Foster, J.R. and Lucas, S. G. R.M., eds., 2006, Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin 36. 233 THE FAUNA AND FLORA OF THE MORRISON FORMATION: 2006
DANIEL J. CHURE1, RON LITWIN2, STEPHEN T. HASIOTIS3, 4, EMMETT EVANOFF5 AND KENNETH CARPENTER6
1 Dinosaur National Monument, Box 92, Jensen, UT 84035; 2 MS 970, U.S. Geological Survey, Reston, VA 22092; 3 Department of Geology and 4 Natural History Museum and Biodiversity Research Center, University of Kansas, 120 Lindley Hall, 1475 Jayhawk Blvd., Lawrence, KS 66045-7613; 5 Geology Section, University of Colorado Museum, 265 UCB, Boulder, CO 80309-0265; 6 Department of Earth Sciences, Denver Museum of Natural History and Science, 2001 Colorado Boulevard, Denver, CO 80205-5798
The Morrison Formation is best known for its abundant and diverse sent hidden biodiversity—that is diversity not counted in any taxonomic dinosaur remains. Exposures of the Morrison provided much of the mate- compilations based solely on body fossils (Hasiotis, 2000, 2002, 2003). rial for the “dinosaur gold rush” in the western US during the last half of For example, the entire record of Morrison insects, a group which must the nineteenth century. These specimens have made immense and funda- have been both abundant and diverse in Morrison ecosystems, is based mental contributions to our understanding of Mesozoic continental eco- solely on ichnofossils. Not a single insect body fossil has yet been reported systems. Thus, dinosaurs dominated the paleontological research in the from this rock unit. We recognize that the assignment of a trace fossil to a Morrison Formation until the latter third of the 20th century and remain a tracemaker is fraught with difficulties. As is clear from Hasiotis (2004), major research effort in the formation. there may be several candidate tracemakers for a single type of trace fossil. Over the last two decades, however, more field work and research In such cases, we have listed the fossil under all potential tracemakers when has focused on the questions related to the composition, structure, and so indicated by the author of the source paper. A good example of this is the evolution of Morrison ecosystems. These efforts revealed an unexpectedly diverse nest types listed under the various families of the termite order rich record of paleobiodiversity. Many of these discoveries have been re- Isoptera. ported across a wide spectrum of scientific periodicals, some of which are We thank Brooks Britt, William Tidwell, Fred Peterson, and John obscure. These publications range from broader systematic reviews to a Foster for their reviews. focus on a single species or a small taxonomic group. Chure et al. (1998) was the first modern attempt to compile a complete floral and faunal list for I. BODY FOSSILS the Morrison using all the literature, including trace fossils. Significant ad- vances have been made since that publication, and an update is timely. We KINGDOM ANIMALAE have followed the format of our 1998 paper, including both body and trace fossils. We have placed fossils, whether named or not, in a family—gener- PHYLUM PORIFERA ally following the assignment given in the source citations—in the belief CLASS DEMOSPONGEA that such a low-level taxonomic assignment will be helpful to those using Family Spongillidae this list, especially for groups the user many not be intimate with. For each family we have given the citations used to compile that Eospongia morrisonensis entry. While these citations may be the original description, more compre- [Sources: Dunagan 1999] hensive reviews have also served as sources. For example dinosaurs are well reviewed in the various chapters in Weishampel et al. (2004). Simi- CLASS incertae sedis larly, we have followed Kielan-Jaworowska et al. (2004) for the composi- Family incertae sedis tion of the Morrison mammal fauna. These thorough works provide an Unidentified sponge spicules entry into the primary scientific literature for those groups. Spicule-like remains Given space limitations, the Latin binomials listed are only those [Sources: Waldschmidt and LeRoy 1944; formal names considered valid by the present authors. This is not a com- Yen and Reeside 1952 p. 31] prehensive list of all names proposed for Morrison fossils. For the same reasons we have not listed synonyms for those names that we consider PHYLUM MOLLUSCA valid. Such synonymy lists are available elsewhere for some groups (Weishampel et al., 2004; Kielan-Jaworowska et al., 2004). For most groups, CLASS BIVALVIA however, such revisions are yet to be written. Family Unionidae In contrast to our 1998 paper, we have greatly reduced the number Unio baileyi of personal communications. In part this is because the specimens involved U. felchi have subsequently been described and are now in the faunal list as a for- U. iroides mally-named species. We now feel that personal communications about U. knighti unidentified specimens may be of little use to the reader and might actually U. lapilloides confuse matters. So, while a few personal communications remain, they U. macropisthus have by and large been omitted. We have included fossils that are distinct U. mammillaris but have not been named, using the description given by the author in the U. nucalis source paper. U. stewardi Although the Morrison Formation is predominantly a continental U. stewardi utahensis deposit, there are marine units in the Windy Hill Member. Hence, the pres- U. toxonotus ence of what might at first appear to be anomalous fossils, such as the U. willistoni anthozoan trace fossil Conichnus isp., is in fact real occurrences of marine Vetulonaia faberi body and trace fossils. V. mayoworthensis We included named and unnamed ichnofossils in our list because V. whitei our intent is to present the most complete record of biotic diversity that Hadrodon jurassicus existed during deposition of the Morrison Formation. Ichnofossils are ex- H. lateralis ceedingly important for this type of information because trace fossils repre- H. trigonus 234 [Sources: Evanoff, Good, and Hanley PHYLUM ARTHROPODA 1998; Yen and Reeside 1952; Good 2004] CLASS CRUSTACEA Subclass Branchiopoda CLASS GASTROPODA Suborder Conchostraca Family Neritidae Family Cyzicidae Mesoneritina morrisonensis Cyzicus cf. tendaguruensis [Sources: Yen and Reeside 1952; Euestheris sp. Evanoff, Good, and Hanley 1998] Lioestheria tendagurensis Family Cyclophoridae Lioestheria sp. A Amplovalvata cyclostoma Lioestheria sp. B A. scabrida Nestoria krasinetzi A. scabrida leei [Sources: Hassinger 1959; Kirkland and A.? reesidei Armstrong 1992; Lockley 1986; Lucas and [Sources: Yen and Reeside 1952; Kirkland 1998] Evanoff, Good, and Hanley 1998] Family Viviparidae Subclass Ostracoda Viviparus morrisonensis Family Darwinulidae V. reesidei Darwinula acuminata [Sources: Yen and Reeside 1952; D. cf dakotensis Evanoff, Good, and Hanley 1998] [Sources: Branson 1935; Hassinger 1959; Family Pleuroceridae Sohn 1958] Lioplacodes jurassicus Family Cytherideidae [Sources: Yen and Reeside 1952;Evanoff, Cytheridea atlantosaurica Good, and Hanley 1998] Cushmanidea marshi Family Bithyniidae [Sources: Branson 1935; Sohn 1958] Reesidella gilloides Family Paracypridiae R. jurassica Paracypris simplus [Sources: Yen and Reeside 1952; [Sources: Hassinger 1959] Evanoff, Good, and Hanley 1998] Family Cyprididae Family Valvatidae Cypris purbeckensis Liratina jurassica Candona