid-line (Figure 18C). Some Figure 25. Frontal and parietal of Anolis carolinensis Laemanctus longipes show a moderate posterior projec- in ventral view. Note the development of strong supra- tion of the frontal. Paired anterior projections of the orbital flanges and the subsumption into a nuchal parietal on either side of the mid-line cause the appear- casque of the supratemporal processes. Abbreviations: ance of state 1 in Basiliscus (Figure 18A); these species crcr, crista cranii; rpra, recessus processi ascendentis; are scored 0. sofl, supraorbital flanges; stpr, supratemporal process. The state in Sphenodon punctatus is difficult to eval- Scale bar is 3 mm. uate, but the presence of state 0 in most scleroglossans (and most iguanians) leads me to suggest that this state is ancestral. L. longipes also shows (better than L. serratus) the typically agamid condition of a posterior mid-line projec- 50. Parietal foramen II. Camp (1923), Etheridge tion on the frontal. It might be that it is the frontal has (1959), de Queiroz (1987) no. 13, Etheridge and de been altered, not the position of the foramen, which is Queiroz (1988) no. 2, Frost and Etheridge (1989) no. still in line with the trend of the suture. Information on 11, Lang (1989) no. 17, Poe (2004) no. 60. Parietal fora- the developmental mechanics underpinning the ulti- men located (0) in parietal, (1) contiguous with but mate location of the parietal organ is needed. slightly posterior to frontoparietal suture, invading only Many iguanids show state 2, but more rarely state parietal, (2) contiguous with frontoparietal suture, 3 obtains (Etheridge 1959; de Queiroz 1987; Lang invading frontal, if not parietal, or (3) foramen entirely 1989). In Anolis, the parietal foramen is generally con- confined to frontal, well anterior of frontoparietal tiguous with the frontoparietal suture, but invades only suture. the parietal (Figure 18B; Etheridge 1959; Williams The position of the parietal foramen has previously 1989). Anolis carolinensis (Figure 25) is an exception. been described relative to the frontoparietal suture (e.g., State 1 is also shown by Oplurus cuvieri and many other Camp 1923; Estes et al. 1988), a tradition maintained Oplurus (see Blanc 1977) and possibly some phrynoso- here. However, the recognition of a posterior projection matines. Laemanctus longipes is regarded as variable of the frontal in some specimens raises another possibil- (Lang 1989). ity: parsing this character for the main trend of the fron- Polarization follows Estes et al. (1988). The charac- toparietal suture, disregarding the posterior projection. ter is treated as ordered. Previous authors, for instance, have regarded Laemanc- tus as “polymorphic” because in L. longipes the foramen 51. Parietal II. Smith (2009). In adult specimens, the can be wholly in the frontal. Technically this is true, but recessus processi ascendentis (the “parietal fossa” of 282 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Oelrich 1956 and others) is (0) located at or just under tor crests, resulting in a casque, especially clearly devel- the posterior margin of the parietal, (1) well inset under oped in Corytophanes, some Laemanctus, and Poly- the posterior margin of the parietal, or (2) absent. chrus, as well as certain Anolis. It is difficult to determine whether the recess seems On the basis of outgroup comparison with Rhyn- to be inset in certain clades because the contact of the chocephalia and Huehuequetzpalli mixtecus (Reynoso processus ascendens of the synotic tectum (Oelrich 1998), state 0 is considered primitive. 1956) is shifted anteriorly, or because the parietal has grown posteriorly. In Anolis, especially, many species 54. Parietal V. cf. Estes et al. (1988) no. 54. Posterior to seem to have greatly foreshortened supratemporal mid-length for the parietal table, approximately where processes of the parietal, but this appearance seems to the parietal is connected to the epipterygoid, the descen- result from posterior growth of the parietal between sus parietalis extends (0) ventrolaterally or (1) ventrally. these processes (Figure 25). The homolog of the This character more or less corresponds to whether supratemporal process is embedded within the body of the origin of the jaw adductor musculature on the pari- the parietal, recognizable from the increased thickness etal is dorsal or ventral. Its reformulation for the descen- of that bone in anteromedially convergent lines contin- sus parietalis, however, allows certain taxa with strongly uous with the distinct supratemporal processes posteri- developed adductor crests (e.g., Corytophanes, Phryno- orly; this homology statement may be further tested by soma, Uromastyx and Polychrus) to be coded which studying the insertion sites of the nuchal musculature might otherwise be ambiguous. All iguanians lacking relative to other squamates (see Oelrich 1956; Tsuihiji strong adductor crests show state 0. 2005). Thus, in Anolis (also Corytophaninae and other polychrotines), independent data suggest that signifi- Polarization follows Estes et al. (1988). Acquisition cant posterior growth of the parietal contributes to the of state 1 is a synapomorphy of Scleroglossa, conver- inset of the recessus. State 1 occurs in certain larger gently acquired in certain iguanians (Phrynosoma, Uro- forms, in particular some iguanines and agamids, whose mastyx). smaller relatives show state 0; in some clades there is an especially strong ontogenetic component to variation in 55. Parietal VI. Etheridge and de Queiroz (1988) no. 3, the position of the recess. Phrynosoma platyrhinos, Phy- Frost and Etheridge (1989) no. 10, Lang (1989) no. 19. maturus palluma and Liolaemus pictus lack a clear Parietal (0) lacks median blade or (1) has one. recessus processi ascendentis. A median parietal blade is present in many Eublepharis macularius lacks the recess, because the chamaeleonids, but not in Brookesia superciliaris. Polar- processus ascendens in this species, like the rest of ization follows Etheridge and de Queiroz (1988). Gekkota (Bellairs and Kamal 1981), is lacking. The recess is deeply inset in many anguimorphs, including 56. Parietal VII. Lang (1989) no. 20, Frost and Elgaria multicarinata. In some scincomorphs (Cordyli- Etheridge (1989) no. 10. Parietal blade, when present, formes and Xantusiidae: Smith 2006b) the recess is develops (0) postembryonically or (1) embryonically. deeply inset, but in others this is not so. I formally regard This features is inapplicable to species lacking a polarity to be ambiguous. parietal blade. Although chamaeleons with a parietal blade are not included here, note that Brock (1941) 52. Parietal III. cf. Etheridge (1959, 1966), Etheridge described a specimen of Bradypodion in which a small and de Queiroz (1988) no. 3, Frost and Etheridge (1989) parietal crest had developed by a late embryonic stage. no. 10, Lang (1989) no. 19, Poe (2004) no. 57. In adult The character is treated as unpolarized because of specimens, parietal table is (0) trapezoidal, (1) V-shaped, the absence of data from outgroups. or nearly so, or (2) Y-shaped. The recognition of parietal growth between the 57. Parietal VIII. Etheridge and de Queiroz (1988), supratemporal processes suggests this character might Lang (1989). Posttemporal fenestra (0) open or (1) need to be reconceived in the future. closed by parietal. Sphenodon punctatus is considered to show state 2, State 1 is an autapomorphy of Corytophanes among although juvenile specimens (see CM 30660 [Digi- Iguania (Lang 1989). Morph.org 2002–2005]) seem to show state 0. Polariza- Polarization follows Lang (1989). tion follows Etheridge and de Queiroz (1988). The character is treated as ordered. 58. Supratemporal I. de Queiroz (1987), Etheridge and de Queiroz (1988) no. 9, McGuire (1996) no. 10, Frost 53. Parietal IV. Etheridge and de Queiroz (1988), Lang and Etheridge (1989) no. 12. Anterior process of (1989). Adductor crests of parietal (0) weakly developed supratemporal located (0) predominantly on anterolat- or (1) strongly developed, projecting over the descen- eral surface of supratemporal process of parietal or (1) sus parietalis. predominantly on posteromedial surface. Nearly all taxa showing a ventral origin of the jaw The posterior end of the supratemporal in Squa- adductor musculature show state 1, but several taxa with mata generally wraps around the end of the supratem- a more or less dorsal origin (ventrolaterally directed poral process of the parietal, but the disposition of the descensi parietalis) also show strongly developed adduc- anterior process of the bone differs (de Queiroz 1987). Eocene Lizards of the Clade Geiseltaliellus • Smith 283

The depth of the facet in which the supratemporal bone articulates also differs (e.g., Frost and Etheridge 1989; McGuire 1996), but the facet’s depth is here regarded as a different character (which is not further systematized). In some taxa with a deep supratemporal facet, the anterior process is strongly reduced (e.g., Liolaemus: see Frost and Etheridge 1989, fig. 2C) and such taxa are scored ?. In contrast, Crotaphytus, which also has a deep supratemporal facet but retains the anterior process of the bone (McGuire 1996), is scored 0. I agree that the supratemporal in Oplurus and Chalarodon is somewhat posteromedially displaced, but I consider them still to show state 0. Leiolepis shows state 1. Although the condition in Sphenodon is not so easily disposed in this scheme, the anterior process of the supratemporal lies lateral to the portion of the parietal that forms the dorsal edge of the supratemporal process. Polarization follows Frost and Etheridge (1989).

