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Smithsonian Contributions to Paleobiology

Displaying 51 - 97 from the 97 total records
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Megaspores and a Palynomorph from the Lower Potomac Group in Virginia
Francis M. Hueber
69 pages, 1 figure, 24 plates
1982 (Date of Issue: 22 February 1982)
Number 49, Smithsonian Contributions to Paleobiology
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A plant microfossil assemblage comprising seven species of megaspores; Verrutriletes carbunculus (Dijkstra) Potonié, Echitriletes cf. E. lanatus (Dijkstra) Potonié, Erlansonisporites erlansonii (Miner) Potonié, Thylakosporites retiarius (Hughes) Potonié, Arcellites disciformis (Miner) Ellis and Tschudy, Arcellites cf. A. pyriformis (Dijkstra) Potter, and Paxillitriletes species Hall and Nicolson; two species of the microspore Crybelosporites Dettmann, C. striatus (Cookson and Dettmann) Dettmann adherent to specimens of Arcellites disciformis, and Crybelosporites species adherent to specimens of Echitriletes cf. E. lanatus; and the palynomorph Dictyothylakos pesslerae Horst; is recorded from the Patuxent Formation, Potomac Group, Lower Cretaceous (Barremian-Aptian) in Virginia, USA. A preliminary analysis of the enclosing matrix for microspores and pollen has related the collection site closely to lowermost Zone I of the Potomac Group as described by Hickey and Doyle (1977). The megaspore assemblage supported by acceptance of the oldest possible date derived from the microspore and pollen analysis suggests correlation with the Barremian-Aptian horizons in the English Wealden, Lower Cretaceous, and specifically with the “Arcellites Flora” of Hughes. Megafossils comprising two seed cones belonging to the Pinaceae, Pityostrobus hueberi Robison and Miller and Pityostrobus virginiana Robison and Miller have been reported from the site. A fruit or cupule of Caytonia has been found along with numerous seeds, fern fragments, coniferous woods, and cycadopsid cuticles. This array of megafossils is not described or illustrated herein. A backswamp area of sedimentation and type of habitat is suggested on the basis of the lithofacies and generalized composition of the flora. The writer fully agrees with Tschudy (1976) as to the importance of searching for megaspores in continental Mesozoic rocks to aid in correlating and subdividing the deposits more effectively.

The Terebratulacea (Brachiopoda), Triassic to Recent: A Study of the Brachidia (Loops)
G. Arthur Cooper
445 pages, 17 figures, 77 plates, 86 tables
1983 (Date of Issue: 3 October 1983)
Number 50, Smithsonian Contributions to Paleobiology
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The narrowly rostrate brachiopods of the Terebratulacea have long been a challenging subject. The interior details of many have long remained unknown. The usual method of studying the interior is by serial grinding (sectioning). Some workers are content with the information revealed by the sections; others give visual reality to the sections by reconstructing the loop and cardinalia. Noting the unlikely results of many of these reconstructions, a better way of revealing the interior suggested itself, i.e., revealing the loop by excavation of the matrix. This method, when matrix is workable, makes possible more accurate measurement and depiction of the loop. Making statistics of parts of the loop to use in comparing these structures in different genera is described. Certain characters and proportions of parts of the loop used in classifying genera are explained. External characters of generic importance are noted for each genus.

The systematics of 208 genera of short-looped brachiopods of which 70% have revealed their loops by excavation or by being silicified are considered along with revision of families and the making of additional categories. It is indicated, with evidence, that the type of the Loboidothyrididae (Loboidothyridacea of Makridin) of the Jurassic has a short loop without conspicuous terminal points (flanges) rather than a long, long-flanged loop. It is recommended that the family definition of the Loboidothyrididae be altered to suit the facts of the structure of its leading genus, and that the Loboidothyridacea be abandoned. The Lobothyrididae, supposedly with short-flanged loop, has a loop with fairly long terminal points, and is thus in need of revision.