coloradensis Tropidina jurassica C. morrisonensis “Pentagoniostoma” altispiratum [Sources: Hassinger 1959; Schudack [Sources: Yen and Reeside 1952; 1995, 1998] Evanoff, Good, and Hanley 1998] Family Limnocytheridae Family Amnicolidae Bisulcocypris pahasapensis Amnicola gilloides B. pustulosa A. jurassica Metacypris bradyi [Sources: Yen and Reeside 1952; M. minnekahtensis Evanoff, Good, and Hanley 1998] M. whitei Family Elobiidae Gomphocythere sp. Mesauriculstra accelerata Helmdachia petersoni M. morrisonensis H. prima M. morrisonensis ovalis H. turneri Mesochilina spiralis Timiriasevia guimarotensis [Sources: Yen and Reeside 1952; [Sources: Branson 1935; Hassinger 1959; Evanoff, Good, and Hanley 1998] Peck 1959; Sohn 1958; Schudack 1998] Family Otinidae Family Dryelbidae Limnopsis jurassica Dryelba pustulosa [Sources: Yen and Reeside 1952; Theriosynoecum wyomingense Evanoff, Good, and Hanley 1998] [Sources: Branson 1935; Sohn 1982] Family Lymnaeidae Family Ilyocyprididae Lymnaea ativuncula Cypridea acuticyatha L. consortis Rhinocypris jurassica L. morrisonensis [Sources: Schudack 1998] [Sources: Yen and Reeside 1952; Family incertae sedis Evanoff, Good, and Hanley 1998] Cetacella armata Family Planorbidae C. striata Gyraulus veternus Cetacella sp. [Sources: Yen and Reeside 1952; Trapezoidella aff. rothi Evanoff, Good, and Hanley 1998] [Sources: Schudack 1995] 235 Subclass Malacostraca Order Caudata Family Cambaridae Family incertae sedis Undescribed crayfish Comonecturoides marshi [Sources: Callison 1987; Hasiotis and Iridotriton hechti Demko 1998] New long bodied salamander (D.J. Chure, personal observation 1999). PHYLUM CHORDATA [Sources: Evans and Milner 1993; Hecht CLASS OSTEICHTHYES and Estes 1960; Evans, Lally, Chure, and Order Paleonisciformes Elder 2005] Family Coccolepididae CLASS REPTILIA Morrolepis schaefferi Order Testudines [Sources: Kirkland 1998] Family Pleurosternidae Family incertae sedis Dinochelys whitei ?paleoniscid dentary Glyptops plicatulus [Sources: Jensen and Padian 1989] Uluops uluops Order Semionotiformes [Sources: Gaffney 1979; Carpenter and Family Semionotidae Bakker 1990; Bakker 1998] Undescribed semionotid Family Baenidae [Sources: Kirkland 1998] Dorsetochelys buzzops [Sources: Bakker 1998] Order Halecostomi Family Leptolepididae Order Sphenodonta cf. Leptolepis sp. Family Sphenodontidae [Sources: Kirkland 1998] Opisthias rarus Family incertae sedis Eilenodon robustus Hulettia hawesi Theretairus antiquus [Sources: Kirkland 1998] [Sources: Simpson 1926; Callison 1987; Gilmore 1909; Rasmussen and Callison Order Amiiformes 1981a] Family incertae sedis Ophiopsis sp. Order Choristodera Unnamed amioid Family incertae sedis [Sources: Prothero 1981; Kirkland 1998] Cteniogenys antiquus Order Ceratodontiformes [Sources: Gilmore 1928; Evans 1990] Family Ceratodontidae Order Squamata Ceratodus felchi Suborder Lacertilia C. fossanovum Family Paramacellodidae C. cf. C. frazieri Paramacellodus sp. C. guentheri [Sources: Evans and Chure 1998 a,b, C. robustus 1999] [Sources: Kirkland 1987, 1998; Foster Family Dorsetisauridae 2003] Dorsetisaurus sp. Order Pycnodontoidea [Sources: Evans and Chure 1999] Family incertae sedis. Family incertae sedis Unnamed genus and species Parviraptor gilmorei [Sources: Kirkland 1998] Saurillodon sp. Schilleria utahensis CLASS AMPHIBIA New scincomorph lizard A Order Anura New scincomorph lizard B Family Discoglossidae New dorsetisaur lizard Enneabatrachus hechti New lizard [Sources: Evans and Milner 1993] [Sources: Callison 1987; Evans 1996; Family Pelobatidae Evans and Chure 1999] Unnamed pelobatid [Sources: Evans and Milner 1993] Suborder ?Serpentes Family incertae sedis Family ?Booiidae Eobatrachus agilis ?New genus and species of snake Comobatrachus aenigmatis [Sources: Callison 1987; Foster 2003] Rhadinosteus parvus Order Crocodylomorpha [Sources: Hecht and Estes 1960; Henrici Family Sphenosuchidae 1998] Macelognathus vagans [Sources: Ostrom 1971; Göhlich, Chiappe, Clark, and Sues 2005] 236 Family Atoposauridae [Sources: Chure 1994; Hartman, Hoplosuchus kayi Lovelace, and Wahl 2005] [Sources: Gilmore 1926; Clark 1985] Family Tyrannosauridae Family Goniopholidae ?Aviatyrannis jurassica Amphicotylus lucasii Stokesosaurus clevelandi Goniopholis felix [Sources: Britt 1991; Rauhut 2003] G. gilmorei Family incertae sedis G. lucasii Elaphrosaurus sp. G. stovalli Marshosaurus bicentissimus Eutretauranosuchus delfsi Dromaeosaurid-like teeth [Sources: Clark 1985; Foster 2003] Small maniraptoran ulna Family incertae sedis [Sources: Britt 1991; Galton 1982; “Fruitachampsa callisoni” Jensen and Padian 1989] New mesoeucrocodilian very similar to Family incertae sedis “Fruitachampsa” (James Ornitholestes hermanni Clark, personal communication Coelurus fragilis 2005) Tanycolagreus topwilsoni Hallopus victor Unnamed maniraptoran [Sources: Steel 1973; Clark 1985; Chure, [Sources: Callison 1987; Molnar, Madsen, and Armstrong; 1991; Foster Kurzanov, and Dong 1990a; Makovicky 2003] 1997; Holtz, Molnar, and Currie 2004; Carpenter, Miles, and Cloward 2005] Order Pterosauria Suborder Rhamphorhynchoidea Suborder Sauropoda Family Rhamphorhynchidae Family Cetiosauridae Harpactognathus gentryii Haplocanthosaurus priscus [Sources: Carpenter, Unwin, Cloward, H. delfsi Miles, and Miles 2003] [Sources: McIntosh 1990; Upchurch, Family incertae sedis Barrett, and Dodson 2004] Comodactylus ostromi Family Brachiosauridae Utahdactylus kateae Brachiosaurus altithorax [Sources: Wellnhofer 1991; Czerkas and Dystylosaurus edwini Mickelson 2002] [Sources: McIntosh 1990; Upchurch, Suborder Pterodactyloidea Barrett, and Dodson 2004] Family incertae sedis Family Camarasauridae Mesadactylus ornithosphyos Camarasaurus supremus Dermodactylus montanus C. grandis Kepodactylus imperatus C. lentus Laopteryx priscus C. lewisi [Sources: Callison 1987; Wellnhofer 1991; [Sources: McIntosh 1990; Upchurch, Harris and Carpenter 1996; Ostrom 1986] Barrett, and Dodson 2004] Family Diplodocidae Order Saurischia Amphicoelias altus Suborder Theropoda A. fragillimus Family Ceratosauridae Apatosaurus ajax Ceratosaurus nasicornis A. excelsus C. magnicornis A. louisae C. dentisulcatus Barosaurus lentus [Sources: Madsen and Welles 2000; Diplodocus longus Tykoski and Rowe 2004] D. carnegii Family Megalosauridae D. hayi Torvosaurus tanneri D. lacustris [Sources: Britt 1991; Holtz, Molnar, and Dyslocosaurus polyonychius Currie 2004] Seismosaurus hallorum Family Allosauridae Supersaurus vivianae Allosaurus fragilis Suuwassea emilieae A. jimmadseni [Sources: Gillette 1991; McIntosh 1990; Saurophaganax maximus McIntosh, Coombs, and Russell 1992; [Sources: Madsen 1976; Chure 1995, Harris and Dodson 2004] 2000; Holtz, Molnar, and Currie 2004] Family incerate sedis Family ?Troodontidae Dystrophaeus viaemalae Koparion douglassi Unnamed cetiosaurid (= “Morosaurus” Unnamed troodontid agilis) 237 Unnamed cetiosaurid (= “Apatosaurus” Order Docodonta minimus) Family Docodontidae [Sources: McIntosh 1990; Gillette 1996; Docodon victor Turner and Peterson 1999; D. sp. Upchurch, Barrett, [Sources: Kielan-Jaworowska, Cifelli, and and Dodson 2004] Luo 2004] Order Ornithischia Order Multituberculata Suborder Ornithopoda Family Allodontidae Family Heterodontosauridae Ctenacodon laticeps Echinodon sp. C. scindens [Sources: Callison 1987; Galton 2002] C. serratus Family Hypsilophodontidae C. sp. Othnielia rex Psalodon fortis [Sources: Sues and Norman 