59. Quadrate I. Lang (1989) no. 25. A medial concha is (0) well developed or (1) absent, or nearly so. Many iguanians have a distinct medial concha on the quadrate, and in certain iguanines (Iguana, Cyclura) it is greatly expanded. The concha is often better developed dorsally than ventrally (Figure 26). Among previously identified clades, loss of the concha is nearly Figure 26. Right quadrate of Enyalioides oshaugh- confined to Anolis, Polychrus, Corytophanes, Laemanc- nessyi (SMF 67590) in posterior view. Abbreviations: tus (Lang, 1989), Phymaturus, Phrynosoma and some cnd, dorsal or cephalic condyle; cnv, ventral condyle; “sand lizards.” Geiseltaliellus maarius (see Figure 3) has col, lateral concha; com, medial concha; lr, lateral a medial concha. The quadrate of chamaeleons is too ridge; pcr, posterior crest; tcr, tympanic crest. Scale bar transformed to score. is 1 mm. The morphology of the quadrate in Sphenodon punctatus and other rhynchocephalians cannot be acco- tympanic crest can be taller than the posterior crest.) In modated in this system, and they are scored ?. The Dipsosaurus dorsalis, hoplocercines, tropidurines, medial concha is absent in Eublepharis macularius and phrynosomatines (Figure 19A) and oplurines, the ven- Plestiodon fasciatus but is present in Elgaria multicari- tral condyle tends to be located more anteriorly, so that nata. The character is left unpolarized. it seems as if the cephalic condyle had been rotated back- wards on the ventral pivot. The tympanic crest extends 60. Quadrate II. Lang (1989) no. 26. In lateral view, pos- more vertically than the posterior crest. but reaches terior crest is (0) posteriorly concave or (1) vertical, or approximately the same horizontal level, so it is distinctly nearly so. shorter than the latter (Figure 26). In juveniles of Iguana The derived state is restricted in distribution to iguana the quadrate is distinctly rotated, but the vertical Corytophanes, Laemanctus (Lang 1989), Polychrus, Ano- condition occurs in adults. This ontogenetic shift is con- lis ricordi and Brookesia, among ingroup taxa. sistent with previous observations (Bellairs and Kamal Polarization follows Lang (1989). 1981; Rieppel and Zaher 2000). The quadrate in Sphenodon punctatus is vertical, 61. Quadrate III. Lee (1998) no. 50, Rieppel and Zaher but otherwise difficult to fit into this scheme, and the (2000) no. 50. Orientation: (0) vertical condition or (1) disarticulated and fragmentary nature of fossil rhyncho- rotated condition. cephalians further complicates the evaluation of polarity. In contrast, the tympanic crest of the quadrate seems In acrodontans, crotaphytines, corytophanines and to arc broadly anteriorly in most scleroglossans, and the polychrotines, as well as many iguanines and large ventral condyle is anterior of the cephalic condyle. I Priscagama gobiensis, the ventral condyle of the quadrate leave this character unpolarized. is located in nearly the same transverse plane as the cephalic condyle (Figure 19D). Because the tympanic crest is vertical and its dorsal edge generally reaches close 62. Quadrate IV. Novum. A weak ridge of bone on the to the same height as the cephalic condyle, this means posterior surface of the lateral concha is (0) absent or that the tympanic crest is approximately as tall as the (1) present. quadrate itself. (In adult Agamidae, the ventral condyle In some taxa, notably corytophanines, many poly- is often posterior to the level of the cephalic one and the chrotines and Enyalioides oshaughnessyi (Figure 26), as 284 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 well as Leiolepis belliana and many agamines, there is a from the lateral edge of these elements to the pyriform low ridge of bone that extends dorsolaterally from recess (state 0). In a few taxa, primarily Iguaninae, this approximately mid-height on the posterior crest. The suture has a kink, extending first anteromedially, then reason for this ridge is unclear, although in my speci- posteromedially for a short distance, and finally antero- men of E. oshaughnessyi and others, where the bone is medially again until it reaches the pyriform recess. In semi-translucent, it seems that a small intraosseous some taxa, the kink occurs in line with a ridge on the canal runs through the ridge. In one Anolis ricordi (UF palatine that would probably bear the teeth if these were 64820), the ridge extends only partially, and at its ter- present. State 2 is primarily found in Anolis, but also in mination a vessel exits through a foramen and contin- some Sceloporus. ues in the same direction as the ridge in a tiny groove. In The states for Eublepharis macularius and one Petrosaurus thalassinus (CM 38327), a small fora- Plestiodon fasciatus are difficult to fit into this scheme; men on the lateral side of the posterior crest opens into the species are scored ?. Sphenodon punctatus and a tiny groove that runs dorsolaterally in the same direc- Elgaria multicarinata show state 0, as does Diphydon- tion as the ridge in other taxa; a small foramen is pres- tosaurus avonis, on the basis of the palatine (Whiteside ent in a similar position in many other taxa lacking the 1986). This state is consequently considered primitive. ridge, although it is only rarely accompanied by a The character is treated as unordered. groove. The ridge probably indicates that a plesiomor- phically present nerve or blood vessel has taken an 66. Pterygoid I. Etheridge (1959), de Queiroz (1987) intraosseous course. State 1 was also found in some no. 51, Etheridge and de Queiroz (1988) no. 18, Frost Sceloporus and Ctenosaura. Brookesia superciliaris is and Etheridge (1989) no. 28. Teeth (0) usually present or scored ?, because the lateral concha in this, as in other (1) usually absent. chamaeleons, is highly reduced. Frost and Etheridge (1989) and Pregill (1992) noted The character cannot be evaluated in Sphenodon variability in the presence of pterygoid teeth in Leio- punctatus, because its morphology is so radically differ- cephalus; I found them to be absent in both L. personatus ent. In fossil rhynchocephalians, the quadrate is gener- and L. melanochlorus. I also found the presence of ptery- ally incomplete or the posterior surface of the lateral goid teeth in Anolis cristatellus and A. chlorocyanus to be concha inscrutable. The ridge, however, was not discov- polymorphic, but Poe (2004) found them to be present ered in any scleroglossan examined, suggesting that state in all his A. chlorocyanus and most A. cristatellus; for 0 is ancestral. now, these species are scored 0. 63. Vomer I. Moody (1980) no. 55, Frost and Etheridge Polarization follows Etheridge and de Queiroz (1989) no. 4. (0) Flat or convex or (1) ventrally concave. (1988). Polarization and coding follows Frost and Etheridge (1989). 67. Nasal capsule I. Hallermann (1994) no. 1. Fenestra septi nasi (0) absent, (1) slit-like or (2) large (half or 64. Palatine II. Etheridge (1959), Estes et al. (1988) no. more of total length of septum nasi). 82, Etheridge and de Queiroz (1988) no. 17, Frost and The cartilaginous, median septum nasi divides the Etheridge (1989) no. 27, McGuire (1996) no. 32; see also nasal capsule into left and right halves. In many iguan- Frost et al. (2001) no. 66. Teeth (0) generally absent or ian taxa the septum is imperforate, but in others there is (1) often present. an opening in the posterior portion. This fenestra can be Frost et al. (2001) noted variability in the appear- narrow and slit-like, or larger and more equidimen- ance of palatine teeth in many polychrotines. Palatine sional, and probably arises through resorption of carti- teeth are very rarely present in some Oplurus (in my O. lage in ontogeny (Bellairs and Kamal 1981; Rice 1920; quadrimaculatus, just 1 tooth, unilaterally; cf. Frost and Hallermann 1994). Rice (1920) in particular studied Etheridge 1989); they were absent in O. cuvieri. many developmental stages in Plestiodon fasciatus and Etheridge and de Queiroz (1988) noted that out- found that the septum becomes a little thinner late in group comparison provides ambiguous evidence for ontogeny, followed by the relatively abrupt appearance polarization of this character: rhynchocephalians typi- of a large fenestra. It is possible, then, that the study of cally have palatine teeth, while outside Iguania they are additional stages in the ontogeny of some ingroup known to be present only in one extant and two fossil species may require modifications to the scoring used species of anguimorph. I follow Estes et al. (1988) in here. I have scored Anolis cristatellus as unknown (cf. polarizing this character and regard palatine teeth, Hallermann 1994). where present, as a reacquisition. According to Hallermann (1994), a fenestra septi nasi is absent in Sphenodon punctatus, but is found in 65. Palatine III. Novum. Palatine–pterygoid suture, in many scleroglossans. The character is treated here as ventral view, (0) is a straight, anteromedially trending unpolarized, but ordered. line, (1) trends anteromedially, but with a strong kink, or (2) is primarily a horizontal line. 68. Nasal capsule II. de Beer (1937), Malan (1946), H. In many iguanians, the ventral exposure of the pala- (0) Fenestra superior nasi perforates tectum nasi; (1) tec- tine–pterygoid suture extends as a straight, oblique line tum nasi whole. Eocene Lizards of the Clade Geiseltaliellus • Smith 285

The dorsal roof of the anterior portion of the nasal capsule (Figure 27A) is perforate in some iguanians. In the species examined by Hallermann (1994), a fenestra was found only in Phymaturus palluma, Leiocephalus, Basiliscus basiliscus, Corytophanes cristatus and Physig- nathus cocincinus. Malan (1946: 84) described Uta stansburiana as showing “incipient” fenestration in this area, but did not elaborate; she also found a large one in Phrynosoma douglassi. Outgroup comparison is ambiguous: perforate in Sphenodon punctatus (Werner 1962) and in many, but far from all scleroglossans. Malan (1946) and Haller- mann (1994) felt it was primitive for Lepidosauria. I treat it as unpolarized.

69. Nasal capsule III. Hallermann (1994) no. 2. Pre- nasal process (0) present or (1) absent. The processus praenasalis (“rostrum” of Oelrich 1956) is a small, cartilatinous projection from the anterior end of the nasal capsule (Figure 27A, B). This process is present in all iguanids and absent in all acrodontans examined by Hallermann (1994). Polarization follows Hallermann (1994).

70. Nasal capsule IV. Hallermann (1994) no. 7. M. nasalis externus (0) absent or (1) well developed. This muscle originates fleshily on the anterolateral surface of the facial process of the maxilla and inserts in and on various structures surrounding the fenestra exonarina (including the skin and the processi alaris superior and inferior). This feature is unique to Acrodonta Figure 27. Nasal capsule in iguanians. A, Nasal cap- among taxa studied by Hallermann (1994), who implied sule of Crotaphytus collaris (after Hallermann 1994) in that it may be an adaptation to hot, dry climates, like Steb- dorsal view. On the left, the roofing cartilages are bins (1948) did for the nasal valve (see below). shown intact, on the right they are removed to reveal Polarization follows Hallermann (1994). the passages of the vestibulum, cavum nasi and glan- dula nasalis lateralis. B, Same, in ventral view. On 71. Nasal capsule V. Stebbins (1948), Hallermann the left, only the cartilages septomaxilla are shown; (1994) no. 8. Nasal valve (0) absent or poorly developed, the right side shows nonskeletal structures. Abbrevia- or (1) well developed. tions: cco, cavum conchale; co, nasal concha; cse, commissura sphenethmoidalis; cvp, commissura The wall of the vestibulum nasi (“anterior chamber” vomeronasalis posterior; dl, lacrimal duct; ech, exter- of Oelrich 1956) in most iguanians is permeated with nal choana; en, external nares; gnl, lateral nasal gland; radially arrayed smooth muscle fibers that lie alongside jo, opening of Jacobson’s organ; nc, cavum nasi; nem, vascular lacunae and space-filled connective tissue medial ethmoidal nerve; prpa, processus rostralis plani (Hallermann 1994). In phrynosomatines, the floor of antorbitalis; prpn, processus praenasalis; sm, sep- the vestibulum is modified into a special valve that, tomaxilla; vs, vestibulum nasi. Scale bar is 0.5 mm. on erection, functions to narrow the nasal passage (Stebbins 1948). Hallermann’s (1994) observations on Petrosaurus thalassinus contradict those reported by Etheridge and de Queiroz (1988); the supposed absence The cleft between the parietotectal and paranasal of a nasal valve, according to the latter, would put Pet- cartilages is named the fissura lateralis nasi. In many rosaurus outside of the remaining Phrynosomatinae. iguanians it is fully developed and plugged by connec- Stebbins (1948) notes that a well-developed nasal tive tissue or the lateral nasal gland, or both, but in many valve is present in Elgaria multicarinata, but is only others it is partly closed during the course of ontogeny. “moderate” in Plestiodon skiltonianus. Polarization fol- In yet others it is lacking entirely. lows Hallermann (1994). Outgroup ambiguity (not developed in Sphenodon punctatus or autarchoglossans, developed in many 72. Nasal capsule VI. Hallermann (1994) no. 10. Fis- gekkotans: Hallermann 1994) requires that the charac- sura lateralis nasi (0) fully developed, (1) partially closed ter be treated as unpolarized; it is, however, treated as or (2) absent. ordered. 286 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

73. Nasal capsule VII. Hallermann (1994) no. 11. Con- and Chamaeleonidae) the commissure is entirely absent cha nasalis (0) well-developed, with cavum conchale, (1) (Malan 1946; Hallermann 1994). reduced to a ridge, cavum conchale reduced, or (2) con- Polarization follows Hallermann (1994). cha nasalis absent. In squamates in which it is well developed, the con- 76. Nasal capsule X. Hallermann (1994) no. 16. Com- cha nasalis is an invagination of the wall of the nasal cap- missura vomeronasalis posterior is (0) incomplete or sule, forming a cavum conchale that holds the lateral absent, or (1) present and complete. nasal gland (Figure 27A). In many iguanians the con- This commissure extends posterior to Jacobson’s cha is small in size, and the cavum conchale accordingly organ and connects the lamina transversalis anterior diminished; more rarely (Chamaeleonidae, Anolis, Poly- with the cartilago paraseptalis (Figure 27B). Haller- chrus and many or all phrynosomatines), these struc- mann’s (1994) coding is different than mine; his states tures are absent altogether (Hallermann 1994). were absent, present, and lost; that is, he assumed that Malan (1946:74) concluded that a concha nasalis the lack of a commissure in some Acrodonta, Poly- (present in Scleroglossa and Sphenodon punctatus, chrotinae*, and Corytophaninae represents reversal. I seemingly contra de Beer 1937) with a cavum conchale do not make this assumption and instead code these lat- (present in Scleroglossa, absent in S. punctatus) is ances- ter taxa as state 0. State 0 thus obtains in most agamids tral for Lepidosauria and Squamata. Hallermann (1994) studied by Hallermann (except Uromastyx acanthinu- followed her lead. Given that the concha is represented rus and an Australian agamid) and in Chamaeleonidae only by a ridge in S. punctatus and many agamids as well as in Anolis, Polychrus and Corytophaninae. (excepting three taxa Hallermann examined, one of Polarization follows Hallermann (1994). which is Physignathus cocincinus), and is absent in Chamaeleonidae, it is not obvious to me that the presence of a concha with a cavum conchale (state 0) should 77. Nasal capsule XI. Hallermann (1994) no. 17. Con- be considered primitive for Squamata. This character is cha of Jacobson’s organ (0) present (well developed or left unpolarized. It is ordered. not) or (1) absent. A solid cartilaginous projection of the lamina trans- 74. Nasal capsule VIII. Malan (1946), Hallermann versalis anterior enters the lumen of Jacobson’s organ, (1994) no. 14. (0) Ventral margin of planum antorbitale dividing it into upper and lower portions (Oelrich straight or nearly so, (1) a strong process projects ante- 1956). Although its shape differs considerably, a concha riorly from the ventral margin of the planum antorbitale is developed in most iguanians. It is absent, however, in (connected by cartilage with the lamina transversalis Anolis, some Chamaeleonidae and rare agamids anterior or not). (Hallermann 1994). Brock (1941) notably found that This generally narrow, cartilaginous process proj- Jacobson’s organ is much better developed in Bradypo- ects anteriorly from the ventral margin of the planum dion occidentale than in Chamaeleo. antorbitale, lateral to the paraseptal cartilage but medial A concha is present in scleroglossans (Hallermann to the choanal groove, which it supports (Figure 27B). It 1994), but is lacking in Sphenodon punctatus. Polariza- was present in all iguanids studied by Hallermann tion follows (Hallermann 1994). (1994) except Basiliscus and was absent in almost all acrodontans. Hallermann’s figure of Basiliscus basilis- 78. Nasal capsule XII. Hallermann (1994) no. 18. Car- cus (1994, fig. 13) inclines me to view it as having this tilago paraseptalis (0) present and complete or (1) process as well, although it is certainly not as well grossly incomplete or absent. demarcated or acutely pointed as in other iguanids. In The paraseptal cartilage runs parallel to the ventral a few taxa (Phrynosoma and Petrosaurus) the processus margin of the septum nasi, connecting the medial mar- rostralis plani antorbitalis is connected anteriorly by car- gin of the planum antorbitale with the lamina transver- tilage or tough connective tissue with the lamina trans- salis anterior (Figure 27B). It is present in most versalis anterior (Hallermann 1994). iguanians, but grossly incomplete or entirely absent in a The process is lacking in Sphenodon punctatus few taxa (Anolis, Polychrus, Tropidurus melanopleurus, (Bellairs and Kamal 1981, fig. 43A). Polarization follows some agamids and Chamaeleonidae; Hallermann 1994). Hallermann (1994). Polarization follows Hallermann (1994).