It is suggested that Pseudodielasma of the Permian is stucturally a possible ancestor of Terebratulidae with abbreviated terminal points. The origin of the Loboidothyrididae (now Tchegemithyrididae) is postulated to be from the Triassic family Angustithyrididae of the Dielasmatacea according to Dagis (1974). The Permian Ectoposia is a possible ancestor of the Angustithyrididae. Available collections were inadequate for the study of the loop development of any genera except living ones. Information on the development of a Mesozoic long-flanged loop is known from the work of Dagis (1968) on Viligothyris.

All known genera of the Terebratulacea were studied either by original preparations of the loop or from the literature. Lack of adequate collections prevented the detailed study of the Pygopidae. Fifty-eight genera of Jurassic brachiopods were developed to show the loop, fifteen of them Buckman's hitherto poorly known genera. An additional four unplaced genera were developed. Thirty-six Cretaceous genera revealed the loop, and fourteen genera of Tertiary Terebratulaceans are known from excavated loops. Forty-seven genera of short-looped brachiopods are known from description and serial sections only and without reconstruction of the loop. The loops of seven genera are unknown. Illustrations of all prepared genera are presented, and in addition, some reconstructions from serial section in the literature. Diagrammatic drawings of important loops of dissected specimens are illustrated. It is concluded that preparation of the loop, when possible, is preferred and is more accurate and less time consuming than reconstruction from serial grinding (section).

The Relationships of Megaoryzomys curioi, an Extinct Cricetine Rodent (Muroidea: Muridae) from the Galápagos Islands, Ecuador
David W. Steadman and Clayton E. Ray
23 pages, 11 figures, 1 table
1982 (Date of Issue: 24 August 1982)
Number 51, Smithsonian Contributions to Paleobiology
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Megaoryzomys curioi is a thomasomyine, not an oryzomyine as previously believed. This rodent was originally described, from three bony fragments found in a cave on Isla Santa Cruz, Galápagos, as a new species of the Antillean oryzomyine genus Megalomys. The genus Megaoryzomys was named recently for this species, based on new material from Isla Santa Cruz. Our study of additional material indicates that Megaoryzomys curioi is not closely related to Oryzomys but is most similar to large species of Thomasomys, a genus confined to mainland South America. The Galápagos have been colonized by cricetine rodents at least three times, once by a thomasomyine and twice by oryzomyines. Of these colonists, Megaoryzomys curioi is the most divergent from mainland relatives and thus is probably derived from the earliest immigrant. Although the time of extinction of Megaoryzomys curioi has not been determined, and it has never been recorded from life, it probably survived into historic time.

North American Eocene Sea Cows (Mammalia: Sirenia)
Daryl P. Domning, Gary S. Morgan and Clayton E. Ray
69 pages, 34 figures, 4 tables
1982 (Date of Issue: 3 September 1982)
Number 52, Smithsonian Contributions to Paleobiology
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The record of Eocene sea cows in North America is reviewed in detail, and that of the world is summarized. The North American record includes some 20 localities, mostly yielding fragments identifiable only as sirenian. Of these, the most extensive materials are a partial skeleton from the Cook Mountain Formation of Texas, numerous isolated elements from the Avon Park Formation of Florida, and a partial skeleton and other specimens from the Castle Hayne Formation of North Carolina. The materials from North Carolina and Florida are middle Eocene in age and are referred to Protosiren species. These specimens provide further confirmation of the fact that Eocene sirenians had a dental formula and were the latest eutherians known to exhibit five premolars. The implications of this for the higher classification of mammals are discussed. The distribution of sirenians suggests a homogeneous middle Eocene Tethyan fauna and also seems to be a more useful guide to the former distribution of seagrasses than are the distributions of Foraminifera. Eocene sirenians have potential value in intercontinental biostratigraphic correlation.

