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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

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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 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 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. 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 difficul- ties 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 spec- imens (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 pres- ence 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 re- lationships remains to be resolved. Keywords Iguanidae, Corytophaninae, Messel, Geiseltal, historical biogeography, molecular-genetic. Introduction of Gauthier et al. 1988 and Schulte et al. 2003; pre- viously 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-
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