75. Nasal capsule IX. Malan (1946), Hallermann (1994) 79. Nasal capsule XIII. Stebbins (1948), Frost and no. 15. Commissura sphenethmoidalis (0) well devel- Etheridge (1989) no. 53–57, Hallermann (1994) no. 20. oped, fully connected with interorbital cartilages, or (1) Vestibulum nasi (0) short, with rostral entrance into poorly developed, that is, incomplete or absent. cavum nasi, (1) intermediate in length, with entrance The commissura sphenethmoidalis is a strut of car- into cavum nasi just posterior to Jacobson’s organ, or tilage that connects the posterior surface of the planum (2) elongate, about as long as nasal capsule or longer. antorbitale with the interorbital cartilages. In at least This character describes only the length of the some taxa (e.g., Crotaphytus collaris; Figure 27A) it is vestibulum (“anterior chamber” of Oelrich 1956), which poorly developed in very small embryos, but complete connects the external naris with the cavum nasale; the in hatchlings (Hallermann 1994). In a few taxa (Anolis junction between vestibulum and cavum is generally Eocene Lizards of the Clade Geiseltaliellus • Smith 287 marked by the duct of the lateral nasal gland (Figure the root of the problem, for Etheridge (1959) presum- 27A). (The geometry of the vestibulum is described in ably examined adult individuals, Hallermann (1994) the next character.) embryos. Even if ontogeny is the problem, even embry- Born (1879) described the nasal capsule of one onic phrynosomatines show the peculiar morphology “Leiosaurus Bellii” whose vestibulum is similar to that characteristic of that clade (Stebbins 1948; Hallermann of Liolaemus pictus. The specific identity of the speci- 1994); it is possible that corytophanines show delayed men has been questioned (Hallermann 1994:59), extension of the vestibulum compared to other because Born failed to mention the supposed and pre- iguanids. Pending further study, I regard the length of sumably obvious lack of a septomaxilla; Hallermann the vestibulum in B. basiliscus, Corytophanes cristatus, (1994) suggested that he may actually have had a spec- Polychrus acutirostris and Anolis cristatellus as imen of Liolaemus belli. It is unlikely to have been a slip unknown. of the pen, as Born (1879) repeatedly uses the generic Stebbins (1948:201) notes that the vestibulum of name Leiosaurus. Furthermore, he (Born 1879:111) Plestiodon skiltonianus is “exceptionally short,” which specifically indicates that the species was named by is also characteristic of many lygosomines (Gabe and “DB” (Dumeril et Bibron). Finally, the specimen of Saint Girons 1976). The “scincine” Chalcides mionecton Leiosaurus bellii available to me does not lack a sep- has a longer vestibulum, but one that still enters the pri- tomaxilla. Thus, there is little reason to believe that the mary nasal cavity at the level of Jacobson’s organ (Gabe species identification, Leiosaurus bellii, was inaccurate; and Saint Girons 1976). Bracketing therefore suggests its seemingly advanced vestibulum could represent an that the vestibulum of Plestiodon fasciatus ought also independent adaptation to the relatively arid habitat of to be short, and it is scored 0. Stebbins (1948) found the this species (see van Devender 1977). The examination vestibulum to be shorter in Elgaria multicarinata than of the nasal capsule of mesic, sylvan Enyalius (Etheridge in any other studied species except Plestiodon skiltoni- 1969) and (western) Pristidactylus (Etheridge and anus; the former, like the latter, is scored 0. Polariza- Williams 1985) may shed light on this problem. tion follows Hallermann (1994). The character is The phylogenetic position of chamaeleonids ordered. whose nasal capsules have been studied is frustrating. Rieppel and Crumly (1997) hypothesized that Malagasy 80. Nasal capsule XIV. Stebbins (1948), Frost and Brookesia is the sister taxon to the remainder of Etheridge (1989) no. 53–57, Hallermann (1994) no. 20. Chamaeleonidae; of that remainder, mainland African Geometry: vestibulum nasi is (0) straight, apart from Rhampholeon (formerly in Brookesia) is the sister taxon the inturning near the external naris, (1) S-shaped in a to the rest. In species of Chamaeleon (Pratt 1948), the horizontal plane, or (2) U-shaped in a vertical plane vestibulum is elongate and enlarged, in the shape of an (“sink trap”). S. However, Gabe and Saint Girons (1976:23) note that This character describes how increased vestibular the vestibulum in Rhampholeon spectrum is “fairly length is accomodated in the nasal capsule. A short short,” as it is in Bradypodion occidentale (Brock 1941). vestibulum is probably closely linked to a straight one, If Malagasy Brookesia also have a short vestibulum, it but even a vestibulum of intermediate length (e.g., seems possible that the elongate, S-shaped vestibulum Iguana iguana) can be S-shaped. The S-shaped condi- evolved independently within Chamaeleonidae as an tion is common among ingroup species with a vestibu- adaptation to arid environments (Pratt 1948; Lemire lum of moderate to great length (Figure 27A). In 1985). Note that the vestibulum of Physignathus cocinc- phrynosomatines, however, the vestibulum extends inus also is short (Hallermann 1994). Hallermann posterodorsally to the end of the nasal capsule before (1994:46) suggested that an S-shaped vestibulum is turning ventrally or anteroventrally (Stebbins 1948; independently derived in Uromastyx, “higher” Lemire 1985); only on the ventral leg does the vestibu- chamaeleons (e.g., Chamaeleo) and iguanids. This is a lum enter the cavum nasi. This geometry corresponds to very reasonable interpretation of the data, but further the “sink trap” of Stebbins (1948). It is also developed examination of basal acrodontans will be necessary to in Agama agama (Lemire 1985). Etheridge’s (1959) confirm it. observations on the vestibulum of Polychrus acutirostris, As discussed also below, there is conflicting evi- which he describes as showing a geometry nearly like dence on the length and geometry of the vestibulum in that in Phrynosomatinae, conflict with Hallermann’s Corytophaninae and Polychrotinae. In particular, (1994), as noted also above. In the case of Anolis spp., Etheridge (1959) describes the vestibulum of Polychrus however, Stebbins (1948), Etheridge (1959) and Haller- acutirostris as highly elongate and that of Basiliscus vit- mann (1994) agree that the vestibulum in many species tatus as intermediate in length. Possibly it is a semantic is short and straight, whereas Gabe and Saint Girons problem on the exact meaning of terms like “elongate.” (1976:20) found the vestibulum of A. cristatellus to be Alternatively, it might be a problem of geometry: the “relatively long.” Clarification is needed. Pending fur- external nares of Polychrus are posteriorly displaced on ther study, I regard the geometry of the vestibulum in the snout (Etheridge 1959; Smith 2006a), so the vestibu- Basiliscus basiliscus, Corytophanes cristatus, P. acu- lum can form an anteriorly directed U, which is differ- tirostris and A. cristatellus as unknown. ent from the corytophanine condition. Finally, Polarization follows Hallermann (1994). This char- dependency on different ontogenetic stages could lie at acter is treated as unordered. 288 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

81. Septomaxilla I. Lang (1989) no. 2, Hallermann A highly reduced crista proötica is found in all (1994) no. 30. (0) present and well developed; (1) very examined hoplocercines as well as scattered small or absent. tropidurines, Corytophanes cristatus and Petrosaurus Polarity follows Lang (1989). thalassinus, among ingroup taxa included here. Outgroup comparison, and rarity, suggest state 1 is 82. Saccus endolymphaticus. Camp (1923), Etheridge derived within Squamata. (1959), Hallermann (1994) no. 34. Endolymphatic sacs (0) located wholly within cranial cavity or (1) extending 87. Braincase IV. Frost et al. (2001) no. 58. Posterior into neck musculature. processes of basisphenoid are (0) well developed and Camp (1923) and Etheridge (1959) had long distin- contribute to anterior margin of sphenoöccipital tuber- guished anoles and certain scleroglossans by the exten- cle or (1) strongly reduced. sion of calcified endolymphatic sacs into the nuchal These processes are well developed in most ingroup musculature. In Agaminae extension of the sacs occurs taxa. The only examined taxa showing significant reduc- through the epiotic foramen (Moody 1980; but see tion of the processes are Anolis, some Polychrus (Frost Hallermann 1994). Hallermann (1994) distinguished et al. 2001), Liolaemus pictus, and Leiolepis belliana. I between merely calcified endolymphatic sacs and those find P. marmoratus to have them. that also extend into the neck musculature, and found These processes are well developed in Sphenodon the former in several other iguanian taxa (especially punctatus (Smith pers. obs.). Their presence in some corytophanines, certain tropidurines, and Chamaeleo). fossil lepidosauromorph outgroups is uncertain because Apparently calcified deposits occur intracranially in sev- of breakage (e.g., Whiteside 1986), but I am inclined to eral phrynosomatines (Smith pers. obs.). Clearly, much view them as present in Clevosaurus hudsoni, in which more remains to be learned about these structures in the basisphenoid seems to be posteriorly expanded on lizards. the mid-line (Fraser 1988, fig. 14). Limited outgroup For the present, I do not parse this character in any comparison as well as the predominance of state 0 in detailed way. However, I regard Agaminae as showing the ingroup suggest this state is primitive. state 1 (a primary homology statement), even if the endolymphatic sacs enter the nuchal musculature by a 88. Dentition I. Etheridge and de Queiroz (1988) no. different route than in polychrotines. Assuming com- 19, Lang (1989) no. 39. Crowns of posterior cheek teeth plete phylogenetic ignorance of Iguania, there is no rea- (0) distinctly tapered, (1) approximately parallel-sided, son to exclude the theoretical possibility (admittedly or (2) distinctly flared. very unlikely in view of what we do in fact know) that the derived state could be a synapomorphy of the two This character is difficult to score in acrodont taxa, groups, with further transformation in agamines (devel- which generally superficially taper but have mesiodis- opment of an epiotic foramen, treated as a separate tally expanded tooth shafts. In Brookesia superciliaris, character below). tricuspid tooth crowns are generally not tapering (Smith pers. obs.). Etheridge and de Queiroz (1988) document 83. Hyoid I. Camp (1923), Hallermann (1994) no. 33. considerable variation in some taxa (e.g., Pristidactylus Epibranchial II (0) present, free, (1) connected to otic and Sceloporus). Coding by Etheridge and de Queiroz capsule or (2) absent. (1988) and Lang (1989) seems to be in conflict in Cory- tophaninae. While I agree with Lang (1989) that tooth Camp (1923) found the second epibranchial to be crowns tend to taper slightly in Laemanctus, the crowns absent in “most” iguanids, a result at variance with more of many teeth are indistinguishable from those that are recent studies. The “second épibranchial est libre” in parallel-sided, and compared with “sand lizards” they Chalarodon madagascariensis (Blanc 1965:110). do not taper substantially. Laemanctus longipes is given Polarization follows Hallermann (1994). state 1 (but details may be important more locally, that is, within Corytophaninae, Lang’s focus). Tooth crowns 84. Braincase I. Siebenrock (1895:1094), Moody (1980) in some Anolis give the impression of tapering, but this no. 11, Frost and Etheridge (1989) no. 15, Hallermann is because their bases are expanded (see below). (1994) no. 32. Epiotic foramen (0) absent or (1) present. The character is treated as ordered, but unpolar- Polarization follows Frost and Etheridge (1989). ized. 85. Braincase II. Etheridge (1959), Etheridge and de Queiroz (1988) no. 8, Lang (1989) no. 29, Frost and 89. Dentition II. Smith (2006a). Shafts of posterior Etheridge (1989) no. 13. Elevation of osseous labyrinth cheek teeth (0) roundish to mesiodistally compressed above opisthotics (0) low to moderate or (1) high. or (1) posterior cheek tooth shafts mesiodistally Polarization follows Etheridge and de Queiroz expanded. (1988). The shafts of the posterior cheek teeth in most iguanids are rounded or mesiodistally compressed, but 86. Braincase III. Conrad and Norell (2007) no. 95. in a few taxa (Anolis and Crotaphytus) they are Crista proötica (0) moderately to highly developed or expanded in this direction and are superficially more (1) strongly reduced. like the teeth of some teiids. Gambelia and occasional Eocene Lizards of the Clade Geiseltaliellus • Smith 289 phrynosomatines can approach state 1. In many acrodontans the crown and what exists of the shaft seem mesiodistally expanded; these species are scored 1. The state of this character cannot reasonably be evaluated in Sphenodon punctatus. State 0 is, however, treated as primitive because most scleroglossans and, seemingly, several fossil lepidosauromorphs retaining pleurodont teeth (Marmoretta oxoniensis, Gephy- rosaurus bridensis, Diphydontosaurus avonis and Hue- huequetzpalli mixtecus) show this state.