Geology and Paleontology of the Lee Creek Mine, North Carolina, I
Clayton E. Ray, editor
529 pages, 95 figures, 101 plates, 8 tables
1983 (Date of Issue: 13 September 1983)
Number 53, Smithsonian Contributions to Paleobiology
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This volume of papers on the geology and paleontology of the Lee Creek Mine is the first of three to be dedicated to the late Remington Kellogg, who initiated Smithsonian studies of the mine. It includes the first 14 papers, as well as a biography of Remington Kellogg by Frank C. Whitmore, Jr., and a prologue by Clayton E. Ray. This study places the Lee Creek Mine in the larger context of the history of Neogene geology and paleontology of the middle Atlantic Coastal Plain. Jack H. McLellan outlines the development and operation of Texasgulf's phosphate mine and manufacturing plant at Lee Creek, particularly as they relate to geological and paleontological studies. Thomas G. Gibson describes the regional patterns of Miocene-Pleistocene deposition in the Salisbury and Albemarle embayments of the central Atlantic Coastal Plain. On the basis of cluster analysis of 16 samples, including 149 taxa of ostracodes from fossiliferous beds above the Pungo River Formation, Joseph E. Hazel determines that the Yorktown Formation at the Lee Creek Mine is early Pliocene in age and the Croatan Formation spans the Plio-Pleistocene boundary. Among the ostracodes, 2 genera, 31 species, and one subspecies, are diagnosed as new. Walter H. Wheeler, Raymond B. Daniels, and Erling E. Gamble survey the post-Yorktown development in the region of the Neuse-Tar-Pamlico rivers. Primarily on the basis of auger holes, they begin with the Aurora paleoscarp marking the top of the Yorktown Formation, on which the organic-rich Small sequence (Croatan or James City Formation) was deposited, followed unconformably by the Pamlico morphostratigraphic unit; the inner edge of the Pamlico msu is associated with the Minnesott Ridge. H. Allen Curran and Patricia L. Parker divide the “Upper Shell” unit at the mine into three bivalve assemblage zones, probably formed through mass mortality in a series of local catastrophic events. Edward S. Belt, Robert W. Frey, and John S. Welch interpret Pleistocene deposition at the mine on the basis of biogenic and physical sedimentary structures, enabling them to recognize five major unconformities and four depositional sequences, indicative of a progradational shoreline under tectonically stable conditions. Their fourth depositional cycle includes a freshwater peat member thought to be of Sangamon interglacial age, on the basis of Donald R. Whitehead's pollen analysis. This analysis reveals high percentages of sedge and grass pollens, an absence of boreal indicators, tree pollen frequencies similar to those of interglacial deposits to the north and south, and general similarity of the fossil pollen spectrum to modern pollen assemblages of eastern North Carolina. Francis M. Hueber identifies the gymnospermous genera Pinus, Juniperus, and Taxodium, and tentatively the angiospermous genus Gleditsia, among the quartz-permineralized woods from the lower part of the Yorktown Formation at the mine; he also discusses the resin-like specimens, which are of unknown biological source and for which the stratigraphic source (Yorktown Formation, above the source of the woods) is known for only one specimen. William H. Abbott and John J. Ernissee report one silicoflagellate and two diatom assemblages (equivalent to Blow's zones N9 and N11) in a diatomaceous clay of the Pungo River Formation from two cores in Beaufort County; one new species of diatom is described. On the basis of 30 species of planktonic Foraminifera and a few radiometric dates, Thomas G. Gibson assigns ages from latest Oligocene through early Pleistocene to 10 stratigraphic units in the central Atlantic Coastal Plain; he describes 37 species and subspecies of benthic Foraminifera, of which 10 species and 2 subspecies are new. Scott W. Snyder, Lucy L. Mauger, and W.H. Akers assign an age of late-early to early-late Pliocene for a 15-meter section of the Yorktown Formation at the mine, based on 29 taxa of planktonic Foraminifera. Druid Wilson describes as a new genus and species of barnacle a puzzling fossil from inside the shell of the bivalve Mercenaria from the Croatan Formation. Porter M. Kier reports one species of echinoid from the Pungo River Formation, three from the Yorktown Formation, of which one is new, and two from the Croatan Formation. John E. Fitch and Robert J. Lavenberg record 45 taxa of teleost otoliths from the Yorktown Formation, representing 27 genera, of which 22 are new to the Pliocene of North America, and 6 are first fossil records.

