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1、.ONLINE SUPPLEMENTARY MATERIAL 1Contents1. Faunal List2. Stratigraphy3. Species Descriptions4. Phylogenetic Analysisa. Revisions to Gauthier et al. 2012 matrixb. New Characters / Character Illustrationsc. Materials Studiedd. Methodse. Resultsf. Discussion5. Resampling6. Morphometric Analysis7. Phylo
2、genetic Independent Contrasts Analyses8. References9. Character-Taxon Matrix1. Faunal List, Upper Maastrichtian of Western North AmericaBased on literature (1-6) and museum specimens (see SI 3 for complete references and specimen numbers) . Species in bold are either named or recognized here for the
3、 first time. Table S1.IGUANIAIguanidae incertae sedisPariguana lancensisPOLYGLYPHANODONTIAPolyglyphanodontia incertae sedisObamadon gracilisPolyglyphanodontidaePolyglyphanodon sternbergiChamopsiidaeChamops segnisLeptochamops denticulatusMeniscognathus altmaniHaptosphenus placodonStypodontosaurus mel
4、letesTripennaculus n. sp.Frenchman chamopsiidPeneteius aquiloniusSocognathus brachyodonLaramie chamopsiidSCINCOMORPHAScincomorpha incertae sedisLonchisaurus trichurusScincoidea incertae sedisEstescincosaurus cookiGlobauridaeContogenys sloaniANGUIMORPHAXenosauridaeExostinus lancensisAnguidaeOdaxosaur
5、us piger“Gerrhonotus” sp.Platynota incertae sedisLitakis gilmoreiColpodontosaurus cracensPalaeosaniwa sp.Paraderma bogertiCemeterius monstrosusParasaniwa wyomingensisOPHIDIAConiophidaeConiophis precedensAlethinophidia incertae sedisCerberophis robustusLance SnakeSQUAMATA INCERTAE SEDISLamiasaurus fe
6、roxSweetwater County lizard2. StratigraphyA. MaastrichtianThe Cretaceous lizards included in this study come from the Hell Creek Formation of Montana, the Lance and Ferris formations of Wyoming, the Frenchman Formation of Saskatchewan, the Scollard Formation of Alberta, the Laramie Formation of Colo
7、rado, and the North Horn Formation of Utah. All formations are inferred to be late Maastrichtian in age and contain typical late Maastrichtian faunas (i.e., Lancian Land Vertebrate Age). Age constraint (Table S2) is based on several lines of evidence, including radiometric dating, magnetostratigraph
8、y, palynostratigraphy, and vertebrate biostratigraphy, which is primarily based on mammalian faunas.i. Frenchman FormationThe entire Frenchman Formation is constrained on the basis of magnetostratigraphy to the final reversed interval of the Cretaceous, C29r (7), which spans the final 300,000 years
9、of the Cretaceous (8) and so the Frenchman lizards are no older than 300,000 years. It is unclear where the Gryde locality (which produced the Frenchman lizards discussed here) lies in section, but it lies 20 meters up the exposed outcrop, and the entire formation is a little more than 50 meters thi
10、ck in this area (9). This suggests that the Gryde lizards date to significantly less than 300,000 years before the K-Pg boundary.ii. Lance FormationThere are fewer constraints on the age of the Lance Formation in eastern Wyoming because no radiometric dates or magnetostratigraphic studies have been
11、published. Here, the K-Pg claystone occurs just below the basal lignite in the Fort Union Formation, at the very top of the Lance Formation; it displays the signature features of the Chicxulub impact, including an iridium anomaly, a layer of spherules, shock-metamorphosed mineral grains, and the abr
12、upt disappearance of many pollen species, followed by the appearance of a layer rich in fern spores (10). In this area, the Lance Formation is roughly 2,500 feet thick, and UCMP V5620 (Lull 2) lies 2,100 feet above the base of the Lance, and UCMP V5711 (Bushy Tailed Blowout) lies in the top half of
13、the formation (11). Based on some assumptions that are admittedly simplistic, i.e. that the Lance was deposited at a constant rate over the final 1.3 Ma of the Cretaceous (as found for the Hell Creek Formation of North Dakota: Hicks et al., 2002), this would indicate ages of 200,000 and <650,000
14、years before the K-Pg boundary for UCMP V5620 and UCMP V5711, respectively. The occurrence of the marsupial Glasbius in both sites, a genus that appears between 900,000 and 600,000 years before the K-T boundary in Montana (12) also supports a latest Maastrichtian age, assuming that the introduction
15、of Glasbius occured more or less simultaneously across the region. Palynostratigraphic studies of this area (13) also show that UCMP V5620 lies well above the first occurrence of pollen species that occur in the Latest Cretaceous in North Dakota. These are Striatopollis tectatus and Liliacidites alt
16、murus. In the Hell Creek Formation, these species appears about 40 meters below the boundary (14), which would correspond 500,000 years before the K-Pg boundary (8). Again, assuming that they appeared simultaneously across the region, then this would support a latest Maastrichtian age for the lizard
17、s occurring in the Lull 2 locality.The same assumptions regarding stratigraphy would suggest an age of 200,000 years before the K-Pg boundary for the Black Butte assemblage in Sweetwater County, southern Wyoming (UW V-79032) which lies 33 meters below the K-T transition in a 233 meter section (15).