90. Dentition III. de Queiroz (1987) no. 46, Etheridge and de Queiroz (1988) no. 19, Frost and Etheridge (1989) no. 25, Wiens and Etheridge (2003) no. 48. Mid to posterior cheek teeth are (0) unicuspid, (1) tricuspid or (2) polycuspate. Smith (2006a) noted that tricuspid teeth in iguanians differ from those in most other squamates whose teeth have been described as “tricuspid” in that distinct grooves separating the central from accessory cusps are developed both lingually and labially. Tricuspid teeth of this description otherwise occur primarily in teiids and Figure 28. Tooth crowns in iguanians, from the mid- cannot be maintained as primitive for Squamata. dle of the right dentary, in lingual view. A, Basiliscus Etheridge and de Queiroz (1988) note that Phymaturus galeritus (UF 61491). B, Enyalioides oshaughnessyi shows state 2. Enyalioides oshaughnessyi seems to be (SMF 67590). Scale bar is 0.5 mm. intraspecifically variable, as my observations differ from those of Wiens and Etheridge (2003). Brookesia superciliaris is peculiar in that tricuspid teeth are developed anteriorly, whereas posterior teeth are unicuspid; it is scored 1. Polarization follows Etheridge and de Queiroz ever, as shown by the asymmetric but tricuspid teeth of (1988). The character is ordered. Enyalioides oshaughnessyi (Figure 28B). Sphenodon punctatus is not really comparable. 91. Dentition IV. Novum. Main cusp in crown (0) Teeth in scleroglossans can be asymmetric (e.g., some located above center of tooth or (1) shifted distally with gekkotans, lacertiforms and anguimorphs), which ren- respect to the tooth shaft, resulting in a distinctly asym- ders polarity equivocal. Yet all known early Eocene and metric crown. older iguanian fossils show state 0, which suggests that In most iguanians with multicuspid cheek teeth, the state 1 is derived. main cusp is located centrally, such that the mesial and distal halves of the tooth are (near) mirror images of one 92. Dentition V. Estes (1961), Smith (2006a, 2006b). another (Figure 28A); in species with strongly curved Tooth crowns (0) smooth or with only weak irregular- teeth (i.e., the longitudinal axis of the shaft is curved), ities or (1) with distinct striations. the main cusp is centrally located with respect to this Distinct striations are uncommon in Iguania, but axis of curvature (e.g., Gambelia wislizenii). In a few taxa Smith (2006a) reports several species, in particular Poly- the main cusp is shifted distally and the cuspiture is chrus (first noted by Estes 1961), in which these are pres- asymmetrical (Figure 28B). This is often apparent in the ent labially and lingually. maxillary teeth even when the dentary cheek teeth seem Striations have not, to my knowledge, been described symmetric. Most examined iguanines clearly show this in Rhynchocephalia. In Scleroglossa they are characteris- feature, as does Enyalioides oshaughnessyi. tic of certain clades (Scincoidea, Anguidae and some There is an ontogenetic component to tooth asym- Varanus), but there is no reason to consider them prim- metry in some taxa. In juvenile Ctenosaura similis, for itive for that clade. State 0 is taken to be primitive. instance, one or two maxillary teeth may have a slightly displaced main cusp, but most are symmetrical. The 93. Dentition VI. Cope (1864), Camp (1923), Estes et al. teeth are all tricuspid. It is only in larger individuals that (1988) no. 84, Frost and Etheridge (1989) no. 26. Tooth a fourth cusp appears mesially on the teeth and the teeth implantation (0) mostly pleurodont or (1) mostly acquire a distinct asymmetry; this ontogenetic change acrodont. cannot be ascribed solely to the posterior addition (de Diphydontosaurus avonis is scored 0 because many Queiroz 1987) of asymmetric teeth, because teeth in the cheek teeth have pleurodont implantation. See Estes same position in juvenile and adult differ in symmetry. et al. (1988) and Zaher and Rieppel (1999) for careful Teeth need not be polycuspate to be asymmetric, how- discussion. Polarization follows Estes et al. (1988). 290 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

(2) closed by suturing or fusion for Ն50% of length anterior to splenial. This character describes not the mode of restriction of the Meckelian groove, but rather how extensive restriction is. The character is treated as polarized (following Etheridge and de Queiroz 1988) and ordered. 96. Dentary III. Denton and O’Neill (1995) no. 23, Smith (2009). Intramandibular lamella (0) well developed for bracing of splenial and anteromedial process of coronoid, (1) absent or only weakly developed. Denton and O’Neill (1995) described in Teiidae what is here called the intramandibular lamella as a posterior extension of the intramandibular septum that depends from the dorsal roof of the intramandibular canal and braces the anteromedial process of the coronoid and the splenial laterally. Smith (2009) independently described a strong intramandibular lamella in certain Iguanidae, noting that the lamella in iguanids can be structurally independent of the intramandibular septum. A moderately to strongly developed lamella is Figure 29. Bones of the mandible in iguanians. A, found in Basiliscus (Figure 29C), Corytophanes, Lae- Right mandible of Brookesia superciliaris (TMM manctus, Crotaphytus, Gambelia, Oplurus cuvieri (other M8676) in medial view. B, Right dentary of Physig- species of the genus unknown), Chalarodon madagas- nathus cocincinus (TMM M8441) in medial view. C, cariensis, Enyalioides oshaughnessyi, some Anolis and Left dentary (reversed) of Basiliscus vittatus (YPM some Sauromalus. It is weak in Polychrus gutturosus, the R11132) in ventromedial view. Abbreviations: cn, coro- only disarticulated specimen of that genus available. noid; cnf, coronoid facet on dentary; d, dentary; iml, A strong lamella is absent in most outgroups, intramandibular lamella; iMl, infra-Meckelian lip; ims, including Sphenodon punctatus, Diphydontosaurus avo- intramandibular septum; Mgr, Meckelian groove; sMl, nis (Whiteside 1986, fig. 28b), Plestiodon fasciatus and supra-Meckelian lip, sp, splenial. Scale bars are 1 mm. Elgaria multicarinata. State 0 is consequently considered primitive. A moderately developed lamella, however, is found in Eublepharis macularius. 94. Dentary I. Etheridge (1959), Etheridge and de Queiroz (1988) no. 11–12, Lang (1989) no. 31, Frost and 97. Dentary IV. Novum. Anteromedial process of coro- Etheridge (1989) no. 20. Meckelian sulcus (0) open from noid and splenial (when present) articulate (0) medial to anterior end of splenial anterior to end of dentary, (1) supra-Meckelian lip of dentary for a distance of at least dorsal and ventral flange sutured but not fused, enclos- three teeth, or (1) almost entirely lateral to supra-Meck- ing a canal, or (2) dorsal and ventral flanges indistin- elian lip (i.e., internal to mandible). guishably fused, enclosing a canal. In many acrodontans (Agama, Laudakia, Draco, In Diphydontosaurus avonis the medial opening of Acanthosaura, Uromastyx, Leiolepis and Chamaeleonidae; the Meckelian canal is greatly restricted posteriorly Figure 29A), the anteromedial process of the coronoid (Whiteside 1986) and essentially closed (Whiteside articulates on the medial surface of the supra-Mecke- 1986, fig. 28b). Polarization follows Etheridge and de lian lip (see Bhullar and Smith 2008) of the dentary Queiroz (1988:338), who also implied that closure could below the tooth row for the space of at least two or three be seen as preceding fusion in phylogeny. The distribu- teeth. (Anteriorly, it may continue interior to the tion of closure and fusion in certain taxa seems consis- mandibular canal.) The splenial, when present, also tent with this notion (e.g., polymorphic in Chalarodon articulates on the medial to ventromedial surface of the madagascariensis: Frost and Etheridge 1989; interspecif- lip, although it too can continue anteriorly interior to ically variable in Basiliscus: Smith pers. obs.). Denser the mandibular canal. The combination of medial expo- taxon sampling (for instance, using fossil taxa described sure of the splenial and the anteromedial process of the in Smith 2006b) may be necessary for ordering of this coronoid, and the development of only medial articula- character to produce meaningful results. tion surfaces for these bones on the supra-Meckelian lip, is state 0. In some agamids (Physignathus, Hydrosaurus and 95. Dentary II. Etheridge and de Queiroz (1988) no. some Australian agamids, including Pogona) the artic- 11–12, Lang (1989) no. 31, Frost and Etheridge (1989) ulation for the anteromedial process of the coronoid no. 20. Meckelian sulcus (0) open, (1) closed by sutur- is almost exclusively on the lateral (internal) surface of ing or fusion for <50% of length anterior to splenial or the supra-Meckelian lip (Figure 29B). In these species, Eocene Lizards of the Clade Geiseltaliellus • Smith 291 the anteromedial process of the coronoid and the sple- not be calculated because the Meckelian groove is fused nial fit in a slot developed between the supra-Mecke- so far posteriorly. However, the intramandibular sep- lian lip and an internal mandibular lamella. The tum cannot be more than 75% of the length of the tooth lamella braces them laterally and the lip braces them row, or it would be visible when an isolated dentary is medially. viewed end-on. Its relative length could be estimated in Most iguanids lack the highly developed intra- Microlophus occipitalis at approximately 0.5. These mandibular lamella seen, for example, in Physignathus, species are scored 0. but in other respects of the coronoid–splenial–dentary In Sphenodon punctatus, the intramandibular sep- articulation they are similar. In all iguanids, a well-devel- tum is longer than the tooth row and so receives 1. It is oped supra-Meckelian lip extends far posteriorly, similarly long in Diphydontosaurus avonis. State 0 is approaching or reaching the posterior end of the tooth found in many scincomorphs (except part of Amphis- row. Here the anteromedial process of the coronoid baenia; Smith 2006b), but state 1 is common to sometimes articulates on the external surface of the lip Anguimorpha. This character is treated as unpolarized. for a short distance (usually less than the space of two teeth) before rotating and articulating on its internal 99. Dentary VII. Etheridge and de Queiroz (1988), surface. The articulation is entirely internal in all exam- Frost et al. (2001) no. 63. Ratio of dentary length to ined members of the Tropidurus group, Phrynosoma, mandibular length, from anterior tip to anterior end of Phymaturus, Sauromalus, Dipsosaurus and Enyalioides glenoid fossa is (0) Յ0.30, (1) 0.31 to 0.50, or (2) >0.50. oshaughnessyi, in almost all Anolis, and in most Leio- As Etheridge and de Queiroz (1988) noted, the den- cephalus. It articulates externally for the space of about tary is elongate in some iguanids, including Anolis and one tooth or less in Sator, some Sceloporus (e.g., S. for- Oplurus, as well as many tropidurines. It is elongate as mosus and S. serrifer), Enyaliosaurus, Ctenosaura, well in Chamaeleonidae and in Leiolepis, Uromastyx Morunosaurus groi, Laemanctus, some Basiliscus (Fig- and many other agamids, but notably shorter in ure 29C), Polychrus, Pristidactylus torquatus, Leiosaurus Priscagama gobiensis. bellii and a few Anolis (A. ricordi, A. garmani and A. whitemani). More uncommonly, the anteromedial The dentary is highly elongate in Sphenodon punc- process of the coronoid articulates externally for a space tatus (Smith pers. obs.), Diphydontosaurus avonis of about two teeth; taxa in this class are Basiliscus (Whiteside 1986, figs. 28 and 31), and other Late Juras- plumifrons, Stenocercus scapularis, Oplurus cuvieri, sic rhynchocephalians (Cocude-Michele 1963). It is some Sceloporus (e.g., S. cyanogenys), Petrosaurus tha- shorter (state 0), however, in Huehuequetzpalli mixte- lassinus, Hoplocercus spinosus, Corytophanes, many cus (Reynoso 1998) and scleroglossan outgroups, and I Pristidactylus (P. achalensis, P. casuhatiensis and P. vol- take the shorter dentary (state 0) to be plesiomorphic. canensis), Aperopristis and Diplolaemus bibronii. The The character is ordered. splenial only ever articulates on the internal surface of the supra-Meckelian lip. State 1 refers to the primarily 100. Coronoid I. Etheridge (1966), Moody (1980) no. internal articulation of the coronoid on the supra-Meck- 59, Lang (1989) no. 33, Frost and Etheridge (1989) no. elian lip (lateral exposure for about one tooth space or 16,18. (0) Dentary weakly overlaps coronoid or vice less). Notably, Priscagama gobiensis also seems to show versa, anterolateral process of coronoid weak or absent; this state. (1) coronoid expanded at least weakly ventrolaterally State 0 is found in Sphenodon punctatus and other with strong anterolateral process overlapping dentary. rhynchocephalians, as with many gekkotans, but in the Lack of polarization follows Frost and Etheridge eublepharid Coleonyx external articulation of the coro- (1989). noid on the supra-Meckelina lip is restricted, and anteriorly the coronoid and splenial articulate primarily 101. Coronoid II. Lang (1989). Coronoid (0) shows no ventrally. In Autarchoglossa, the anteromedial process strong posterolateral process or (1) has strong postero- of the coronoid articulates externally for at least a space lateral process that extends laterally unto the posterior of two teeth; more anteriorly, it and the splenial gener- margin of the anterior surangular foramen. ally articulate on the ventromedial or ventral surface of Only a few ingroup taxa (Anolis ricordi, Geiseltaliel- the supra-Meckelian lip. Most autarchoglossans would lus) show state 1. A partly developed posterolateral also therefore receive state 0. (Exceptions include many process is seen, however, in Laemanctus and Coryto- species of Celestus, in which the coronoid and splenial phanes (Lang 1989). Acrodontans generally show a pos- articulations are internal on the supra-Meckelian lip.) terolateral process along the top of the surangular. The distribution of state 0 outside Iguania suggests that Because it is dorsomedial to the anterior surangular state 1 is derived in Iguania. foramen, it seems on positional grounds to be homologous with a weak posterior elaboration found on the 98. Dentary V. Smith (2006a). (0) Intramandibular sep- descending posteromedial process of the coronoid in tum Յ75% as long as tooth row; (1) intramandibular many iguanids. The acrodontan condition is not incor- septum >75% as long as tooth row. porated in the phylogenetic analysis. In Leiocephalus and Tropidurus torquatus, the exact State 0 is found in Sphenodon punctatus and most proportional length of the intramandibular septum can- scleroglossans and is deemed ancestral. 292 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