The Carnivora of the Edson Local Fauna (Late Hemphillian), Kansas
Jessica A. Harrison
42 pages, 18 figures
1983 (Date of Issue: 16 November 1983)
Number 54, Smithsonian Contributions to Paleobiology
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The late Hemphillian Edson Quarry Local Fauna contains 36 species of amphibians, reptiles, birds, and mammals. The eight species of carnivorans are Canis davisi, a primitive dog; Osteoborus cyonoides, a large borophagine; Agriotherium species, a long-limbed bear; Plesiogulo marshalli, a wolverine; Pliotaxidea nevadensis, a badger; Martinogale alveodens, a skunk; Adelphailurus kansensis, a metailurine felid; and Machairodus coloradensis, a machairodontine felid. Edson is one of several fossil localities in Sherman County, Kansas, and was deposited in a series of fine sands within the Ogallala Formation. A secondary channel in a braided stream system is proposed as the environment of deposition. The high percentage of juveniles, as well as the vast numbers of the salamander Ambystoma kansensis, indicate accumulation during the spring of the year. The Edson Quarry Local Fauna compares very well with such typically late Hemphillian faunas as Coffee Ranch, Texas, and Optima, Oklahoma. Although only the carnivorans have been treated in depth, a listing of the vertebrate taxa is offered as well.

Fossil Spatangoid Echinoids of Cuba
Porter M. Kier
336 pages, 45 figures, 90 plates, 6 tables
1984 (Date of Issue: 21 March 1984)
Number 55, Smithsonian Contributions to Paleobiology
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The fossil spatangoid echinoids of Cuba are described based for the most part on specimens in the Sánchez Roig Collection. Seventy-nine species are recognized including 10 from the Late Cretaceous, 36 from the Eocene, 20 from the Oligocene-Miocene, 11 from the Miocene, and 2 of uncertain age. Three of the Eocene species are new: Schizaster formelli, Linthia monteroae, and Antillaster albeari. A new genus of schizasterid is described, Caribbaster, with the Eocene Prenaster loveni Cotteau as the type-species. A new Asterostoma, A. pawsoni, is described from the Eocene of Jamaica.

The Eocene age of the Cuban echinoid-bearing localities is confirmed by the presence outside Cuba of many of the same species in beds dated on other fossils. Some evidence supports the Miocene determinations, but the echinoids are of little assistance in resolving the question whether the Cuban beds attributed to the Oligocene are Oligocene or Miocene. Cuban, and in general, the Caribbean Tertiary echinoid faunas are distinct from those in Europe and the Mediterranean. Many genera are confined to the Caribbean. The Cuban fauna is also different from that found nearby in Florida. This difference may be due to a suggested greater depth of water in Cuba.

Se describen los equinoideos espatangoideos de Cuba, incluyendo los especímenes de la Colección Sánchez Roig. Se reconocen setena y ocho especies: 10 del Cretáceo Superior, 36 del Eoceno, 20 del Oligo-Mioceno, 11 del Mioceno, y 2 cuya edad no se ha determinado con certeza. Tres de las especies del Eoceno son neuvas: Schizaster formelli, Linthia monteroae, y Antillaster albeari. Se establece un nuevo género de schizasterid, Caribbaster, y se elige a Prenaster loveni Cotteau, del Eoceno, como especie-tipo. También se describe una nueva especie de Asterostoma, A. pawsoni, del Eoceno de Jamaica.

La edad de los equinoideos cubanos del Eoceno ha sido confirmada por la presencia, en foraciones de áreas fuera de Cuba, de muchas de las mismas especies en localidades donde se hallan otros fósiles del Eoceno. Las determinaciones del Mioceno están basadas en ciertas evidencias, sin embargo, los equinoideos ofrecen poca ayuda para dilucidar la cuetión de si las formaciones cubanas que se atribuyen al Oligoceno son de este período o del Mioceno. Las faunas de equinoideos del Terciario de Cuba, y en general del Caribe, son muy distintas a las de Europa y del Mediterráneo. Muchos géneros están confinados al Caribe y la fauna cubana es, asimismo, muy diferente a la que se encuentra en la cercana Florida. Esta diferencia puede ser debida a la mayor profunidad, según se ha sugerido, de las aguas de Cuba.