18、iii. Hell Creek FormationThe Hell Creek of Garfield/McCone Counties in Montana spans the final 1.8 million years of the Cretaceous (12); unfortunately no published stratigraphic data are available for the localities that produced the lizards described here. No stratigraphic data are available for th
19、e lizards from Carter County and Dawson County, Montana, but similar ages are assumed here. iv. Scollard FormationThe Cretaceous portion of the Scollard can be constrained to the final 1.5 million years of the Cretaceous (16).v. Laramie FormationThe Laramie site UCM 77062 is the oldest site in this
20、study, dating to 2.5 Ma before the K-T (17).vi. North Horn Formation.The age of the Cretaceous exposures of the North Horn Formation in Utah remains uncertain, but on the basis of the mammal fauna it is assigned to the Late Maastrichtian (18).vii. Bighorn Basin LanceThe stratigraphy of the Bighorn B
21、asin sites is not well-known but they contain characteristic Lancian mammals (19) suggesting a Late Maastrichtian age; furthermore the occurrence of Glasbius in the Hewitts Foresight locality (19) supports this assignment.viii. Ferris FormationFinally, the age of the Cretaceous part of the Ferris Fo
22、rmation is not well-constrained, but on the basis of the fauna these assemblages are late Maastrichtian in age (20). The sole lizard reported from the Ferris, Odaxosaurus, occurs high in section but given that it crosses the K-Pg boundary, the precise age of this specimen is not an issue.It should b
23、e kept in mind that the the point here is not to make definitive claims about the precise age of last occurrence for each taxon, but rather to emphasize that the fauna described here is as close as one can possibly come to providing a picture of what the squamate fauna looked like just before the K-
24、T impact, within the inevitable constraints imposed by an imprecise understanding of the chronostratigraphy of these formations, and an extremely patchy fossil record. Given that a number of lizard genera appear to have stratigraphic ranges of 10 Ma or more (SI3), it is unlikely that a significant p
25、ercentage of the species described here became extinct in the hundreds of thousands of years that separate these localities from the K-T boundary, and the various localities can also be assumed to represent, more or less, a picture of a single fauna rather than a succession of faunas.B. PaleoceneThe
26、 ages of the Palaeocene localities are less of an issue: a survivor is a survivor regardless of whether it occurs a few tens of thousands of years after the K-T boundary, or millions of years later. Therefore, for the purposes of inferring survival and for rarefaction, the analysis of Paleocene spec
27、ies lumped together localities from the Puercan and Torrejonian land vertebrate ages, a period spanning some 4 million years (21), because relatively few Puercan localities are known. Inferences of Puercan and Torrejonian age have previously been made on the basis of the co-occurring mammals. Insofa
28、r as this lumping of assemblages might tend to bias the study, treating these various assemblages as a single fauna would inflate Paleocene diversity and make the fauna appear more diverse and less well-sampled than it really is. Given that this would bias the study against the hypothesis we are inv
29、estigating (i.e. a major drop in diversity from the Maastrichtian to Palaeocene) we argue that this assumption is conservative. The Bug Creek fauna, however, is problematic in representing a mixed assemblage (22, 23) containing Palaeocene and reworked Maastrichtian species. For examining Paleaocene
30、diversity, we assumed the Bug Creek lizard fossils were Palaeocene if they were (a) from species known to cross the K-Pg boundary based on undisputed Palaeocene fossils occurring elsewhere, or (b) they represented species entirely unknown from the Maastrichtian. Although it is conceivable that some
31、of these specimens could be reworked from the Maastrichtian, in this case, it would only bias the data against the conclusions of this paper by inflating Paleocene diversity, and in this sense our assumptions are again conservative. Table S2. Stratigraphically highest occurrences for species failing
32、 to cross the K-T boundary. Occurrence data: Estes, 1964; Gao and Fox, 1996, this study (SI3). TaxonOccurrenceDateChamops segnisFrenchman Fm., SK300,000 years before K-TLeptochamops denticulatusFrenchman Fm., SK300,000 years before K-TMeniscognathus altmaniFrenchman Fm., SK300,000 years before K-THa