1990; P. marshi Norman, Sues, Witmer, and Coria 2004, ?P. marshi Galton in press] P. potens Family Dryosauridae ?P. sp. Dryosaurus altus Gen. et sp. indet. [Sources: Sues and Norman 1990] [Sources: Engelmann and Callison 1999; Family Camptosauridae Kielan-Jaworowska, Cifelli, and Luo Camptosaurus dispar 2004 ] C. amplus Family Plagiaulacidae [Sources: Norman and Weishampel “Ctenacodon” brentbaatar 1990] [Sources: Kielan-Jaworowska, Cifelli, and Family incertae sedis Luo 2004] Drinker nisti Family Zofiabataaridae [Sources: Sues and Norman 1990; Norman, Zofiabaatar pulcher Sues, Witmer, and Coria 2004] [Sources: Kielan-Jaworowska, Cifelli, and Luo 2004] Suborder Stegosauria Family incertae sedis Family Stegosauridae Glirodon grandis Stegosaurus armatus [Sources: Engelmann and Callison 1999; S. longispinus Kielan-Jaworowska, Cifelli, and Luo 2004] S. stenops S. ungulatus Order Symmetrodonta Hesperosaurus mjosi Family Amphiodontidae [Sources: Galton 1990; Galton and Amphiodon superstes Upchurch 2004] [Sources: Kielan-Jaworowska, Cifelli, and Luo 2004] Suborder Ankylosauria Family Tinodontidae Family Polacanthidae Tinodon bellus Gargoyleosaurus parkpinorum T. sp. Mymoorapelta maysi [Sources: Kielan-Jaworowska, Cifelli, and [Sources: Kirkland and Carpenter 1994] Luo 2004] CLASS MAMMALIA Order Eupantotheria Order Triconodonta Family Dryolestidae Family Amphilestidae Dryolestes priscus Comodon gidleyi cf. D. priscus Aploconodon comoensis Laolestes eminens Triconolestes curvicuspis L. goodrichi [Sources: Engelmann and Callison 1998; L. oweni Kielan-Jaworowska, Cifelli, and Luo L. sp. 2004] Amblotherium gracilis Family Triconodontidae A. minimum Priacodon ferox A. sp. 1 ( = Kepolestes coloradensis) P. robustus A. sp. 2 P. lulli Gen. et sp. indet. P. grandaevus [Sources: Kielan-Jaworowska, Cifelli, and P. fruitaensis Luo 2004] Trioracodon bisulcus Family Paurodontidae Gen et sp. indet. Paurodon valens [Sources: Engelmann and Callison 1999; Archaeotrigon brevimaxillus Kielan-Jaworowska, Cifelli, and Luo 2004] 238 A. distagmus Latochara bellatula Tathiodon agilis L. collina Araeodon intermissus L. concinna Comotherium richi L. latitruncata Euthlastus cordiformis L. aff. L. mensinki Foxraptor atrox L. spherica New paurodontid under study by Rougier Mesochara voluta (K. Carpenter personal Obtusochara oblata communication 2006) Peckinspaera verticillata [Sources: Kielan-Jaworowska, Cifelli, and P. glypta Luo 2004] Praechara voluta P. symmetrica Order incertae sedis Porochara arguta Family incertae sedis P. fusca Fruitafossor windscheffeli P. kimmeridgensis [Source: Luo and Wible 2005] P. minima collina KINGDOM FUNGI Sphaerochara verticillata Stellatochara obovata DIVISION EUMYCOTA [Sources: Litwin, Turner, and Peterson 1998; Ott 1958; Peck 1956; Ross 1960; CLASS ASCOMYCETES Schudack 1995, 1998] Family incertae sedis unidentified remains KINGDOM PLANTAE [Sources: Tidwell 1990; Tidwell, Britt, and Ash 1998; Tidwell, Medlyn, and Hartmann DIVISION BRYOPHYTA 1980; Ash and Tidwell 1998] CLASS HEPATICAE CLASS BASIDIOMYCETES Family Marchantiaceae Family incertae sedis Marchantites sp. unidentified remains Thallites? sp. [Sources: Tidwell 1990a; Tidwell, Medlyn [Sources: Brown 1972; Ash and Tidwell and Hartmann, 1980; Tidwell, Britt, and 1998] Ash 1998; Ash and Tidwell 1998] DIVISION TRACHEOPHYTA CLASS incertae sedis Osteophagous fungal hyphae in dinosaur CLASS SPHENOPSIDA bone and ossified tendons Order Equisetales [Sources: Britt, Scheetz, Stadtman, and Family Equisitaceae Chure 2003] Equisteum laterale E. bakkeri KINGDOM PROTISTA E. comoensis E. cf. burchardtii DIVISION CHLOROPHYTA E. sp. CLASS CHLOROPHYCEAE Neocalamites sp. (W.D. Tidwell, personal Family Dictyospheriaceae comm., 1997) Botryoccus sp. [Sources: Tidwell 1990a; Tidwell, Britt, and [Sources: K. Carpenter, personal comm., Ash 1998; Ash and Tidwell 1998; Tidwell, 1997] Connely, and Britt 2005] CLASS CHAROPHYCEAE CLASS FILICOPSIDA Family Clavatoraceae Family Dipteridaceae Echinocara pecki Hausmania fisheri E. spinosa [Sources: Brown 1972; Ash and Tidwell [Sources: Ott 1958; Peck 1956; Schudack 1998] 1995] Family Dicksoniaceae Family Characeae Coniopteris hymenophylloides Aclistochara bransoni [Sources: Brown 1972; Ash and Tidwell A. complanata 1998] A. jonesi Family Osmundaceae A. latisulcata Osmundacaulis lemonii A. madleri Ashicaulis wadei A. miniscula Cladophlebis alberta A. obovata C. heterophylla A. rotunda C. virginiensis Chara verticillata [Sources: Brown 1972; Tidwell 1990b, C. voluta Ash and Tidwell 1998] 239 Family Gleicheniaceae 1990a; Tidwell, Britt, Gleichenia? sp. A and Ash 1998, Ash and Tidwell 1998] Gleichenia? sp. B Order Czekanowskiales [Sources: Tidwell, Connely, and Britt Family Czekanowskiaceae 2005] Czekanowskia turneri Family incertae sedis C. sp. Adiantites montanensis [Sources: Ash 1994; Tidwell, Britt, and Ash Solenostelopteris leithii 1998, Ash and Tidwell 1998] S. medlynii [Sources: Brown 1972; Miller 1987; CLASS CONIFEROPHYTA Tidwell 1990a; Ash and Tidwell 1998, Order Coniferales Tidwell and Skog 1999] Family Taxodiaceae? Family incertae sedis Hillostrobus axelrodi Fern sporangia Sequoia sp. [Sources: Chin and Carpenter 1998] “Other taxodiaceous cones” (Ash and Tidwell 1998) CLASS PTERIDOSPERMOPHYTA [Sources: Chandler 1966; Tidwell 1990a; Order Caytoniales Ash and Tidwell 1998] Family Caytoniaceae Family Cheirolepidiaceae? Sagenopteris elliptica Brachyphyllum rechtenii [Sources: Brown 1972; Tidwell 1990; Ash B. chaneyi and Tidwell 1998] B. morrisonense CLASS CYCADOPSIDA B. spp. (“2 possible species” Tidwell Order Cycadales 1990a) Family incertae sedis Cupressinocladus spp. Cycadinorachis sp. Protocupressinoxylon medlynii Jensensispermum redmondi [Sources: Francis 1983; Tidwell 1990a; Nilssonia nigricollensis Tidwell, Britt, and Ash 1998; Ash and N. schaumburgensis Tidwell 1998; Tidwell, Connely, and Britt N. cf. compta 2005] [Sources: Brown 1972; Chandler 1966; Family Araucariaceae Miller 1987; Tidwell 1990a; Ash and Araucarites obscurum Tidwell 1998] Araucarioxylon hoodi Podozamites lanceolatus Order Bennettitales [Sources: Tidwell 1990a; Wieland 1935; Family incertae sedis Tidwell and Medlyn 1993; Ash and Zamites arcticus Tidwell 1998] Otozamites sp. Family Podocarpaceae? Cycadolepsis spp. (“3 forms” Tidwell Xenoxylon morrisonense 1990a, Tidwell, Britt, and Ash 1998). X. mooreii Weltrichia sp. [Sources: Medlyn and Tidwell 1975, Anomozamites sp. Tidwell 1990a; Tidwell, Britt, and Ash Ptilophyllum lindstromi 1998] P. arcticum Family Protopinaceae P. spp. (“2 species” Tidwell 1990a) Protopiceoxylon resiniferous Cycadeoidea wyomingensis Mesembioxylon obscurum C. spp. M. carterii Cycadinorachis sp. Protocupressinoxylon medlynii Jensensispermum redmondii [Sources: Medlyn and Tidwell 1979, Cycadospermum montanense 1992; Tidwell, Britt, and Ash 1998; Ash [Sources: Brown 1972; Delevoryas 1960; and Tidwell 1998] Miller 1987; Tidwell 1990a; Tidwell, Family Pinaceae? Britt, and Ash 1998; Ash and Tidwell Pityocladus sp. 1998] Pityophyllum lindstromii Unidentified Cycadophytes Schizolepis sp. Cycadophyte petiole [Sources: Miller 1987; Tidwell 1990a; Cycadophyte laminae Ash and Tidwell 1998] [Sources: Chin and Kirkland 1998] Family Cupressaceae Order Ginkgoales Elatides sp. Family Ginkgoaceae [Sources: Tidwell, Britt, and Ash 1998] Ginkgoites cascandensis Family incertae sedis: leaf genera Ginkgo pluripartita Pagiophyllum sp. G. sp. “Possible male