Echinoids from the Triassic (St. Cassian) of Italy, Their Lantern Supports, and a Revised Phylogeny of Triassic Echinoids
Porter M. Kier
41 pages, 4 figures, 14 plates
1984 (Date of Issue: 8 June 1984)
Number 56, Smithsonian Contributions to Paleobiology
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Three new species of Triassic echinoids are described from the St. Cassian (Karnian) beds of Cortina d'Ampezzo, Italy: Levicidaris furlani, L. pfaifferi, and Zardinechinus giulinii. Hundreds of echinoid fragments from the same beds show that 16 species lack apophyses (interambulacral lantern supports) and 7 possess them. Previously, paleontologists assumed that most Triassic echinoids had apophyses. Their absence from so many species and the presence of slightly developed auricles (ambulacral lantern supports) suggest that two echinoid lineages crossed from the Paleozoic to the Triassic: one, possessing apophyses, is ancestral to all modern cidaroids; a second, lacking apophyses, gave rise to all noncidaroid echinoids.

Giant Camels from the Cenozoic of North America
Jessica A. Harrison
29 pages, 17 figures, 4 tables
1985 (Date of Issue: 14 June 1985)
Number 57, Smithsonian Contributions to Paleobiology
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Seven genera of giant camels occurred in North America during the interval from the late Clarendonian to the early Holocene. Aepycamelus was the first camel to achieve giant size and is the only one not in the subfamily Camelinae. Blancocamelus and Camelops are in the tribe Lamini, and the remaining giant camels Megatylopus, Titanotylopus, Megacamelus, Gigantocamelus, and Camelus are in the tribe Camelini. Megacamelus is a late Hemphillian giant camel most closely related to Gigantocamelus. Titanotylopus is reserved for the brachyodont form from the Irvingtonian of Nebraska, and Gigantocamelus is reinstated for the broad-chinned, Blancan form.

Stratigraphic Record of the Neogene Globorotalid Radiation (Planktonic Foraminiferida)
Richard Cifelli and George Scott
101 pages, 43 figures
1986 (Date of Issue: 11 February 1986)
Number 58, Smithsonian Contributions to Paleobiology
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The histories of lineages forming the Neogene globorotalid radiation in the planktonic foraminifera are reconstructed primarily from stratigraphic distributions. Data on major taxa are synthesized, with particular reference to the development of shell design, and related to biogeograpy and evolutionary strategies.

The radiation was established about the base of the lower Miocene by three groups (Fohsella, Globorotalia zealandica lineage, and G. praescitula plexus), which probably arose from separate paragloborotalid lineages.

Common trends (size increase, chamber compression, keel development, reduced wall relief) early in the radiation culminated in the evolution of disklike taxa which, since the middle Miocene, have been centered in the tropics. The later phase of the radiation (post middle Miocene) was marked by architectural diversification as spiroconical (e.g., G. margaritae), ventroconical (e.g., G. truncatulinoides), and globose (e.g., G. inflata) taxa arose. Architectural diversification may be linked with watermass differentiation in the late Neogene.

Neogene designs have close counterparts in the earlier, but phyletically isolated, Paleogene and Cretaceous radiations. There are also resemblances in ontogenetic strategies and lineage histories. Common adaptations are suggested, but specific functional explanations have not been established.

Periods of major redesign are recognized in most lineages and are not confined to speciation events. Examples of stasis in adult morphology occur particularly in taxa that have evolved compressed, keeled shells. Bifurcations in lineages are indistinctly represented by wide spectra of morphotypes. Within the radiation very rapid speciation events are conspicuously absent, although they possibly occurred at the origin of some lineages. The distinctly sluggish tempo of change may be due to large population sizes and their degree of intercommunication. Good examples of allopatric and parapatric speciation were not found, but the prevalence of polytypic taxa, often distributed in contiguous populations showing clinal variation, would favor the inception of parapatric speciation.