33、ptosphenus placodonFrenchman Fm., SK300,000 years before K-TTripennaculus sp.Frenchman Fm., SK300,000 years before K-TFrenchman chamopsiidFrenchman Fm., SK300,000 years before K-TParaderma bogertiFrenchman Fm., SK300,000 years before K-TParasaniwa wyomingensisFrenchman Fm., SK300,000 years before K-
34、TLamiasaurus feroxLance Formation, Sweetwater County, WY200,000 years before K-T (?)Sweetwater LizardLance Formation, Sweetwater County, WY200,000 years before K-T (?)Cemeterius monstrosusUCMP V5620, Lance Fm., WY250,000 years before K-T(?)Litakis gilmoreiUCMP V5620, Lance Fm., WY250,000 years befor
35、e K-T(?)Palaeosaniwa canadensisUCMP V5620, Lance Fm., WY250,000 years before K-T(?)Colpodontosaurus cracensUCMP V5620, Lance Fm., WY250,000 years before K-T(?)Estescincosaurus cookiUCMP V5620, Lance Fm., WY250,000 years before K-T(?)Lance snakeUCMP V5711, Lance Fm., WY650,000 years before K-T(?)Lonc
36、hisaurus trichurusUCMP V5711, Lance Fm., WY650,000 years before K-T(?)Pariguana lancensisUCMP V5711, Lance Fm., WY650,000 years before K-T(?)Stypodontosaurus melletesScollard Fm., AB1.5 Ma before K-TCerberophis robustusUCMP V80096, Hell Creek Fm., MT1.8 Ma before K-T Obamadon gracilisUCMP V74116, He
37、ll Creek Fm., MT1.8 Ma before K-TPeneteius aquiloniusUCMP V74116, Hell Creek Fm., MT1.8 Ma before K-TLaramie chamopsiidUCM 77062, Laramie Fm., CO2.5 Ma before K-TSocognathus brachyodonLance Formation, Park County, WYLate MaastrichtianPolyglyphanodon sternbergiNorth Horn Formation, UTLate Maastrichti
38、an3. Species DescriptionsInstitutional abbreviations. AMNH, American Museum of Natural History, New York, New York; UCM, University of Colorado Museum, Boulder, Colorado; UCMP, University of California Museum, Berkeley, California; USNM, Smithsonian Institution National Museum of Natural History, Wa
39、shington, DC; YPM, Yale Peabody Museum, New Haven, Connecticut; YPM-PU, Yale Peabody Museum, Princeton University collection.REPTILIA Laurenti, 1768SQUAMATA Oppel, 1811IGUANIA Cope 1864IGUANIDAE Oppel 1811Pariguana new genusEtymology. The genus name is derived from Greek para, near, and the genus na
40、me Iguana.Diagnosis. As for the type and only species.Type species. Pariguana lancensis. Pariguana lancensis new speciesEtymology. The specific name refers to the Lance Formation.Diagnosis. Small iguanid characterized by the following combination of characters: teeth tall, slender, with tapering cro
41、wns and weak accessory cusps; coronoid extends forward on lateral surface of jaw beneath last tooth, Meckelian groove constricted suddenly ahead of anterior inferior alveolar foramen.Holotype. AMNH 22208, partial mandible. Locality and Horizon. UCMP V5711, Bushy Tailed Blowout; upper Maastrichtian L
42、ance Formation, Niobrara County, eastern Wyoming.Description. The type and only known specimen consists of the posterior half of a dentary and the associated parts of the splenial, coronoid, and angular. The dentary is weakly bowed in lateral view and extends posteriorly well beyond the toothrow and
43、 below the coronoid. In medial view, the Meckelian groove is broad posteriorly, but exhibits a sharp taper just ahead of the anterior inferior alveolar foramen, where the splenial is pinched between the subdental ridge and the ventral margin of the dentary. This distinctive taper distinguishes Parig
44、uana from the Paleocene iguanid Swainiguanoides milleri (24). The subdental ridge is also flat medially, in contrast to the more rounded subdental ridge of Swainiguanoides. The ventral margin of the dentary wraps beneath the splenial, an iguanian synapomorphy. The splenial has a large, elliptical an
45、terior inferior alveolar foramen, and below and slightly behind it a smaller mylohyoid foramen. A long, slender sliver of the angular extends well forward between the splenial and angular. The coronoid extends beneath the subdental ridge medially, and on the lateral surface of the dentary there is a
46、 facet where the coronoid would extended onto the dorsolateral surface of the dentary. This anterolateral process of the dentary is a derived feature seen in a number of crown iguanids.There are 13 tooth positions preserved, and the complete mandible would probably have had between 20 and 30 teeth.