cones attached to [Sources: Holt 1940; Brown 1975; Tidwell 240 Pagiophyllum-type foliage DIVISION BRYOPHYTA [Sources: Brown 1972, Tidwell 1990a; Ash and Tidwell 1998] CLASS HEPATICAE Family incertae sedis: short shoots Family Sphagnaceae Behuninia provoensis Stereisporites cf. S. clavus B. joannei [Sources: Dodson, Behrensmeyer, and B. bassii Bakker 1980] B. scottii Family incertae sedis Carpolithus sp. Couperisporites jurassicus Hillostrobus axelrodii Foraminisporis sp. Steinerocaulis radiatus Staplinisporites caminus [Sources: Chandler 1966; Tidwell 1990a; [Sources: Tschudy, Tschudy, and Loenen Tidwell and Medlyn 1992; Ash and 1980, 1981, 1988a] Tidwell 1998; Tidwell, Britt, and Ash DIVISION TRACHEOPHYTA 1998] Family incertae sedis CLASS LYCOPSIDA Conites sp. Family Lycopodiaceae [Source: Tidwell, Britt, and Ash 1998] Lycopodium cf. L. crassimacerius Lycopodiacidites baculatus DIVISION TRACHEOPHYTA: CLASS incertae sedis Lycopodiumsporites austroclavitidites Order Hermanophytales L. crassiamcerius Family unnamed L. circolumenus Hermanophyton kirkbyorum L. cf. L. cerniidites H. taylorii Retritiletes concavus H. glismannii [Sources: Dodson, Behrensmeyer, and H. owensii Bakker 1980; Tschudy , Tschudy, and [Sources: Tidwell 1990a; Tidwell and Ash Loenen 1980, 1981, 1988a] 1990] CLASS FILICOPSIDA Order incertae sedis Family Dipteridaceae Families incertae sedis Dictyophyllidites harrissi Thorn (?) [Sources: Dodson, Behrensmeyer, and “Literally thouands of seeds from Bakker 1980] the Morrison Formation repre- Family Cyatheaceae sent either conifers, Cyathidites australis ginkgophytes,..members of the C. minor Cycadophyta....some of these [Sources: Tschudy, Tschudy, and Loenen specimens may also belong to the 1980, 1981, 1988a] seed ferns as well.” Tidwell Family Polypodiaceae 1990: p. 5 Seeds (at least 4 types) Laevigatosporites gracilis (Chin and Tidwell 1998) L. ovatus Cuticle [Sources: Dodson, Behrensmeyer, and Pycnoxylic secondary wood Bakker 1980; Tschudy, Tschudy, and [Sources: Chandler 1966; Tidwell Loenen 1980] 1990a;Chin and Kirkland 1998; Tidwell Family Osmundaceae 1998] Acanthotriletes midwayensis PALYNOMORPHS Baculatisporites comaumensis Biretisporites modestus The Morrison Formation has traditionally been viewed as being Leptolepidites major poor in palynomorphs. However, Litwin, Turner, and Peterson (1998: 313) L. psarosus report the recognition of more than 225 pollen and spore types in their Osmundacidites wellmani study of Morrison palynomorphs. A detailed study of that material has yet Todisporites minor to be published. [Sources: Dodson, Behrensmeyer, and KINGDOM RHODOPHYTA Bakker 1980; Tschudy, Tschudy, and Loenen 1980, 1981] CLASS incertae sedis Family Matoniaceae Scriniodinium crystallinum Concavisporites montuosus Wanea clathrata C. subgranulosi W. spectabilis C. variverrucatus [Sources: Litwin, Turner, and Peterson [Sources: Dodson, Behrensmeyer, and 1998] Bakker 1980] Family Gleichenaceae KINGDOM PLANTAE Gleichiniidites senonicus 241 [Sources: Tschudy, Tschudy, and Loenen 1981, 1998a, b] 1988b] Order Cycadales or Bennetitales Family Gleichenaceae or Matoniaceae Monosulcatus carpenteri Deltoidospora halli M. minimus D. junctum [Sources: Dodson, Behrensmeyer, and D. psilostoma Bakker 1980; Tschudy, Tschudy, and [Sources: Dodson, Behrensmeyer, and Loenen 1980, 1988a] Bakker 1980; Tschudy, Tschudy, and Loenen 1980, 1988a] Order Bennetitales or Ginkgoales Family Schizaeaceae Clavatipollenites hughsii Ceratosporites coulinensis Ginkgocycadophytus nitidus C. cf cretaceous [Sources: Dodson, Behrensmeyer, and C. dettmanii Bakker 1980; Tschudy, Tschudy, and Concavissimisporites irroratus Loenen et al. 1980, 1988b] C. jurienensis Order Coniferales C. montuosus Family ?Taxodiaceae C. penolaensis Perinopollenites asaccatus C. punctatus P. elatoides Klukisporites pseudoreticulatus [Sources: Dodson, Behrensmeyer, and Lygodiois sp. Bakker 1980; Tschudy, Tschudy, and Lygodiosporites perverrucatus Loenen 1980, 1988a] Schizosporis sp. Family Cheirolepidiaceae Trilobosporites jurassicus Classopoliis ituensis [Sources: Dodson, Behrensmeyer, and Corollina sp. Bakker 1980; Tschudy, Tschudy, and [Sources: Tschudy, Tschudy, and Loenen Loenen 1980, 1988a] 1980, 1988a] Family Salviniaceaea or Marseiliaceae Family Araucariaceae Crybelosporites vectensis Araucariacites australis [Sources: Tschudy, Tschudy, and Loenen A. fissus 1980, 1988a] [Sources: Dodson, Behrensmeyer, Family Selaginellaceae and Bakker 1980; Tschudy, Tschudy, Heliosporites cf. altmarkensis and Loenen 1981, 1988a] [Sources: Tscuhdy, Tschudy, and Loenen Family Araucariaceae or Cupressidaceae 1988b] Spheripollenites psilatus Family incertae sedis S. scabratus Coverrucosisporites proxiganulatus S. subgranulatus C. utriculosus [Sources: Dodson, Behrensmeyer, and Januasporites reticularis Bakker 1980] [Sources: Dodson, Behrensmeyer, and Bakker 1980; Tschudy, Tschudy, and Family ?Podocarpaceae Loenen 1980] Callialasporites crenulatus C. dampieri CLASS GYMOSPERMOPSIDA C. rugulatus Order Caytoniales C. segmentatus Family Caytoniaceae C. trilobatus Vitreisporites pallidus C. turbatus [Sources: Tschudy, Tschudy, and Loenen Podocarpidites cf. P. ellipticus 1981, 1988a, b] P. cf. P. granulatus P. langii Order Cycadales P. multesimus Family Cycadaeae P. cf. P. multicinus Cycadopites carpenteri P. rousei C. fragilis P. unicus C. nitidus [Sources: Dodson, Behrensmeyer, and C. parvus Bakker 1980; Tschudy, Tschudy, and Eucommiidites troedssonii Loenen 1981, 1988a, b] [Sources: Tschudy, Tschudy, and Loenen Family Pinaceae 1981. 1988a, b] Inapperturopollenites australis Order Bennetitales I. cf. I. australis Family incertae sedis I. dubius Exesipollintes scabratus I. magnus E. tumulus I. turbatus [Sources: Tschudy, Tschudy, and Loenen Pityoproties divulgatus 242 P. nigraeformis [Sources: Dodson, Behrensmeyer, and Bakker 1980; Litwin, Turner, [Sources: Dodson, Behrensmeyer, and and Peterson 1998; Tschudy, Tschudy, and Loenen 1980, 1981, 1988a, Bakker 1980; Tschudy, Tschudy, and 1988b] Loenen 1981, 1988a, b] Family incertae sedis II. TRACE FOSSILS Alisporites bilateralis A. grandis KINGDOM incertae sedis A. lowoodensis A. microsaccus STROMATOLITES A. thomasii cf. Rivularis sp. Cedripites canadensis “algal balls” C. cf. C. cretaceous “algal biscuts” [Sources: Dodson, Behrensmeyer, and [Sources: Lockley, Houck, and Prince 1986; Neuhauser, Lucas, Stephen Bakker 1980; Tschudy, Tschudy, and de Albuquerque, Hayden, Kietzke, Oakes, and Des Marais, Loenen 1980, 1981, 1988a, b] 1987; Hasiotis 2004] Order Gnetales KINGDOM FUNGI Family Ephedraceae Equisetosporites sp. DIVISION incertae sedis [Sources: Dodson, Behrensmeyer, and “Numerous bore holes in tracheids of these fossil woods are very similar Bakker 1980; Tschudy, Tschudy, and to the holes produced by hyphae of the white rot fungus Polyporus.” Loenen 1980, 1981, 1988a, b] [Sources: Tidwell 1990a: p. 7) CLASS GYMONSPERMOPSIDA incertae sedis Cerebropollenites macroverrucosus Wood rotting fungi present in Mesembioxylon carteri, C. mesozoicus Protocupressinoxylon medlynii, and Xenoxylon, also short shoots, and [Sources: Dodson, other species of fungi in species of Solenosteleopteris. Behrensmeyer, and Bakker 1980; [Sources: Tidwell, Britt, and Ash 1998; Tidwell and Medlyn 1992; Tschudy, Tschudy, and Loenen 1980, Tidwell and Skogg 1999)] 1981, 1988a, b] PALYNOMORPHS OF UNCERTAIN AFFINITY “Small to large cavities 1-5 cm in