Two New Oligocene Desmostylians and a Discussion of Tethytherian Systematics
Daryl P. Domning, Clayton E. Ray and Malcolm C. McKenna
56 pages, 23 figures
1986 (Date of Issue: 28 May 1986)
Number 59, Smithsonian Contributions to Paleobiology
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A new genus, comprising two new species of desmostylians, is described from marine Oligocene deposits of the Pacific Northwest. Behemotops proteus, new genus, new species, is based on an immature mandibular ramus and apparently associated skeletal fragments from the middle or (more likely) upper Oligocene lower part of the Pysht Formation of Clallam County, Washington. A related new species, Behemotops emlongi, is founded on a mandibular ramus of an old individual and a mandibular fragment with canine tusk from the uppermost Oligocene (early Arikareean equivalent) Yaquina Formation of Lincoln County, Oregon. The two new species are the most primitive known desmostylians and compare favorably with the primitive Eocene proboscideans Anthracobune and Moeritherium, and to the still more primitive tethythere Minchenella from the Paleocene of China.

For many years the Desmostylia were widely regarded as members of the mammalian order Sirenia before being accepted as a taxon coordinate with the Sirenia and Proboscidea (Reinhart, 1953). On the basis of cladistic analysis we go a step further and regard the Desmostylia as more closely related to Proboscidea than to Sirenia because the Desmostylia and Proboscidea are interpreted herein to share a more recent common ancestor than either order does with the Sirenia. This analysis also suggests that the common ancestor of the Proboscidea and Desmostylia (but not the Sirenia) had suppressed P5 and the original last molar. These characters may be convergent with some other mammals. The Superorder Tokotheria McKenna, 1975, was originally thought to be characterized by loss of both P5 and M3. However, because early sirenians do not show these losses, they may have occurred independently in the common ancestor of proboscideans and desmostylians and in various other tokotheres.

The late Paleocene genus Minchenella Zhang, 1980, from China, is a suitable candidate to be the common ancestor of both the Desmostylia and the Proboscidea. It possesses a small entoconid II on M3. The Eocene genus Lammidhania Gingerich, 1977, from Pakistan, and the late Paleocene and/or early Eocene Chinese and Mongolian phenacolophids had not acquired an entoconid II on M3 but are otherwise similar to Minchenella and the anthracobunids. The Asiatic occurrence of phenacolophids, Lammidhania, Minchenella, and anthracobunids suggests an Asian origin for the Proboscidea and is in accord with the exclusively Pacific distribution of the Desmostylia.

We believe that desmostylians were amphibious herbivores that fed on marine algae and angiosperms, and that at least the earlier taxa depended to a large extent on plants exposed in the intertidal zone.

Microdistribution of Foraminifera in a Single Bed of the Monterey Formation, Monterey County, California
Roberta K. Smith and Martin A. Buzas
33 pages, 4 figures, 2 plates, 7 tables
1986 (Date of Issue: 27 October 1986)
Number 60, Smithsonian Contributions to Paleobiology
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While several papers exist on the small scale spatial distribution of living foraminifera, almost no work exists on the small scale spatial distribution of fossils. The present study took 24 (5 ml) replicates 10 cm apart along one bed of the Monterey Formation in California.

The mean density for all replicates is 6084.96 with a standard deviation of 8776.95. Both inspection and a cluster analysis of the data indicate replicates 20-24 have a much higher density and different rank order of abundance than replicates 1-19. The mean density for the total of all species in replicates 1-19 is 2387.47 with a standard deviation of 1175.58. For replicates 20-24 the mean density is 20135.40 with a standard deviation of 11181.40. The spatial variability is so great that four replicates (more than commonly taken) would only allow us to be 95% confident that we are within 50% of the true mean. Because age determination is based on presence of particular taxa rather than on densities, stratigraphic assignment would still be possible.

The three species dominating the 1-19 group make up from 86% to 99% of the fauna. The three species dominating the 20-24 group make up from 77% to 85% of the fauna. Two of these are also dominant in the 1-19 group, but the most dominant species in the 20-24 group constitutes only <1% to 8% in the 1-19 group.

The greatest number of species (22) occurs in the 20-24 group, as would be expected from the densities. The 1-19 group has 16 species. The information function is also highest in the 20-24 group.