47、Implantation is pleurodont. Several teeth bear large resorption pits at their bases, and four teeth in the back of the jaw are short and implanted at odd angles, a pathology. Tooth crowns are tall and slender, with tapered tips. The tip is tricuspid, with a broad, blunt central cusp flanked by tiny
48、mesial and distal cusps. Medially, the accessory cusps are separated from the central cusp by grooves. The teeth generally resemble those of Swainiguanoides, but differ in having a narrower crown. Discussion. Pariguana can be referred to the Iguania on the basis of the constriction of the Meckelian
49、groove by the ventral margin of the dentary. Pariguana exhibits two features that are found in crown Iguanidae but not among stem Iguanidae from the Cretaceous of Mongolia (e.g. Temujinia, Zapsosaurus): anterior extension of the coronoid beneath the toothrow on the lateral surface of the jaw, and po
50、sterior extension of the dentary beneath the coronoid process. On this basis, Pariguana appears to represent either the sister taxon of crown Iguanidae, or a member of the crown; in the present analysis Pariguana is the sister taxon to Hoplocercidae. Given the uncertainty over the relationships of c
51、rown iguanids, and the limited material available for Pariguana, this conclusion should be considered tentative. Although Pariguana closely resembles the Paleocene Swainiguanoides milleri, it differs in the strongly tapering shape of the Meckelian groove, the flat subdental ridge, and in having more
52、 strongly tapered tooth crowns.The earliest and most primitive known iguanids and acrodonts are Asian, suggesting that the Iguania originated in Asia and later migrated into North America (Gauthier et al., 2012). Until now, the earliest known iguanids were Paleocene, which suggested that this migrat
53、ion occurred in the Palaeocene (Gauthier et al., 2012); the presence of Pariguana in the Maastrichtian of North America shows that this immigration event had occurred prior to the end of the Cretaceous. The end-Cretaceous mass extinction appears to have eliminated the Asian iguanids, but spared the
54、North American lineage, leading to a North American radiation of iguanids in the Palaeogene. Although Gao and Fox (6) identify several Cretaceous lizards as iguanids, these specimens do not exhibit any derived features that allow confident referral to either Iguania or Iguanidae. In particular, the
55、Maastrichtian fossils they refer to Iguanidae can be referred to Tripennaculus, an unusual polyglyphanodontian with strongly tricuspid teeth (25); the others appear to represent polyglyphanodontians as well. Accordingly, Pariguana represents the earliest definitive evidence of Iguanidae in North Ame
56、rica. Fig. S1. Pariguana lancensis n. gen et sp., AMNH 22208 left dentary. In A, medial view; B, dorsal view; C, ventral view, D, lateral view. Abbreviations: aiaf, anterior inferior alveolar foramen; cf, coronoid facet; mf, mylohyoid foramen; sdr, subdental ridge; vl, ventral lip of dentary.POLYGLY
57、PHANODONTIA Alifanov 2000Definition. Polyglyphanodon sternbergi and all species closer to Polyglyphanodon sternbergi than to any extant species. This taxon includes Borioteiioidea as defined by Nydam et al. (26). However, Borioteiioidea is a node-based taxon, whereas Polygyphanodontia is a stem-based taxon, and would therefore include any species which were found to lie along the stem of Borioteiioidea. POLYGLYPHANODONTIA INCERTAE SEDISObamadon new genusEtymology. The genus name refers to Barack Hussein Obama and odon (Greek) = tooth, in referen
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