102. Surangular I. Frost and Etheridge (1989) no. 19. Polarization follows Estes et al. (1988). Acquisition Anterior surangular foramen (0) near apex of surangular, of state 1 is a synapomorphy of Iguania. It is reduced in dorsal to the dentary (when elongate), or (1) foramen dis- certain iguanians (e.g., Uromastyx) and lost in placed ventrally and bordered dorsally by posterior Chamaeleonidae (Estes et al. 1988). extremity of dentary when the latter is elongate. In most iguanids, the anterior surangular foramen 108. Hyoid II. McDowell (1972), Lang (1989) no. 37. is close to the posterior base of the coronoid, near the Ceratobranchial II is (0) shorter than ceratobranchial I, apex of the surangular (state 0). State 1 is confined to or (1) longer than it. Acrodonta and Hoplocercinae among ingroup taxa. States 0 and 1 are described as the “Z” and “X” pat- The foramen is notably ventrally displaced in certain terns, respectively, of Lang (1989). Iguaninae as well (de Queiroz 1987). Polarization follows Lang (1989). Polarization follows Frost and Etheridge (1989). 109. Ribs I. Etheridge and de Queiroz (1988) no. 28, 103. Angular I. Etheridge and de Queiroz (1988) no. Frost and Etheridge (1989) no. 38. First cervical rib 15, McGuire (1996) no. 23. In adult, anterior process of located on (0) the third, (1) the fourth or (2) the fifth angular (0) does not extend far anteriorly, never reaches cervical vertebra. antepenultimate dentary tooth, rarely beyond ultimate Following Frost and Etheridge (1989), Brookesia tooth, or (1) extends at least as far forward as antepenul- superciliaris is scored ?. I find state 2 to be present in timate dentary tooth. Oplurus, Chalarodon (cf. Frost and Etheridge 1989), Iguana iguana seems to depart from other igua- Iguaninae, Corytophaninae, Anolis and Polychrus acu- nines in showing a strong ontogenetic transformation in tirostris, but not in Pristidactylus torquatus (Frost and the length of the angular, which is almost entirely Etheridge noted that it varies in Pristidactylus). The first behind the transverse level of the posterior end of the rib is on the fifth cervical in Geiseltaliellus maarius (con- tooth row in juveniles, but extends well anterior of this tra Rossmann 2000). According to Moody (1980), most level in large adults. agamids have state 1, but I follow Frost and Etheridge This character is treated as unpolarized. (1989) in coding included agamids 2. Sphenodon punctatus shows state 1. This character 104. Splenial I. Frost and Etheridge (1989) no. 22. Pos- is not polarized (following Frost and Etheridge 1989), teriorly, splenial terminates (0) anterior to anterior edge but it is ordered. of mandibular fossa or (1) at or posterior to this level. Scoring from Frost and Etheridge (1989), except 110. Vertebrae I. Siebenrock (1893), Hoffstetter and for Enyalioides oshaughnessyi and outgroup scleroglos- Gasc (1969), Frost and Etheridge (1989). Number of sans. This character cannot be scored in species lacking cervical vertebrae is (0) eight or (1) five. a splenial. A reduction in the number of cervical ribs, among iguanians, is an autapomorphy of Chamaeleonidae 105. Splenial II. Smith (2006a), Conrad and Norell (Frost and Etheridge 1989). (2007) no. 117. Anterior inferior alveolar foramen of mandible located (0) within splenial or (1) at boundary 111. Vertebrae II. de Queiroz (1987) no. 56, McGuire between splenial and dentary. (1996) no. 38. Accessory vertebral articulations (0) Outgroup comparison is ambiguous (state 0 is absent or only weakly developed, facing dorsolaterally, common to many scleroglossans, state 1 is common to (1) distinct, ventrolaterally facing accessory articulation Rhynchocephalia), and the character is treated as unpo- surface developed, but zygosphene connected by con- larized. tinuous arc of bone to prezygapophysis on anterior presacral vertebrae, or (2) distinct accessory articulation surfaces, zygosphene separated by deep notch from 106. Angular II. Frost and Etheridge (1989) no. 24, prezygapophysis on anterior presacral vertebrae. McGuire (1996) no. 24. Posterior mylohyoid foramen located (0) anterior to or approximately at level of apex Weak accessory articular surfaces that face dorso- of coronoid or (1) posterior to level of superior apex of laterally could be ancestral for Squamata (Gauthier et al. coronoid. 1988). When these become better developed (state 1), they face laterally to ventrolaterally; the most highly I find state 0 to be present in Crotaphytus collaris elaborated accessory articulations (state 2) occur in (cf. Frost and Etheridge 1989). The foramen is far ante- Iguaninae, Corytophaninae, Polychrus and possibly rior to the level of the apex of the coronoid in many Hoplocercinae, in which the zygosphene is separated agamids, and this variation ought eventually to be incor- from the prezygapophysis by a notch (de Queiroz 1987). porated. The peculiar accessory intervertebral connections in Polarization follows Frost and Etheridge (1989). Brookesia superciliaris are not homologous (Siebenrock 1893). 107. Articular I. Estes et al. (1988) no. 80. Larger angu- Zygosphene development can differ among regions lar process on articular (0) absent or (1) present. of the vertebral column. In Polychrus gutturosus, for Eocene Lizards of the Clade Geiseltaliellus • Smith 293 instance, anterior presacral vertebrae tend to show fully (Camp 1923; Cocude-Michel 1963; Evans 1981) as well developed zygosphenes, but on posterior presacrals the as Sphenodon punctatus. Curiously, Schauinsland zygosphenes are connected by a continuous sheet of (1900) stated that the autotomic septum in the caudal bone with the prezygapophyses. In other species (e.g., vertebrae of Sphenodon punctatus did not arise by Oplurus cuvieri), posterior vertebrae show better devel- resorption of bone, as in Squamata (Etheridge 1967), opment of accessory articulations than anterior ones. but rather represents a primary failure of the two halves State 0 is found in Sphenodon punctatus and most of the vertebra to unite across the sclerotomal boundary. scleroglossan clades (including the outgroups selected For the present, I continue to consider the septa in here) and is deemed ancestral. The character is treated Sphenodon as homologous. as ordered. 116. Vertebrae V. cf. Etheridge (1967), Etheridge and 112. Vertebrae III. de Queiroz (1987), Etheridge and de Queiroz (1988) no. 33, Frost and Etheridge (1989) de Queiroz (1988), Etheridge (1995). Posterior presacral no. 34, Poe (2004) no. 49. Autotomic septum (0) passes neural spines (0) very short, (1) moderate or (2) highly posterior to transverse process (“Sceloporus type,” in elongate. part), (1) bisects transverse process, at least anteriorly, Neural spines are extremely short in a few iguani- which may be split into laterally diverging splines ans (e.g., Sauromalus, Phymaturus: de Queiroz 1987; (“Iguana type,” in part), or (2) passes anterior to trans- Etheridge 1995; Smith pers. obs.). Although they are also verse process (“Anolis type,” in part). reduced in certain other iguanians (e.g., Uromastyx and Etheridge (1967) recognized in Iguania several Petrosaurus), reduction is not as great as in the afore- recurring patterns in the morphology of the caudal ver- mentioned clades. Neural spines are also greatly elon- tebrae. These patterns were taken up as character states gate in certain Basiliscus (Lang 1989; see above). They by Etheridge and de Queiroz (1988) and Frost and are also elongate in the tail in certain Anolis (e.g., A. Etheridge (1989). Like Frost et al. (2001), I break these cristatellus). types into constituent characters. Cases in which the Outgroup comparison suggests state 1 is primitive. autotomic plane is absent, and no known variants or This character is treated as ordered. ontogenetic data might permit character state discrim- ination, species are scored ?. Variants of Basiliscus will 113. Ribs II. Hoffstetter and Gasc (1969), Etheridge show an autotomic plane posterior to the transverse (1964), Lang (1989) no. 45. Number of sternal ribs is (0) processes on the ultimate vertebra bearing them four, (1) three or (2) two. (Etheridge and de Queiroz 1988; Smith pers. obs. in B. Some variation in this feature has been discussed basiliscus). In species of Stenocercus (Ophryoessoides) by Etheridge (1964) and Lang (1989). other than scapularis, the septum passes posterior to the processes (Etheridge 1967). In Iguaninae, the anterior Outgroup comparison suggests state 1 is primitive. transverse processes are bisected by the autotomic sep- The character is treated as ordered. tum and are divergent; more posteriorly, the septum passes posterior to a single pair of transverse processes, 114. Ribs III. Etheridge and de Queiroz (1988) no. 31, consistent with the derivation of this state from the Sceo- Lang (1989) no. 41. (0) All post-sternal presacral verte- porus type (Etheridge 1967). brae bearing ribs; one or more posterior presacral vertebrae (1) with ribs absent or (2) with ribs fused. Polarization follows Etheridge (1967). In Plestiodon fasciatus the autotomic septum bisects the transverse Many agamids (including Physignathus cocincinus process, although the two halves of the latter do not and P. lesueurii) show state 2. Uromastyx and Leiolepis become divergent (Etheridge 1967); it is scored 1. In show state 0. Sphenodon punctatus the septum passes anterior to the Polarization follows Etheridge and de Queiroz main axis of the process (Hoffstetter and Gasc 1969). (1988). The character is not considered ordered.