An attempt was made to achieve the faunal composition of the 1-19 group for replicates 20-24 by removal of percents of small-sized taxa. Comparable relative abundances are best achieved by removing 100% of Epistominella subperuviana and 95% of Bolivina brevior and other significant small-sized species. Total specimen numbers for both small- and large-sized species remains higher in replicates 20-24 than in 1-19, however. Thus, analysis of species percentages and species specimen size indicates that while transportation—winnowing—of small specimens (or large specimens) into or out of the environment of deposition may be significant, it does not account for the differences between replicates 1-19 and 20-24. Therefore, either two habitats or some other mode of allocthonous enrichment or depletion rather than particle size winnowing must be invoked to account for the observed distribution.

The low numbers of species, especially with so many individuals, indicates the fauna probably lived under stressful conditions. Low amounts of available oxygen may have caused the stress.

Geology and Paleontology of the Lee Creek Mine, North Carolina, II
Clayton E. Ray, editor
283 pages, 49 figures, 80 plates, 21 tables
1987 (Date of Issue: 15 June 1987)
Number 61, Smithsonian Contributions to Paleobiology
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Volume I of this projected series of three volumes included the prologue to the series, a biography of Remington Kellogg, and 13 papers on geology and paleontology other than Mollusca and Vertebrata (except otoliths). It was published in 1983 as Smithsonian Contributions to Paleobiology number 53. The present volume consists of a foreword and five chapters devoted to molluscan paleontology. The foreword recounts the earliest scientific publication of New World fossils, all mollusks, and reproduces Martin Lister's illustrations of them. William M. Furnish and Brian F. Glenister record the nautilid genus Aturia from the Pungo River Formation and discuss its occurrence elsewhere. Druid Wilson describes a new pycnodont oyster from the Pungo River Formation and lists the Cenozoic pycnodonts from the Atlantic and Gulf Coastal Plain; he also summarizes the stratigraphic and geographic occurrences of the subgenera of Ecphora, Ecphora and Stenomphalus, naming a new species of each from the Pungo River Formation, and a new species of the former from the St. Marys Formation of Maryland. Thomas G. Gibson clarifies the relationships and stratigraphic utility of 17 taxa (including one new species from the Pungo River Formation) of pectinid bivalves on the basis of biometric study of large samples from lower Miocene to lower Pleistocene beds in and near the mine. Lauck W. Ward and Blake W. Blackwelder describe a molluscan fauna of 194 species, including 30 new species and 3 new subspecies, from the Chowan River (upper Pliocene) and James City (lower Pleistocene) formations, and conclude that the fauna reflects a subtropical thermal regime and that it was deposited under open marine conditions at depths not exceeding 25 meters.

Fossil Birds from the Oligocene Jebel Qatrani Formation, Fayum Province, Egypt
D. Tab Rasmussen, Storrs L. Olson and Elwyn L. Simons
20 pages, 15 figures
1987 (Date of Issue: 1 December 1987)
Number 62, Smithsonian Contributions to Paleobiology
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Fossils from fluvial deposits of early Oligocene age in Egypt document the earliest known diverse avifauna from Africa, comprising at least 13 families and 18 species. Included are the oldest fossil records of the Musophagidae (turacos), Pandionidae (ospreys), Jacanidae (jacanas), and Balaenicipitidae (shoebilled storks). Other families represented are the Accipitridae (hawks and eagles), Rallidae (rails), Gruidae (cranes), Phoenicopteridae (flamingos), Ardeidae (herons), Ciconiidae (storks), and Phalacrocoracidae (cormorants). A highly distinctive rostrum is described as a new family, Xenerodiopidae, probably most closely related to herons. A humerus lacking the distal end is tentatively referred to the same family. Two new genera and three species of large to very large jacanas are described from the distal ends of tarsometatarsi. This Oligocene avifauna resembles that of modern tropical African assemblages. The habitat preferences of the constituent species of birds indicate a tropical, swampy, vegetation-choked, fresh-water environment at the time of deposition.