diameter, which occur in the heart- wood of permineralized coniferous wood.” Aequitriradites acusus [Sources: Hasiotis 2004] cf. Anapiculatisporites dawsonensis Appendicisporites sp. KINGDOM PLANTAE Cadargasporites reticulatus Cicatricosisporites sp. A DIVISION TRACHEOPHYTA Circulisporites sp. Coptospora sp. Rhizoliths Crassitudisporites problematicus Equisetites rhizoliths Densoisporites velatus [Sources: Chin and Kirkland 1998; Hasiotis and Demko 1998; Hasiotis Distalanulisporites spurius 2002, 2004] Distaltriangulisporis sp. Interulobites sp. KINGDOM ANIMALIA Ischyosporites disjunctus PHYLUM COELENTERATA Micocachrydites antarticus Murospora florida CLASS ANTHOZOA Neveisporites radiatus Family incertae sedis N. vallatus Conichnus isp. Obtusisporites canadensis [Sources: Hasiotis 2004] Paleoconiferus minor Polycingulatisporites reduncus PHYLUM BRACHIOPODA Pristinuspollenites microsaccus CLASS INARTICULATA P. microreticulatus Family incertae sedis cf. Protoconiferus funarius Lingulichnus isp. Protopicea exilioides [Sources: Hasiotis 2004] Pseudowalchia ovalis Psilatriletes circumundulatus PHYLUM MOLLUSCA Rubinella major CLASS GASTROPODA Rugubivesiculites woodbinensis Family incertae sedis Sphagnumsporites cooksoni Curvolithus isp. Crawling traces (S.T. Stereisporites cf. S. clavus Hasiotis, personal comm., 1997) Taurocusporites reduncus cf. Bergaueria isp. Resting traces (S.T. Undulatisporites sp. Hasiotis, personal comm., 1997) Veryhachium sp. 243 Borings in the upper surface of stromato- Vertical, bent Y-tube lites [Sources: Hasiotis and Demko 1998; Large patterned surface trail Hasiotis 2004] Elongate furrows Order Hemiptera Furrows with striations Family incertae sedis [Sources: Hasiotis 2004] Adhesive meniscate burrows CLASS BIVALVIA [Sources: Hasiotis 2002, 2004] Family Unionidae Dwelling/resting , locomotion, and escape Order Homoptera (cicadas) traces Family incertae sedis [Sources: Good 2004; Hasiotis 2004] Adhesive meniscate burrows Family incertae sedis Quasivertical striated burrows Pelecypodichnus amygdaloides (S.T. Horizontal, striated burrows Hasiotis, personal comm., 1997) [Sources: Hasiotis 2002, 2004] Lockeia isp. Borings in upper surface of stromatolites Order Diptera [Sources: Hasiotis 2002, 2004] Family Chironomidae Cochlichnus isp. PHYLUM NEMATODA Compressed U-shaped tubes with reinforced tops CLASS incertae sedis Ghost traces of U-shaped tubes Family incertae sedis Vertical, bent Y-tube Cochlichnus isp. [Sources: Hasiotis 2002, 2004] [Sources: Hasiotis 2002, 2004] Family Tipulidae PHYLUM ANNELIDA Oblique burrows (S.T. Hasiotis, personal comm., 1997) CLASS POLYCHAETA Horizontal striated burrows Family incertae sedis [Sources: Hasiotis 2004] Arenicolides isp. Family incertae sedis Cochlichnus isp. Scoyenia isp. Planolites isp. (S.T. Hasiotis, personal [Sources: Hasiotis 2004] comm. 1997) cf. Phycodes isp. Order Coleoptera Palaeophycus isp. Family Staphylinidae “Terebellina” isp. J-shaped dwelling burrows Pustulose marks [Sources: Hasitois 2004] [Sources: Hasiotis 2002, 2004] Family Carabiidae Adhesive meniscate burrows PHYLUM ARTHROPODA Horizontal striated burrows [Sources: Hasiotis 2004] CLASS MEROSTOMATA Family Scarabaenidae Order Xiphiosura Coprinisphaera isp. Family incertae sedis Adhesive meniscate burrows Kouphichnium isp. [Sources: Hasiotis 2002, 2004] [Sources: Hasiotis 2002, 2004 ] Family Cupedidae CLASS INSECTA Borings, including both feeding galleries Order Ephemeroptera and pupation cavities (S.T. Hasiotis, Family incertae sedis personal comm., 1997) Fuersichnus isp. Family Cicindelidae Compressed U-shaped tubes with Cylindricum antiquus reinforced tops [Sources: Hasiotis and Demko 1998; Ghost traces of U-shaped tubes Hasiotis 2004] Horizontal U-shaped tubes Family Heteroceridae Vertical, bent Y-tube Steinichnus isp., horizontal feeding/ [Sources: Hasiotis 2004] dwelling burrows [Sources: Hasiotis 2002, 2004] Order Orthoptera Family Buprestidae Family Gyrillidae Paleobuprestis isp. Horizontal, striated burrows [Sources: Hasiotis 2004] [Sources: Hasiotis 2004] Family Scolytidae Order Trichoptera Paleoscolytus isp. Family incertae sedis [Sources: Hasiotis 2004] Tektonargus kollaspilas Family Dermestidae Fuersichnus isp. Circular to elliptical borings in dinosaur 244 bone Rhizolith specific nest [Sources: Hasiotis 2004] Spherical structure Family incertae sedis Ramp Structure Concentrated galleries “Grub holes” in Behuninia [Sources: Hasiotis 2004] cf. Ancorichnus isp. Family incertae sedis Scoyenia isp. Smooth cavities in wood Smooth cavities in wood [Sources: Hasiotis 2004] [Sources: Tidwell and Medlyn 1992; Hasiotis 2004] CLASS CRUSTACEA Order Decapoda Order Hymenoptera Family Cambaridae Family Halictidae Camborygma araioklados Celliforma spirifera C. litonomos Rosellichnus isp. C. eumekenomos [Sources: Hasiotis and Demko 1998; [Sources: Hasiotis and Demko 1998; Hasiotis 2002, 2004] Hasiotis 2002, 2004] Family Sphecidae Family incertae sedis Spindle- to tablet- shaped cocoons (large Thalassinoides isp. (S.T. Hasiotis, and small) personal comm., 1997) Vertical tubes of variable dimensions Diplocraterion isp. (S.T. Hasiotis, [Sources: Hasiotis 2002, 2004] personal comm., 1997) Family Formicidae Scolicia isp. Interconnected polydomal chambers and Crab-like burrows (S.T. Hasiotis, personal galleries comm., 1997) Concentrated systems: hundreds to Large patterned surface trail thousands of chambers and [Sources: Hasiotis 2004] galleries Dispersed systems: dispersed polydomal PHYLUM ECHINODERMATA chambers and galleries CLASS ECHINOIDEA Low concentrated systems: 10-30 closely spaced chambers and galleries Family incertae sedis Smooth cavities in wood Scolicia isp. [Sources: Hasiotis 2002, [Sources: Hasiotis 2004] 2004] PHYLUM CHORDATA Order Isoptera CLASS REPTILIA Family Prototermitidae Order Testudines Large and deep nests Family incertae sedis Rhizolith engulfing nest Chelonipus isp. Rhizolith specific nest Unnamed turtle trackways Spherical structure [Sources: Foster, Lockley, and Brockett Ramp Structure 1999; Lockley and Foster 2006] Concentrated galleries [Sources: Hasiotis 2004] Order Sphenodontia Family Kalotermitidae Family incertae sedis Large and deep nests Single large-diameter, inclined burrows Rhizolith engulfing nest Single track Rhizolith specific nest [Sources: Hasiotis 2004; Lockley and Spherical structure Foster 2006] Ramp Structure Order Crocodilia Concentrated galleries Family incertae sedis [Sources: Hasiotis 2004] Hatcherichnus sanjuanensis Family Stolotermitidae Single large-diameter, inclined burrows Large and deep nests [Sources: Foster and Lockley 1997; Hasiotis Rhizolith engulfing nest 2004] Rhizolith specific nest Spherical structure Order Pterosauria Ramp Structure Family incertae sedis Concentrated galleries Pteraichnus saltwashensis [Sources: Hasiotis 2004] [Sources: Lockley, Foster, and Hunt Family Mastotermitidae 1998; Hasiotis 2002, 2004] Large and deep nests Order Saurischia Rhizolith engulfing nest Suborder Theropoda Family incertae sedis 245 ?Allosaurid track [Sources: Chin and Kirkland 1998] “Grallatorid” track CLASS MAMMALIA Unidentified small theropod footprints Family incertae sedis Unidentified “carnosaur” footprints Complex, large-diameter burrows Unnamed theropod tooth marks on [Sources: Hasiotis 2002, 2004; Hasiotis, sauropod bones have been Wellner, Martin, and Demko, 2004] attributed to theropods. [Sources: Bakker 1997; Lockley, Foster, FOSSIL