115. Vertebrae IV. Etheridge (1959, 1967), de Queiroz 117. Vertebrae VI. Etheridge (1967), Etheridge and de (1987) no. 60, Etheridge and de Queiroz (1988) no. 34, Queiroz (1988) no. 33, Frost and Etheridge (1989) no. Lang (1989) no. 42, Frost and Etheridge (1989) no. 41, 34, Poe (2004) no. 49. Transverse processes (0) devel- McGuire (1996) no. 40, Poe (2004) no. 54. Caudal auto- oped on most caudal vertebrae or (1) absent from a sig- tomy septa (0) present or (1) absent in adult, even ante- nificant proportion of caudal vertebrae (two-thirds or riorly. more). Autotomic planes are absent in Stenocercus scapu- Tranverse processes are absent on caudal vertebrae laris, but present in most other Stenocercus (Etheridge in Stenocercus scapularis from roughly the middle of the 1966). They are apparently absent in Geiseltaliellus, tail posteriorly. Most iguanids retain transverse including juvenile specimens (Capitolacerta dubia) processes on the caudal vertebrae (Etheridge 1967). referred to G. longicaudus (Smith pers. obs.). Scoring for Hoplocercinae follows Wiens and Etheridge (2003). 118. Ribs IV. Etheridge (1959, 1965), de Queiroz Polarization follows Gauthier et al. (1988). Auto- (1987) no. 63, Etheridge and de Queiroz (1988) no. 32, tomic septa are known in Mesozoic rhynchocephalians Frost and Etheridge (1989) no. 40. Post-xiphisternal 294 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009 inscriptional ribs (0) all attached proximally to dorsal In the ingroup, a moderately to well-developed ante- ribs, none confluent midventrally, or (1) one or more rior process of the interclavicle has been found in Leiolepis pairs attach dorsally and are confluent mid-ventrally, (2) and Uromastyx (Moody 1980) and in Leiocephalus none is attached to dorsal ribs or continuous midven- (Etheridge and de Queiroz 1988; Pregill 1992). I also find trally, present as isolated splints in body musculature. a moderately developed anterior process in Stenocercus Etheridge (1965) indicates that at least one pair of (Ophryoessoides) scapularis (cf. Etheridge 1966). inscriptional ribs forms a midventral chevron in Iguana, Lack of polarization follows Frost and Etheridge Brachylophus and some Conolophus, Amblyrhynchus, (1989). Cyclura and Ctenosaura; Sauromalus shows state 0. Hoplocercus spinosus and at least two species of 123. Interclavicle II. Etheridge (1959), de Queiroz Enyalioides (E. laticeps and E. praestabilis) show state 1 (1987) no. 69. Interclavicle (0) T-shaped or (1) arrow- (Etheridge 1965). It is possible that state 1 is primitive shaped. for Hoplocercinae, but Morunasaurus groi shows state Outgroup comparison suggests state 0 is primitive. 0; more data are desirable. Polarization follows Frost and Etheridge (1989). 124. Interclavicle III. Moody (1980) no. 100, de The character is not considered ordered. Queiroz (1987) no. 68, Etheridge and de Queiroz (1988) no. 24. Posterior process is (0) long, extends posteriorly 119. Clavicle I. Moody (1980) no. 101, de Queiroz beyond the lateral corners of the sternum, or (1) dis- (1987) no. 67, Etheridge and de Queiroz (1988) no. 21 tinctly shorter than this. and 22, Lang (1989) no. 48, Frost and Etheridge (1989) no. 30. Ventromedial end of bone is (0) slender, rod- I find that the posterior process is distinctly shorter like or tabular, or (1) flat, wide and generally fenes- in Polychrus acutirostris and P. marmoratus and have trated. scored the former 1 (cf. Etheridge and de Queiroz 1988). A hole develops in some taxa with expanded por- Polarization follows Etheridge and de Queiroz tions of the clavicle, perhaps related to the action of M. (1988). clavodeltoideus (Camp 1923; Russell 1988). If more information on the development and functional 125. Scapulocoracoid I. Lécuru (1968a), de Queiroz anatomy of this region in lizards generally were known, (1987) no. 65, Etheridge and de Queiroz (1988) no. 25, it might be desirable to rewrite this character for the Frost and Etheridge (1989) no. 35. Scapular fenestra is muscular basis. Conrad and Norell (2007: 24) stated that (0) absent or (1) present. the medial portion of the clavicle was “broad” in There are four fenestrae in the squamate scapuloco- Saichangurvel davidsoni, but on the basis of the figures racoid (Camp 1923; Romer 1956). From posteroventral it is unclear exactly how broad; I code that species as to anterodorsal these are the posterior and anterior unknown. coracoid fenestrae, the scapulocoracoid fenestra and the Moody (1980) indicates that 0 is the state most scapular fenestra. Each fenestra does not only emar- commonly present in Agaminae, but Physignathus, ginate the bone but also the epicoracoid cartilage. These some Hypsilurus and Hydrosaurus show state 2, as does fenestrae could be related to the action of particular Leiolepis. muscles and Romer (1956) suggested that the first three Lack of polarization follows Frost and Etheridge fenestrae correspond to the origins of the biceps, scapu- (1989). lohumeralis anterior and supracoracoideus (see also Russell 1988). These characters could be considered 120. Clavicle II. Etheridge and de Queiroz (1988) no. more than simply binary, as the definitions of states 23. Lateral margin is (0) smoothly rounded or (1) drawn might imply, and if we knew more about the structure out into an angular flange or a hook. and function of the pectoral muscles it might be desirable to rewrite this character on a muscular basis. It is Polarization follows Etheridge and de Queiroz not obvious that “thinning” of bone in the position of a (1988). posterior coracoid fenestra implies that the presence of a fenestra is primitive for Iguania (cf. Frost and 121. Clavicle III. Estes et al. (1988) no. 117, Lang (1989) Etheridge 1989). Only the anterior coracoid and scapu- no. 47, Frost and Etheridge (1989) no. 31. Clavicle locoracoid fenestrae are found in all iguanians. inserts predominantly (0) on suprascapula or (1) on Oplurus quadrimaculatus has a scapular fenestra. scapula. Notably, however, a phylogeny of Oplurinae (Titus and In most examined agamines (including Physig- Frost 1996) found this species to be nested within Oplu- nathus, excepting Hydrosaurus amboinensis) the clavi- rus, not basal, and other examined species show state 0. cle inserts on the scapula. A scapular fenestra is present in Hydrosaurus Polarization follows Frost and Etheridge (1989). amboinensis, but absent in other agamids I have seen. A scapular fenestra is very weakly present in Pristidacty- 122. Interclavicle I. Camp (1923), Moody (1980) no. lus torquatus. 100, Frost and Etheridge (1989) no. 32. Anterior process Polarization follows Etheridge and de Queiroz (0) absent or poorly developed, or (1) well developed. (1988). Eocene Lizards of the Clade Geiseltaliellus • Smith 295

126. Scapulocoracoid II. Lécuru (1968a), de Queiroz (1987) no. 66, Etheridge and de Queiroz (1988) no. 26, Frost and Etheridge (1989) no. 36, McGuire (1996) no. 43. Posterior coracoid fenestra is (0) absent or (1) present. I tentatively code Saichangurvel davidsoni as having scapulocoracoid and anterior coracoid fenestrae, rather than two coracoid fenestrae (cf. Conrad and Norell 2007). Polarization follows Etheridge and de Queiroz (1988).

127. Sternum I. Moody (1980) no. 98, de Queiroz Figure 30. Sterna in iguanians in ventral view. A, (1987) no. 70, Estes et al. (1988) no. 121, Etheridge and Pristidactylus torquatus (CAS 85234). B, Sceloporus de Queiroz (1988) no. 27, Frost and Etheridge (1989) occidentalis. Sternal cartilages are in gray, interclavicle no. 37. Sternum posterior to interclavicle (0) is imperfo- in black. Abbreviations: cc, coracoid; cl, clavicle; f1, rate or has (1) a large, equidimensional (roundish to car- large sternal fontanelle; f2, narrow sternal fontanelle; diac-shaped) and median fontanelle, or (2) a large dual ic, interclavicle; sc, scapula; stn, sternum. Scale bars are fontanelle, separated on the median by a strut of carti- 3 mm. lage. Large fontanelles in the cartilaginous sternum are thought to arise through resorption during ontogeny 1980), but fontanelles of any kind are absent in (see Bogoljubsky 1914 on Lacerta), although Etheridge Chamaeleonidae. If the present character could be (1962) found no ontogenetic variation in size in his sam- rewritten using a muscular basis (cf. Camp 1923), it ples of larger individuals of Sator grandaevus. A large could clarify the evolution of this feature in Acrodonta. sternal fontanelle has been considered a possible synapomorphy of Phrynosomatinae and Tropidurinae Polarization follows Etheridge and de Queiroz (Frost and Etheridge 1989), although a long, narrow (1988). The character is unordered. fontanelle that is largely hidden by the posterior process of the interclavicle has been reported in other taxa (e.g., 128. Sternum II. Novum. Narrow sternal fontanelle Weiner and Smith 1965; de Queiroz 1987). Although it behind interclavicle (0) absent or (1) present. is tempting to homologize this fontanelle with the larger, This narrow fontanelle (see also above) is usually generally more posteriorly placed fontanelle of phryno- largely concealed behind the posterior process of the somatines and tropidurines, there are specimens of interclavicle, but can extend slightly beyond it posteri- Sceloporus (e.g., S. malachiticus, YPM R11950; S. occi- orly (Figure 30B). It might represent primary failure of dentalis, TMM M8940: Figure 30A; see also Etheridge the two halves of the sternum to fuse completely dur- 1964) in which both a large, posterior, median ing ontogeny (cf. Bogoljubsky 1914). In some species fontanelle and a narrow one behind the interclavicle are (e.g., Leiocephalus melanochlorus; see also Etheridge present. Thus, the hypothesis of primary homology 1964), a large sternal fontanelle has a keyhole shape that would seem to fail the conjunction test (de Pinna 1991; suggests the presence of both a large sternal fontanelle Patterson 1982). Moreover, the median posteriorly (previous character) and a narrow one. The sternal directed prong of a cardiac fontanelle in some taxa (Fig- fontanelle cited by Weiner and Smith (1965) in Gam- ure 30A) and the median dividing bar of the dual belia is considered to represent this narrow one on the fontanelle indicate that in these cases, cartilage tends to basis of shape and position. be retained on the mid-line rather than being absent. The absence of a narrrow sternal fontanelle in out- For these reasons I do not consider the two kinds of groups suggests that state 0 is primitive. The mode of fontanelle to be homologous. appearance of a large sternal fontanelle (by resorption) The occurrence of a sternal fontanelle in Oplurus is does not necessarily apply to this narrow fontanelle. interspecifically variable (e.g., absent in O. cyclurus, pres- There are even cases in which the narrow sternal ent but relatively small in O. quadrimaculatus, O. gran- fontanelle is discontinuous (e.g., one Anolis ricordi, UF didieri, O. fierinensis, O. cuvieri) (cf. Frost and Etheridge 64820). This fontanelle is possibly primary. 1989). Round and found at the tip of the interclavicle, it is tentatively considered homologous with the large sternal fontanelle of phrynosomatines and tropidurines. No 129. Pes I. Frost et al. (2001) no. 39. Digit IV of pes (0) fontanelle was discovered in Chalarodon madagas- longer than digit III or (1) subequal in size to it. cariensis. Most Uromastyx (e.g., U. aegyptia) show state State 1 is considered derived within Iguania and 2 (Moody 1980), which is probably primitive for the constitutes an autapomorphy of Polychrus (Frost et al. clade. These fontanelles differ in U. acanthinurus, com- 2001). paring my observations with those of Moody (1980). Outgroup comparison is ambiguous for the prim- State 2 is otherwise common to Agamidae* (Moody itive state leading up to Iguania. 296 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