EGGS and Hunt 1998; Harris 1998; Kvale et al. 2001; Hasiotis 2002, 2004; Foster and CLASS REPTILIA Lockley 2006; Platt and Hasiotis, 2006] Order Testudines Family incertae sedis Suborder Sauropoda ?Testudoid basic type Family incertae sedis Indeterminate testudoid Brontopodus sp. morphotype Parabrontopodus mcintoshi [Sources: Hirsch 1994; Bray and Hirsch Unidentified sauropod footprints at many 1998] localities [Sources: Lockley, Farlow, and Meyer Order Crocodylia 1994; Engelmann and Hasiotis, 1999; Family incertae sedis Kvale et al. 2001; Hasiotis 2004; Platt and ?Crocodyloid basic type Hasiotis, 2006] Indeterminate crocodyloid morphotype Order Ornithischia [Sources: Hirsch 1994; Zelentitsky and Hill Suborder Ornithopoda 1996; Bray and Hirsch 1998] Family incertae sedis cf. Anomoepus sp. Subclass Dinosauria Dinehichnus socialis Family Prismatoolithidae ?Gypsichnites sp. Preprismatoolithus coloradensis Footprints of ornithopods [Sources: Hirsch 1994; Bray and Hirsch Footprints of small ornithopods 1998] Footprints of large ornithopods Family incertae sedis [Sources: Lockley, Foster, and Hunt 1998; Dinosauroid-Prismatic Basic Type Lockley, Dos Santos, Meyer, and Hunt Oliquiprismatic morphotype 1998; Harris 1998; Kvale et al. 2001; type 2 Hasiotis 2004; Kvale et al. 2004; Platt and Indeterminate prismatic Hasiotis, 2006] morphotype type 1 Indeterminate prismatic Suborder Stegosauria morphotype type 2 Family incertae sedis Stegopodus czerkasi [Sources: Hirsch 1994; Bray and Hirsch [Sources: Lockley and Hunt 1998] 1998] Suborder incertae sedis Family incertae sedis Dinosauroid-Spherulitic Basic Type Herbivorous dinosaur coprolites Tubospherulitic morphotype Indeterminate morphhotype [Sources: Hirsch 1994; Bray and Hirsch REFERENCES 1998]
Ash, S., 1994, First occurrence of Czekanowskia (Gymnospermae, Czekanowskiales) the highest levels of the Morrison Formation at Como Bluff, Wyoming, with in the United States: Review of Palaeobotany and Palynology v. 81, p. 129-140. comments on Morrison Biochronology. Part II. The mammals: a new Ash, S.R., and Tidwell, W.D., 1998, Plant megafossils from the Brushy Basin Mem- multituberculate and a new paurodont: Hunteria v.2, no. 6, p 4-8, 10-14. ber of the Morrison Formation near Montezuma Creek Trading Post, southeast- Branson, C.C., 1935, Fresh-water invertebrates from the Morrison Formation (Ju- ern Utah. in Carpenter, K., Chure, D.J., and Kirkland, J.I., eds., The Upper Ju- rassic?) of Wyoming: Journal of Paleontology, v.9, no. 6, p. 514-522. rassic Morrison Formation: An Interdisciplinary Study. Modern Geology v. 22, Bray, E., and Hirsch, K., 1998, Eggshell from the Upper Jurassic Morrison Forma- p. 321-340. tion. in Carpenter, K., Chure, D.J., and Kirkland, J.I., eds., The Morrison For- Bakker, R.T., 1997, Raptor family values: allosaur parents brought giant carcasses mation: An Interdisciplinary Approach. Modern Geology, v. 23, p. 219-240. into their lair to feed their young: in Wolberg, D.L., Stump, E., and Rosenberg, Britt, B.B., 1991, Theropods of the Dry Mesa Quarry (Morrison Formation, Late G.D., eds., DinoFest International. Proceedings of a Symposium sponsored by Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri: Arizona State University. Philadelphia, The Academy of Natural Sciences, p. Brigham Young University Geology Studies, v. 37, p. 1-72. 51-63. Britt, B.B, Scheetz, R., Stadtman, K., and Chure, D.J., 2003, Relicts of soft tissue Bakker, R.T., 1998, Dinosaur mid-life crisis: the Jurassic-Cretaceous transition in and osteophagous fungi in partially ossified tendons of Ceratosaurus (Theropoda, Wyoming and Colorado: in Lucas, S.G., Kirkland, J.I., and Estep, J.W., eds., Dinosauria) (abs.): Journal of Vertebrate Paleontology, v. 23, (supplement to Lower and Middle Cretaceous Terrestrial Ecosystems. Albuquerque, New Mexico no. 3), p. 36A. Museum of Natural History and Science Bulletin 14, p. 66-77. Brown, J.T., 1972, The flora of the Morrison Formation (Upper Jurassic) of central Bakker, R.T., and Carpenter, K., 1990, A new latest Jurassic vertebrate fauna, from Montana, [Ph.D. Dissertation]: Missoula, University of Montana, 77 p. 246 Brown, J.T., 1975, Upper Jurassic and Lower Cretaceous Ginkgophytes from Mon- Engelmann, G.F. and Hasiotis, S.T., 1999, Deep dinosaur tracks in the Morrison: tana: Journal of Paleontology, v. 49, p.724-730. sole marks that are really sole marks: in Gillette, D. D., ed., Vertebrate Paleon- Callison, G., 1987, Fruita: a place for wee fossils: in Averett, W.R., ed., 1987, Pale- tology in Utah: Utah Geological Survey, Miscellaneous Publication, 99-1, p. ontology and Geology of the Dinosaur Triangle, Grand Junction, The Museum 179-183. of Western Colorado, p. 91-96. Evanoff, E., Good, S., and Hanley, J., 1998, An overview of the freshwater mollusks Carpenter, K., and Bakker, R.T., 1990, A new latest Jurassic vertebrate fauna, from from the Morrison Formation (Upper Jurassic, Western Interior, USA): in Car- the highest levels of the Morrison Formation at Como Bluff, Wyoming, with penter, K.E., Chure, D.J., and Kirkland, J.I., eds., The Morrison Symposium: comments on Morrison Biochronology. Part II. A new baenid turtle: Hunteria, v. An Interdisciplinary Approach: Modern Geology, v. 22, p. 423-450. 2, no. 6, p. 3-4, 6-7. Evans, S.E., 1990, The skull of Cteniogenys, a choristodere (Reptilia: Carpenter, K., Miles, C., and Cloward, K., 2005, New small theropod from the Archosauromorpha) from the Middle Jurassic of Oxfordshire: Zoological Jour- Upper Jurassic Morrison Formation of Wyoming: in Carpenter, K., ed., The nal of the Linnean Society, v. 99, p. 205-237. Carnivorous Dinosaurs: Bloomington and Indianapolis, Indiana University Press, Evans, S.E., 1996, Parviraptor (Squamata: Anguimorpha) and other lizards from p. 23-48. the Morrison Formation at Fruita, Colorado: in Morales, M., ed., The Continen- Carpenter, K., Unwin, D., Cloward, K., Miles, C., and Miles, C., 2003, A new tal Jurassic, Transactions of the Continental Jurassic Symposium. Flagstaff, scaphognathine like pterosaur from the Upper Jurassic Morrison Formation of Museum of Northern Arizona Bulletin 60, p. 243-248. Wyoming, USA: in Buffetaut, E., and Mazin, J.-M., eds., Evolution and Evans, S.E., and Chure, D.J., 1998a, Morrison lizards: structure, relationships, and Paleobiology of Pterosaurs: Geological Society of America Special Publication biogeography: in Carpenter, K., Chure, D.J., and Kirkland, J.I., eds., The Upper no. 217, p. 45-54. Jurassic Morrison Formation: An Interdisciplinary Study: Modern Geology, v. Chandler, M.E.J., 1966, Fruiting organs from the Morrison Formation of Utah, USA: 23, p. 35-48. Bulletin of the British Museum (Natural History) Geology, v. 12, no. 4, p. 139- Evans, S. E., and Chure, D.J., 1998b., Paramacellodid lizard skulls from the Juras- 171 + plates 1-12 sic Morrison Formation at Dinosaur National Monument, Utah: Journal of Ver- Chin, K., and Kirkland, J.I., 1998, Probable herbivore coprolites from the Upper tebrate Paleontology, v. 18, p. 99-114. Jurassic Mygatt-Moore Quarry, western Colorado: in Carpenter, K., Chure, D.J., Evans, S.E., and Chure, D.J., 1999, Upper Jurassic lizards from the Morrison For- and Kirkland, J.I., eds., The Upper Jurassic Morrison Formation: An Interdisci- mation of Dinosaur National Monument, UT: in Gillette, D.D., ed., Vertebrate plinary Study: Modern Geology, v. 23, p. 249-275. Paleontology in Utah: Utah Geological Survey, Miscellaneous Publication 99- Chure, D.J., 1994, Koparion douglassi, a new dinosaur from the Morrison Forma- 1, p. 151-159. tion (Upper Jurassic) of Dinosaur National Monument; the oldest troodontid Evans, S.E., Lally, C., Chure, D.J., Elder, A., and Maisano, J.A., 2005, A Late Ju- (Theropoda: Maniraptora): Brigham Young University Geology Studies, v. 40, rassic salamander (Amphibia: Caudata) from the Morrison Formation of North p. 11-15. America: Zoological Journal of the Linnean Society, v. 143, p. 599-616. Chure, D.J., 1995, A reassessment of the gigantic theropod Saurophagus maximus Evans, S.E., and Milner, A.R., 1993, Frogs and salamanders from the Upper Juras- from the Morrison Formation (Upper Jurassic) of Oklahoma, USA: in Sun, L., sic Morrison Formation (Quarry Nine, Como Bluff) of North America: Journal and Wang, Y.Q., eds., Sixth Symposium on Mesozoic Terrestrial Ecosystems of Vertebrate Paleontology, v. 13, p. 24-30. and Biota, Short Papers: Beijing, China Ocean Press, p. 103-106. Foster, J.R., 2003, Paleoecological analysis of the vertebrate fauna of the Morrison Chure, D.J., 2000, A New Species of Allosaurus from the Morrison Formation, Formation (Upper Jurassic), Rocky Mountain Region, U.S.A.: Albuquerque, Dinosaur National Monument (UT-CO) and a Revision of the Theropod Family New Mexico Museum of Natural History and Science Bulletin 23, 95 p. Allosauridae [Ph.D. Dissertation]: New York, Columbia University, 900 p. Foster, J.R., and Lockley, M.G., 1997, Probable crocodilian tracks and traces from Chure, D.J., Carpenter, K., Litwin, R., Hasiotis, S., and Evanoff, E., 1998, The flora the Morrison Formation (Upper Jurassic) of eastern Utah: Ichnos, v. 5, p. 121- and fauna of the Morrison Formation: in Carpenter, K., Chure, D.J., and Kirkland, 129. J.I., eds., The Upper Jurassic Morrison Formation: An Interdisciplinary Study: Foster, J.R., and Lockley, M.G., 2006, The vertebrate ichnological record of the Modern Geology, v. 23, p. 507-537. Morrison Formation (Upper Jurassic, North America), in Foster, J.R., and Lucas, Chure, D.J., Madsen, S.K., and Armstrong, H.J., 1991, An unusual mesosuchian S. G., eds., Paleontology and Geology of the Upper Jurassic Morrison Forma- crocodilian from the Salt Wash Member of the Morrison Formation, Upper Ju- tion: New Mexico Museum of Natural History and Science, Bulletin 36, this rassic, western Colorado (ABS.): Journal of Vertebrate Paleontology, v. 11, volume. supplement to number 3, 22A-23A. Foster, J.R., Lockley, M.G., and Brockett, J., 1999, Possible turtle tracks from the Clark, J.A., 1985, A new crocodylomorph from the Late Jurassic Morrison Forma- Morrison Formation of southern Utah: in Gillette, D.D., ed., Vertebrate Paleon- tion of western Colorado, with a discussion of relationships within the tology in Utah: Utah Geological Survey, Miscellaneous Publication 99-1, p. “Mesosuchia” [Unpublished M.S. Thesis]: Berkeley, University of California, 186-191. 102 p. Francis, E., 1983, The dominant conifer of the Jurassic Purbeck Formation, En- Czerkas, S.A., and Mickelson, D.L., 2002, The first occurrence of skeletal ptero- gland: Palaeontology, v. 26, p. 277-294. saur remains in Utah: in Czerkas, S.J., ed., Feathered Dinosaurs and the Origin Gaffey, E.S., 1979, The Jurassic turtles of North America: American Museum of of Flight: Blanding, The Dinosaur Museum Journal, v. 1, p. 2-13. Natural History, Bulletin, v.162, p. 91-136. Delevoryas, T., 1960, Investigations of North American cycadeoids: trunks from Galton, P.M., 1982, Elaphrosaurus, an ornithomimid dinosaur from the Upper Ju- Wyoming: American Journal of Botany, v. 47, p.778-786. rassic of North America and Africa: Palaeontologica Zetischrift, v. 56, p. 265- Dodson, P., Behrensmeyer, A.K., and Bakker, R.T., 1980, Taphonomy of the Morrison 272. Formation (Kimmeridgian-Portlandian) and Cloverly Formation (Aptian-Albian) Galton, P.M., 1990, Stegosauria: in Weishampel, D.B., Dodson, P., and Osmolska, of the western United States: Memoires de la Societe geologique de France (N.S.), H., eds., The Dinosauria (1st edition): Berkeley, University of California Press, Tome 49, (Memoire no. 139), p. 87-93. p. 435-455. Dunagan, S.P., 1999, A North American freshwater sponge (Eospongilla Galton, P.M., 2002, New material of ornithischian (?heterodontodsaurid) dinosaur morrisonensis new genus and species) from the Morrison Formation (Upper Echinodon (Early Cretaceous, southern England) from the Late Jurassic of Fruita Jurassic), Colorado: Journal of Paleontology, v. 73, p. 389-393. near Grand Junction, Colorado, USA (abs.): Journal of Vertebrate Paleontology, Engelmann, G.F., and Callison, G., 1998 Mammalian faunas of the Morrison For- v. 22, supplement to no. 3, p. 55A-56A. mation: in Carpenter, K.E., Chure, D.J. and Kirkland, J.I., eds., The Morrison Galton, P.M., in press, Teeth of ornithischian dinosaurs (mostly Ornithopoda) from Symposium: An Interdisciplinary Approach: Modern Geology, v. 23, p. 343- the Morrison Formation (Upper Jurassic) of western USA.: in: Carpenter, K., 380. ed., Horns and Beaks, Ceratopsian and Ornithopod Dinosaurs, Bloomington, Engelmann, G.F., and Callison, G., 1999, Glirodon grandis, a new multituberculate University of Indiana Press mammal from the Upper Jurassic Morrison Formation: in Gillette, D.D., ed., Gillette, D.D., 1991, Seismosaurus halli, gen. et sp. nov., a new sauropod dinosaur Vertebrate Paleontology in Utah: Utah Geological Survey, Miscellaneous Pub- from the Morrison Formation (Upper Jurassic / Lower Cretaceous) of New lication, 99-1, p. 161-177. Mexico, U.S.A.: Journal of Vertebrate Paleontology, v.11, p. 417-433. 247 Gillette, D.D., 1996, Stratigraphic position of the sauropod dinosaur Dystrophaeus D.B., Dodson, P., and Osmolska, H., eds., The Dinosauria: Berkeley, University viaemalae Cope 1877 and its evolutionary implications: in Morales, M., ed., of California Press, (2nd edition), p. 71-110. The Continental Jurassic, Transactions of the Continental Jurassic Symposium. Jensen, J.A., and Padian, K., 1989, Small pterosaurs and dinosaurs from the Flagstaff, Museum of Northern Arizona Bulletin No. 60, p.59-68. Uncompahgre Fauna (Brushy Basin Member, Morrison Formation: ?Tithonian), Gilmore, C.W., 1909, A new rhynchocephalian reptile from the Jurassic of Wyo- Late Jurassic, western Colorado: Journal of Paleontology, v. 63, p. 364-373. ming, with notes on the fauna of Quarry Nine: United States National Museum Kielan-Jaworowska, Z., Cifelli, R.L., and Luo, Z.