130. Scalation I. Smith (1946), Frost and Etheridge The well-preserved squamation in Geiseltaliellus (1989) no. 42, Etheridge and de Queiroz (1988) no. 45. maarius indicates that it shows state 0. Interparietal scale (0) narrow or (1) large, at least as wide Outgroup comparison suggests state 1 is derived. as the interorbital space. Frost and Etheridge (1989) note the “frequently 137. Scalation VIII. Etheridge and de Queiroz (1988), obvious” nature of edge-to-edge scale amalgamation in Wiens and Etheridge (2003) no. 24. Tail (0) not spin- some Tropidurus and Uranoscodon. It is also apparent ous or (1) with strongly projecting spines. in some Stenocercus and some phrynosomatines (Smith Some iguanines, Uromastyx, Oplurus and hoplo- pers. obs.) and need not be taken as evidence against the cercines have a very spiny tail. It may be useful in the homology of these structures. If large cephalic scales in future to quantify this character for more precision. For Iguania are indeed constructed by the amalgamation of the present, the boundary between the two states is indi- many smaller scales, edge structures could persist in cated implicitly by the coding in the matrix. some species or individual organisms. Outgroup comparison suggests state 1 is derived. The interparietal is small in Sphenodon punctatus and Eublepharis macularius. It is larger in Elgaria multicarinata and Plestiodon faciatum, but still narrower 138. Scalation IX. Etheridge and de Queiroz (1988) no. than the interorbital space. Polarization follows 43, Frost and Etheridge (1989) no. 47. Gular fold (0) Etheridge and de Queiroz (1988). complete medially, with or without distinct change in squamation, or (1) incomplete medially or absent. 131. Scalation II. Frost and Etheridge (1989) no. 43. This character is not meant as an exhaustive char- Interparietal black spot (0) absent or (1) present. acterization of variation in the gular fold. For instance, The median black spot in Microlophus occipitalis is Frost and Etheridge (1989) noted that a change in squa- located well behind the interparietal scale. mation (the occurrence of much smaller scales) may or may not accompany the fold. I am inclined to treat the Polarization follows Frost and Etheridge (1988). change in squamation as a separate character, although I have not included it here. 132. Scalation III. Williams (1988), Frost et al. (2001) no. 2. Mental scale (0) single or (1) divided. Polarization follows Etheridge and de Queiroz (1988). State 1 is unique to Anolis and Polychrus.

133. Scalation IV. Etheridge and de Queiroz (1988) no. 139. Scalation X. Camp (1923), Jullien and Renous- 46, Frost and Etheridge (1989) no. 44. Superciliary Lécuru (1972), Etheridge and de Queiroz (1988) no. 41, scales (0) not distinctly elongate and imbricate or Frost and Etheridge (1989) no. 48. Femoral pores (0) (1) distinctly so. present or (1) absent. Unpolarized. Polarization follows Etheridge and de Queiroz (1988). 134. Scalation V. Etheridge and de Queiroz (1988) no. 47, Frost and Etheridge (1989) no. 45. Subocular scales 140. Scalation XI. Jullien and Renous-Lécuru (1972), are (0) all subequal in size or (1) at least one scale below Etheridge and de Queiroz (1988) no. 42, Frost and eye conspicuously enlarged. Etheridge (1989) no. 49. Preanal pores (0) absent or (1) Unpolarized. present. Polarization follows Etheridge and de Queiroz 135. Scalation VI. Estes et al. (1988), Etheridge and de (1988). Queiroz (1988) no. 44, Frost and Etheridge (1989) no. 46. Projecting mid-dorsal scale row (0) present or (1) absent. 141. Scalation XII. Peterson and Williams (1981), Pregill (1992) noted that Leiocephalus pratensis Moody (1980) no. 117, Etheridge and de Queiroz (1988) lacks a projecting mid-dorsal scale row, but according to no. 39, Frost and Etheridge (1989) no. 51. Subdigital his phylogeny this is not the primitive condition for the scale surface (0) strongly carinate or (1) smooth, or only genus. State 1 includes two different conditions, one in very weakly carinate. which no mid-dorsal scale row is present at all and I have tentatively coded Geiseltaliellus 1 on the basis another in which a mid-dorsal scale row is present but of the specimens presented here and of the holotype of not projecting. It may be fruitful to elaborate, drawing G. longicaudus. a distinction between these two conditions, or to differ- Polarization follows Frost and Etheridge (1989). entiate further state 0. Unpolarized. 142. Scalation XIII. Smith (1946), Laerm (1973). Sub- digital lamellae (0) without strong ventral–postaxial 136. Scalation VII. Wiens and Etheridge (2003) no. 16 flaps or (1) with these structures. and 17. Scales of dorsum (0) uniform in size or (1) het- Well-developed subdigital flaps are unique to erogeneous, with a few large scales interspersed among Basiliscus. granular ones. Outgroup comparison suggests state 1 is derived. Eocene Lizards of the Clade Geiseltaliellus • Smith 297

143. Scalation XIV. Moody (1980) no. 122, Peterson 152. Body proportions II. Novum. Leg length (0) less (1983), Williams (1988), Etheridge and de Queiroz than two-thirds snout-vent length or (1) greater than (1988) no. 36, Frost and Etheridge (1989) no. 52. Spin- two-thirds snout-vent length. ules on scale organs (0) absent or (1) present. This feature was discussed in the comparative sec- Spinules are notably known from several iguanians, tion of the text. For want of adequate data on the even when they are lacking on the scale organs (e.g., ontogeny of relative leg length for most species, I used Cole and van Devender 1976; Peterson 1984). Polariza- species averages to score this character. tion follows Etheridge and de Queiroz (1988). State 0 obtains in all outgroup species measured and is consequently considered primitive. 144. Limb innervation I. Lécuru (1968b), Frost and Etheridge (1989) no. 58. Ulnar nerve pathway shows (0) L-condition (superficial to musculature) or (1) V-con- Appendix 4: dition (deep to it). Unambiguous Apomorphies Unpolarized. In support of the single most-parsimonious cladogram 145. Limb innervation II. Jullien and Renous-Lécuru of iguanian relationships nodes (see Figure 16), together (1972), Frost and Etheridge (1989) no. 59. Dorsal shank with unambiguous autapomorphies of terminal species. muscle innervation shows (0) A-condition (peroneus), For character optimization, the tree was rooted between (1) B-condition (interosseus). Rhynchocephalia (Sphenodon punctatus + Diphydon- tosaurus avonis) and Squamata, which only affects the Unpolarized. interpretation of autapomorphies at Node 1. 146. Hemipenis I. Frost and Etheridge (1989) no. 60. → → Posterior lobe (0) not enlarged or (1) enlarged. Node 1 (Saichangurvel + Iguania): 19(0 1), 22(0 1), 43(0→1), 44(0→1), 50(0→1), 90(0→1), 97(0→1), I follow Frost and Etheridge (1989) in not polariz- 107(0→1) ing this character. Saichangurvel davidsoni: No autapomorphies. 147. Hemipenis II. Frost and Etheridge (1989) no. 61. Node 2 (Iguania): 1(0→1/2), 31(0→1), 33(0→1), (0) Unicapitate or weakly bilobate without distinctly 35(0→1), 50(1→2) divided sulci; (1) bilobate with distinctly dividied sulci; or (2) strongly bicapitate. Node 3 (Priscagama + Acrodonta): 1(1/2 →3), 8(0→1), Unpolarized. This is considered an ordered trans- 14(0→1), 38(0→1), 45(0→1), 89(0→1), 93(0→1) formation. Priscagama gobiensis: 22(1→0), 47(0→1), 53(0→1), 90(1→0) 148. Hemipenis III. Arnold (1984) no. 32, Frost and Etheridge (1989) no. 62. M. retractor lateralis posterior Node 4 (Acrodonta): 34(0→2), 66(0→1), 97(1→0), (0) not completely divided or (1) completely divided. 99(0→1), 102(0→1) Unpolarized. Leiolepis belliana: 4(0→1), 48(0→1), 49(0→1), 58(0→1), 63(0→1), 87(0→1), 122(0→1), 148(0→1) . 149. Hemipenis IV Arnold (1984) no. 35, Frost and → → Etheridge (1989) no. 63. M. retractor lateralis posterior Node 5 (Brookesia + Agamidae): 19(0 1), 26(0 1) Brookesia superciliaris: 5(0→1), 9(0→1), 23(0→1), (0) not substantially situated within the hemipenial sheath → → → → → or (1) substantially situated within the hemipenial sheath. 25(0 1), 29(0 1), 31(1 2), 38(1 0), 39(1 2), 47(0→1), 50(2→3), 53(0→1), 81(0→1), 107(1→0), Unpolarized. 110(0→1), 113(1→2), 118(0→1), 129(1→0), 138(0→1), 139(0→1), 144(0→1) 150. Alimentary canal I. Lönnberg (1902), Iverson (1980), Frost and Etheridge (1989) no. 65. Colic septa Node 6 (Physignathus + Agama): 20(0→1), 22(1→0), (0) absent or (1) present. 39(1→0), 84(0→1), 90(1→0), 117(0→1) Polarization follows Frost and Etheridge (1989). Physignathus cocincinus: 16(1→0), 62(0→1), 73(1→0), 79(1→0), 97(0→1), 98(1→0), 106(0→1), 112(1→2) 151. Body proportions I. Novum. Tail length (0) less Agama agama: 4(0→1), 49(0→1), 72(1→0), 79(1→2), than twice snout-vent length or (1) greater than twice 80(0→2), 124(0→1) snout-vent length. Node 7 (Iguanidae): 10(0→1), 74(0→1), 76(0→1), This feature was discussed in the comparative sec- → → → → tion of the text. For want of adequate data on the 79(1 2), 80(0 1), 95(0 1), 125(0 1) ontogeny of relative tail length for most species, I used Node 8 (Clade B): 31(1→2), 37(0→1), 103(1→0), species averages to score this character. 104(0→1), 106(0→1), 127(0→1), 133(0→1), State 0 obtains in all outgroup species except Elgaria 134(0→1) multicarinata. The tail in gerrhonotines and some anguines is notably long compared with most other Node 9 (Phrynosomatinae): 66(0→1), 72(1→0), anguimorphs. Thus, state 0 is considered primitive. 73(1→2), 80(1→2), 146(0→1), 148(0→1) 298 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