-X., 2004, Mammals from the Proceedings, v. 37, p. 35-42 + plate 11. Age of Dinosaurs: Origins, Evolution, and Structure: New York, Columbia Gilmore, C.W., 1926, A new aetosaurian reptile from the Morrison Formation of University Press, xv + 630 p. Utah: Annals of the Carnegie Museum, v. XVI, p. 326-342 + plate xxvii. Kirkland, J.I., 1987, Upper Jurassic and Cretaceous lungfish toothplates from the Gilmore, C.W., 1928, Fossil lizards of North America; U.S. National Academy of Western Interior, the last dipnoan faunas of North America: Hunteria v. 2, 16 p. Sciences, v. XXII, third memoir, 201 p. Kirkland, J.I., 1998, Morrison fishes, in, Carpenter, K., Chure, D.J., and Kirkland, Göhlich, U.B., Chiappe, L.M., Clark, J.M., and Sues, H.D., 2005, The systematic J.I., eds., The Upper Jurassic Morrison Formation: An Interdisciplinary Study: position of the Late Jurassic alleged dinosaur Macelognathus Modern Geology, v. 22, p. 503-533. (Crocodylomorpha: Sphenosuchia): Canadian Journal of Earth Science, v. 42, Kirkland, J.I., and Armstrong, H.J., 1992, Taphonomy of the Mygatt-Moore (M and p. 307-321. M) Quarry, middle Brushy Basin Member, Morrison Formation (Upper Juras- Good, S.C., 2004, Paleoenvironmental and paleoclimatic significance of freshwater sic) Western, Colorado (abs.): Journal of Vertebrate Paleontology, v. 12 (supple- bivalves in the Upper Jurassic Formation, Western Interior, USA: Sedimentary ment to no. 3), p. 37A. Geology, v. 167, p.163-176. Kirkland, J.I., and Carpenter, K., 1994, North America’s first pre-Cretaceous Harris, J.D., 1998, Dinosaur footprints from Garden Park, Colorado: in Carpenter, ankylosaur (Dinosauria) from the Upper Jurassic Morrison Formation of West- K.E., Chure, D.J., and Kirkland, J.I., eds., The Morrison Symposium: An Inter- ern Colorado: Brigham Young University Geology Studies, v. 40, p. 25-42. disciplinary Approach: Modern Geology, v. 23, p. 291-307. Kvale, E.P., Hasiotis, S.T., Mickelson, D., and Johnson, G.D., 2001, Middle and Harris, J.D., and Carpenter, K., 1996, A large pterodactyloid from the Morrison Late Jurassic Dinosaur fossil-bearing horizons: Implications for dinosaur pa- Formation (Late Jurassic) of Garden Park, Colorado: Neues Jahrbuch der leoecology, northeastern Bighorn Basin, Wyoming: Society of Vertebrate Pale- Geologie, Paläontologie, Montash 1996, p. 473-484. ontology Annual Meeting, Bozeman, Montana, Field Trip Guidebook, Museum Harris, J.D., and Dodson, P., 2004, A new diplodocoid sauropod dinosaur from the of the Rockies Occasional Paper No. 3, p. 17-45. Upper Jurassic Morrison Formation of Montana, USA: Acta Palaeontologica Kvale, E.P., Hasiotis, S.T., Mickelson, D., and Johnson, G.D., 2004, The history of Polonica, v. 49, p. 197-210. dinosaur track discoveries in Wyoming with emphasis on the Bighorn Basin: Hartman, S., Lovelace, D., and Wahl, D., 2005, Phylogenetic assessment of a Ichnos, v. 11, p. 3-9. maniraptoran from the Morrison Formation: (ABS.) Journal of Vertebrate Pale- Litwin, R., Turner, C.E., and Peterson, F., 1998, Palynological evidence on the age ontology, v. 25, supplement to no. 3, p. 67A-68A. of the Morrison Formation, Western Interior U.S., in Carpenter, K.E., Chure, Hasiotis, S.T., 2000, The Invertebrate Invasion and Evolution of Mesozoic Soil D.J. and Kirkland, J.I., eds., The Morrison Symposium: An Interdisciplinary Ecosystems: The Ichnofossil Record of Ecological Innovations: in Gastaldo, R. Approach: Modern Geology, v. 22, p. 297-320. and Dimichele, W., eds., Phanerozoic Terrestrial Ecosystems, Paleontological Lockley, M.G., 1986, Dinosaur Tracksites, University of Colorado at Denver, De- Society Short Course, 6, p. 141-169. partment of Geology Magazine, Special issue no. 1, 56 p. Hasiotis, S.T., 2002, Continental Trace Fossils: Society of Economic Paleontolo- Lockley, M.G., Dos Santos, V.F., Meyer, C., and Hunt, A.P., 1998, A new dinosaur gists and Mineralogists, Short Course Notes Number 51, Tulsa, Oklahoma, 132 tracksite in the Morrison Formation Boundary Butte, southeastern Utah, in Car- p. penter, K.E., Chure, D.J,. and Kirkland, J.I., eds., The Morrison Symposium: Hasiotis, S.T., 2003, Complex ichnofossils of solitary to social soil organisms: un- An Interdisciplinary Approach: Modern Geology, v. 23, p. 317-330. derstanding their evolution and roles in terrestrial paleoecosystems: Lockley, M., Farlow, J.O., and Hunt, A.P., 1998, A short summary of dinosaur tracks Palaeogeography, Palaeoclimatology, Palaeoecology, v. 192, p. 259-320. and other fossil footprints from the Morrison Formation, in Carpenter, K.E., Hasiotis, S.T., 2004, Reconnaissance of Upper Jurassic Morrison Formation Chure, D.J., and Kirkland, J.I., eds., The Morrison Symposium: An Interdisci- ichnofossils, Rocky Mountain Region, USA: paleoenvironmental, stratigraphic, plinary Approach: Modern Geology, v. 23, p. 277-290. and paleoclimatic significance of terrestrial and freshwater ichnocoenoses: Sedi- Lockley, M.G., Farlow, J.O., and Meyer, C., 1994, Brontopodus and mentary Geology, v. 167, p. 177-268. Parabrontopodous ichnogen. nov. and the significance of wide and narrow gauge Hasiotis, S.T., and Demko, T.M., 1998, Ichnofossils from Garden Park Paleonto- sauropod trackways: Gaia v.10, p. 135-145. logical Area, Colorado: Implications for paleoecologic and paleoclimatic Lockley, M.G., and Foster, J.R., 2006, Dinosaur and turtle tracks from the Morrison reconstructiuons of the Upper Jurassic: in Carpenter, K.E., Chure, D.J., and Formation Upper Jurassic of Colorado National Monument, with observations Kirkland, J.I., eds., The Morrison Symposium: An Interdisciplinary Approach: on the taxonomy of vertebrate swim tracks, in Foster, J.R., and Lucas, S. G., Modern Geology, v. 22, p. 461-479. eds., Paleontology and Geology of the Upper Jurassic Morrison Formation: New Hasiotis, S.T., Wellner, R.W., Martin, A., and Demko, T.M., 2004, Vertebrate bur- Mexico Museum of Natural History and Science, Bulletin 36, this volume. rows from Triassic and Jurassic continental deposits of North America and Ant- Lockley, M.G., Foster, J.R., and Hunt, A.P., 1998, A short summary of dinosaur arctica: their paleoenvironmental and paleoecological significance: Ichnos, v. tracks and other fossil footprints from the Morrison Formation, in Carpenter, 11, p. 103-124. K.E., Chure, D.J., and Kirkland, J.I., eds., The Morrison Symposium: An Inter- Hassinger, R.N., 1959, Stratigraphy of the Morrison Formation of the Canon City disciplinary Approach: Modern Geology, v. 23, p. 277-290. Embayment, Colorado [MS Thesis]: Norman, University of Oklahoma, Norman, Lockley, M.G., Houck, K.J., and Prince, N.K., 1986, North America’s largest dino- 116 p. saur trackway site: implications for Morrison Formation paleoecology: Geo- Hecht, M.K., and Estes, R., 1960, Fossil amphibians from Quarry Nine: Postilla no. logical Society of America Bulletin, v. 97, p. 1163-1176. 46, 19 p. Lockley, M.G. and Hunt, A.P., 1998, A probable stegosaur track from the Morrison Henrici, A.C., 1998, A new pipoid anuran from the Late Jurassic Morrison Forma- Formation of Utah, in Carpenter, K.E., Chure, D.J., and Kirkland, J.I., eds., The tion at Dinosaur National Monument, Utah: Journal of Vertebrate Paleontology, Morrison Symposium: An Interdisciplinary Approach: Modern Geology, v. 23, v. 18, p. 321-332. p. 331-342. Hirsch, K.F., 1994, Upper Jurassic eggshells from the Western Interior of North Lucas, S.G., and Kirkland, J.I., 1998, Preliminary report on Conchostraca from the America, in Carpenter. K., Hirsch, K.F., and Horner, J.R., eds., Dinosaur Eggs Morrison Formation, Western United States, in Carpenter, K.E., Chure, D.J., and Babies: Cambridge, Cambridge University Press, p. 137-150. and Kirkland, J.I., eds., The Morrison Symposium: An Interdisciplinary Ap- Holt, E., 1940, The Morrison and Summerville formations of the Grand River Val- proach: Modern Geology, v. 22, p. 415-422. ley and their vertebrate and invertebrate fauna [Master’s Thesis]: Denver, Uni- Luo, Z.-X., and Wible, J.R., 2005, A Late Jurassic digging mammal and early mam- versity of Colorado, 71 p. malian diversification: Science, v. 308, p.103-107 + 100 pages of on-line supple- Holtz, T.R., Molnar, R.E., and Currie, P.M. 2004, Basal Tetanurae, in, Weishampel, mentary information at
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