Sceloporus undulatus: 65(0→2), 76(1→0), 120(0→1), Node 22 (Iguaninae, Hoplocercinae): 37(0→1), 138(0→1) 88(1→2), 91(0→1), 100(0→1) Enyalioides oshaughnessyi: 1(1→0), 6(0→1), 9(0→1), Node 10 (Phrynosoma + Petrosaurus): 53(0→1), → → → → → → → → 26(0 1), 42(0 1), 47(0 1), 62(0 1), 63(0 1), 123(1 0), 125(1 0), 145(1 0) 86(0→1), 95(1→0), 102(0→1), 113(0→2), 136(0→1) Phrynosoma platyrhinos: 4(0→1), 15(0→1), 21(0→1), 25(0→1), 32(0→2), 39(0→2), 42(0→1), 46(0→1), Node 23 (Iguaninae): 2(0→1), 18(0→1), 24(0→1), 47(0→1), 51(0→2), 59(0→1), 62(0→1), 88(1→0), 58(0→1), 65(0→1), 95(1→2), 96(1→0), 145(1→0), 90(1→0), 95(1→0), 99(0→1), 114(0→1), 115(0→1), 150(0→1) 124(0→1), 136(0→1) Dipsosaurus dorsalis: 3(1→0), 31(1→2), 38(0→1), Petrosaurus thalassinus: 1(2→1), 83(2→1), 109(1→0), 50(2→3), 105(1→0), 120(0→1), 123(1→0), 113(1→0) 133(0→1), 134(0→1) Node 11 (Oplurinae + Tropidurinae*): 139(0→1) Node 24 (Brachylophus + Iguana): 79(2→1), 108(0→1), → → → → → 111(1 2), 115(0 1) Node 12 (Tropidurinae*): 67(1 2), 68(1 0), 138(0 1) Brachylophus fasciatus: 4(1→0), 31(1→0), 67(1→2) → → → Iguana iguana: 11(0→1), 51(0→1), 52(0→2), Node 13 (Liolaemini): 51(0 2), 125(1 0), 140(0 1) → → Phymaturus palluma: 1(2→3), 2(0→1), 4(0→1), 72(1 0), 126(0 1) 16(1→0), 59(0→1), 86(0→1), 88(1→2), 94(2→0), Node 25 (Corytophaninae + Polychrotinae*): 46(0→1), 95(1/2→0), 112(1→0), 123(1→0), 124(0→1) 51(0→1), 76(1→0), 125(1→0), 139(0→1) Liolaemus pictus: 36(0→1), 87(0→1), 99(0→1), 122(0→1), 126(0→1) Node 26 (Corytophaninae + stem): 27(0→1), 30(0→1), 52(0→2), 119(0→1), 152(0→1) Node 14 (Leiocephalus + Tropidurini): 135(0→1), 145(1→0) Node 27 (early Eocene iguanid + Geiseltaliellus): Leiocephalus personatus → → → 94(2→1) : 5(0 1), 16(1 0), 18(0 2), Geiseltaliellus maarius → → → 66(0→1), 88(1→2), 108(0→1), 122(0→1) : 34(0 1), 100(0 1), 101(0 1) early Eocene iguanid (see Smith 2009): No autapomor- Node 15 (Tropidurini): 12(0→1), 50(2→1), 99(0→1), phies. 126(0→1), 147(0→1/2) Node 28 (Corytophaninae): 1(1→0), 18(0→1), Node 16 (Tropidurus + Plica): 7(0→1), 120(1→0), 36(0→1), 50(2→3), 55(0→1), 62(0→1) 123(1→0) Basiliscus basiliscus: 2(0→1), 65(0→1), 74(1→0), Tropidurus torquatus: 1(2→3), 88(1→2) 112(1→2), 133(0→1), 134(0→1), 142(0→1) Plica umbra → → → → : 3(0 1), 4(0 1), 10(1 0), 42(0 1), → 66(0→1), 98(0→1), 105(0→1), 119(0→1) Node 29 (Corytophanes + Laemanctus): 53(0 1), 59(0→1), 60(0→1), 81(0→1), 114(0→1) Node 17 (Stenocercus + Microlophus): 46(0→1) Corytophanes cristatus: 21(0→1), 27(1→0), 29(0→1), Stenocercus scapularis: 1(2→1), 10(1→0), 19(1→0), 34(0→2), 39(1→2), 42(0→1), 57(0→1), 85(0→1), 37(1→0), 39(0→1), 45(0→1), 48(0→1), 115(0→1), 86(0→1), 95(1→2), 97(1→0), 105(1→0), 108(0→1), 117(0→1), 122(0→1) 121(0→1) Microlophus occipitalis: 86(0→1) Laemanctus longipes: 3(1→0), 22(1→0), 47(0→1), 122(0→1) Node 18 (Oplurinae): 83(2→0), 118(0→2), 131(0→1), 143(0→1), 144(0→1) Node 30 (Polychrotinae*): 47(0→1), 82(0→1), Oplurus quadrimaculatus: 3(0→1), 19(1→0), 36(0→1), 98(0→1), 147(0→2) 62(0→1), 64(0→1), 99(0→1), 109(1→2), 128(0→1), Pristidactylus torquatus: 31(1→2), 64(0→1), 137(0→1) 111(1→0), 120(0→1), 126(0→1), 134(0→1) Chalarodon madagascariensis → → : 27(0 1), 39(0 1), → → 94(1→2), 122(0→1), 127(1→0), 135(1→0) Node 31 (Polychrus + Anolis): 1(1 0), 29(0 1), 87(0→1), 95(1→2), 108(0→1), 113(0→1), Node 19 (Clade A): 96(0→1), 98(1→0), 111(0→1), 121(0→1), 132(0→1), 138(0→1) 113(1→0), 128(0→1) Polychrus acutirostris: 6(0→1), 7(0→1), 9(0→1), 28(0→1), 32(0→1), 66(0→1), 86(0→1), 92(0→1), Node 20 (Crotaphytinae): 18(0→1), 22(1→0), 111(1→2), 113(1→2), 115(0→1), 122(0→1), 34(0→2), 38(0→1), 64(0→1), 126(0→1), 152(0→1) 124(0→1), 129(1→0), 139(1→0) Crotaphytus collaris: 17(0→1), 47(0→1), 89(0→1), 92(0→1), 95(1→0), 115(0→1) Node 32 (Anolis): 12(0→1), 36(0→1), 50(2→1), Gambelia wislizenii: 94(0→1), 106(0→1), 119(0→1) 52(0→2), 65(0→2), 85(0→1), 89(0→1), 100(0→1), 105(1→0), 106(0→1) Node 21 (Iguaninae, Hoplocercinae, Corytophaninae, Anolis ricordi: 30(0→1), 101(0→1) Polychrotinae*): 3(0→1), 16(1→0), 23(0→1), Anolis cristatellus: 3(1→0), 10(1→0), 31(1→2), 135(1→0) 47(1→0), 61(1→0), 111(1→0) Eocene Lizards of the Clade Geiseltaliellus • Smith 299

Appendix 5: R13485), C. quinquecarinata (S: YPM R10392), Dip- Modern Specimens Examined for This Study sosaurus dorsalis (S: TMM M8936–8939, UF 55334, In addition to the modern specimens examined for this YPM R10991, 11128, 12081; W: SMF 65652–65654), study, many disarticulated specimens were examined in Iguana iguana (S: TMM M3587, 8393, 8454, YPM a course of study of fossil material (Smith 2009, online R11775), Sauromalus obesus (S: TMM M293, 4469, appendix). Institutions: FMNH, Field Museum of Nat- 8941, 8949, 8950, UF 45624, YPM R13407) ural History, Chicago, Illinois, USA; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, Hoplocercinae USA; GM, Geiseltalmuseum, Martin-Luther-Univer- Enyalioides heterolepis (W: SMF 11038), E. laticeps (W: sität, Halle-Wittenberg, Germany; MCZ, Museum of SMF 72624–72626), E. oshaughnessyi (S: SMF 67950; Comparative Zoology, Harvard University, Cambridge, W: SMF 75804), Hoplocercus spinosus (S: CAS 231483; Massachussetts, USA; SMF, Herpetology Collection, W: 62417), Morunasaurus annularis (W: SMF Senckenberg Forschungsinstitut und Naturmuseum, 78053–78056), M. groi (S: CAS 98235) Frankfurt am Main, Germany; TMM, Texas Memorial Museum, The University of Texas, Austin, Texas USA; Oplurinae UF, Florida Museum of Natural History, Gainesville, Chalarodon madagascariensis (S: TMM M8509, YPM Florida, USA; YPM, Peabody Museum of Natural His- R12804, 13520, 13521; W: YPM R14576, 14577), Oplu- tory, Yale University, New Haven, Connecticut, USA. rus cuvieri (S: CAS 231484, SMF 79191, TMM M8512), Abbreviations: S, skeletal specimens; W, alcohol-pre- O. cyclurus (S: YPM R12861), O. fierinensis (S: YPM served specimens. R13413; W: YPM R13412), O. grandidieri (S: YPM R13517; W: YPM R13414), O. quadrimaculatus (S: Agamidae* YPM R13418; W: YPM R13415, 13416) Agama agama (S: TMM M6780, 6784, 8448, 8503, 8942, 8944; W: SMF 816), Leiolepis belliana (S: SMF 57471, UF Phrynosomatinae 62046–62048, YPM R10622, 12129, 12864; W: SMF Callisaurus draconoides (S: YPM R13372, 12166; W: 10390, 10399, 10400), Physignathus cocincinus (S: SMF YPM 646, 2832), Cophosaurus texanus (S: YPM R11039; 61415, TMM M8436, 8441, UF 71685, 71686, YPM W: YPM 697, 911, 1007), Holbrookia maculata (W: R12089, 14378; W: 59678, 59579), Acanthocerus atricol- YPM 1014, 1018), Petrosaurus thalassinus (S: CAS 3009, lis (S: TMM M8439), Uromastyx acanthinurus (S: TMM TMM M8948, YPM R11846; W: YPM R14664), M8932, UF 54136, YPM R13525; W: SMF 10407–10409), Phrynosoma cornutum (W: YPM R13395), P. corona- U. geeri (S: YPM R13701), U. maliensis (S: YPM R13524) tum (S: TMM M8644, YPM 16071), P. platyrhinos (S: TMM M8951–8955), Sceloporus graciosus (S: YPM Chamaeleonidae R11087), S. magister (S: YPM R12163; W: YPM 919), S. Brookesia superciliaris (S: TMM M8675, 8676, YPM malachiticus (S: YPM R11950; W: YPM R13003– R11370) 13005), S. occidentalis (S: TMM M8940, YPM R10439), S. olivaceus (S: YPM R11409), S. undulatus (S: TMM Corytophaninae M8934, 8935; W: SMF 41657), S. variabilis (W: YPM R14005, 14006, 14008), Uma notata (YPM 649), U. sco- Basiliscus galeritus (S: UF 61491; W: SMF 11029), B. paria (S: CAS 42135, YPM R11090), Urosaurus gracio- basiliscus (S: UF 99655; W: SMF 24882, 50919–50925), sus (S: YPM R10970), U. microscutatus (W: YPM 6692), B. plumifrons (S: YPM R11644, 12506; W: SMF 24882), U. ornatus (S: YPM R12162), Uta stansburiana (S: YPM B. vittatus (S: YPM R10428, 11129; W: SMF 11019, R12165, R11004; W: YPM 2846, 2851) 42109, 42112, 42113, 42119, 47207, 48080, 52055, 52060, 53574, 53684), Corytophanes cristatus (S: UF 69072, YPM R10443, 11095, 11183, 11184; W: SMF 81504), C. Polychrotinae* hernandesii (S: UF 72492; W: SMF 11015), C. perceri- Anisolepis undulatus (W: SMF 11055, 11056), Anolis natus (W: SMF 80733–80738), Laemanctus longipes (S: chlorocyanus (S: MCZ 57488, 57490, UF 42494, 99949), UF 66061, YPM R10434, 10435, 13196; W: SMF 11017 A. cobanensis (W: SMF 82688), A. cristatellus (S: MCZ or 11018 [jaws closed]), L. serratus (W: SMF 11016) 85134, 85135, UF 99457, YPM R12050; W: YPM R12074), A. cuvieri (W: YPM R12066), A. distichus (W: Crotaphytinae YPM 3993), A. polylepis (W: SMF 81818), A. richteri (W: SMF 60817, 60820), A. ricordi (S: UF 64820, 99672; W: Crotaphytus bicinctores (S: TMM M8947), C. collaris (S: SMF 24859), A. trinitatus (S: TMM M8943), A. wetmorei YPM R11208, 12160, TMM M8395; W: YPM 1144, (W: YPM 1866), Diplolaemus bibronii (W: SMF 58523, 1147, 1220), C. insularis (S: TMM M8933), Gambelia 58524), Enyalius bilineatus (W: SMF 11054), E. catena- wislizenii (S: CAS 200856, TMM M1474, YPM R10367, tus (W: SMF 11040–11041), Leiosaurus belli (S: MCZ 11219; W: YPM 7135, 7818) 162921; W: SMF 11066, 11067), L. catamarcensis (S: MCZ 58116, 58117), L. paronae (S: MCZ 162923), Poly- Iguaninae chrus acutirostris (S: SMF 24870; W: SMF 62421), P. Brachylophus fasciatus (W: SMF 73361–73364), Cyclura femoralis (S: FMNH 81405), P. gutturosus (S: MCZ cornuta (S: YPM R10438), Ctenosaura palearis (S: YPM 46441, UF 49377), P. marmoratus (S: YPM R13556; W: 300 Bulletin of the Peabody Museum of Natural History 50(2) • October 2009

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