The Cretaceous Birds of New Jersey
Storrs L. Olson and David C. Parris
22 pages, 11 figures
1987 (Date of Issue: 8 December 1987)
Number 63, Smithsonian Contributions to Paleobiology
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This is a revision of the fossil birds from Late Cretaceous (Maastrichtian; Hornerstown and Navesink formations) deposits in New Jersey. Material of previously named taxa, described over a century ago, is augmented by more recently collected specimens from a new locality at the Inversand Company marl pits near Sewell, Gloucester County. With about 8 genera and 9 species, this is the most diverse Cretaceous avifauna yet known. Most species belong to a group of primitive Charadriiformes resembling in limb morphology the fossil family Presbyornithidae and the living family Burhinidae. These are tentatively referred to the “form family” Graculavidae Fürbringer, 1888, with its provisional synonyms Palaeotringinae Wetmore, 1940; Telmatornithidae Cracraft, 1972, and Laornithidae Cracraft, 1972. The species included are: Graculavus velox Marsh, 1872; Telmatornis priscus Marsh, 1870 (synonyms: Telmatornis affinis Marsh, 1870; Graculavus pumilus Marsh, 1872; Palaeotringa vetus Marsh, 1870); Anatalavis rex (Shufeldt, 1915); Laornis edvardsianus Marsh, 1870; Palaeotringa littoralis Marsh, 1870; P. vagans Marsh, 1872; and an undescribed genus and species probably different from any of the preceding. Anatalavis is proposed as a new genus for Telmatornis rex Shufeldt, 1915. A new family, genus, and species (Tytthostonychidae, Tytthostonyx glauconiticus) is proposed for a humerus showing similarities to the Pelecaniformes and Procellariiformes and tentatively referred to the latter, along with an ulna of a much smaller species. The species in this fauna appear to be part of the modern radiation of neognathous birds, but none can be referred to modern families.

Some Tertiary Brachiopods of the East Coast of the United States
G. Arthur Cooper
45 pages, 11 figures, 9 plates, 2 tables
1988 (Date of Issue: 28 April 1988)
Number 64, Smithsonian Contributions to Paleobiology
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The Tertiary brachiopod fauna of the United States is discussed and reference is made to the composition of the World Tertiary fauna. The brachiopod fauna of the East Coast of the United States at present numbers 17 genera (4 inarticulates and 13 articulates), of which 11 are discussed herein (asterisk): 2 linguilidids: Glottidia,* Discradisca; 2 craniidinids: Isocrania?* and Crania?*; 2 rhynchonellids: Probolarina* and Cryptopora, the former extinct; 2 cancellothyrids: Eucalathis* and Terebratulina*; 5 extinct terebratulids: Embolosia, Oleneothyris, Plicatoria,* Rhytisoria,* and Tanyoscapha; 2 terebratellids: Argyrotheca* and Platidia; and 2 thecideids: Lacazella* and Thecidellina.* Pliocene Glottidia inexpectans Olsson in adult form is described. New species of Probolarina, Terebratulina, Argyrotheca, Lacazella, and Thecidellina are described and illustrated. Specimens not identified as to species are described: Crania,?, Isocrania?, Eucalathis?, and Terebratulina. The remarkable variation of Plicatoria wilmingtonensis (Lyell and Sowerby) is described and illustrated.

Jurassic Brachiopods of Saudi Arabia
G. Arthur Cooper
213 pages, 48 figures, 37 plates
1989 (Date of Issue: 13 July 1989)
Number 65, Smithsonian Contributions to Paleobiology
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No studies in depth have been made of the brachiopods from the Jurassic deposits of Saudi Arabia. This first study of the brachiopods from this important region is based mainly on a collection presented to the Smithsonian Institution by the Arabian-American Oil Company (Aramco). In addition, the study includes collections made by Drs. P.M. Kier and E.G. Kauffman of the Smithsonian Institution and R.W. Powers, C.D. Redmond and H.A. MacClure of the Arabian-American Oil Company.

Sixty-one genera are described of which 29 are new. Of these 13 rhynchonellid genera are new: Amydroptychus, Baeorhynchia, Colpotoria, Conarosia, Deltarynchia, Echyrosia, Eurysites, Heteromychus, Lirellarina, Nastosia, Pycnoria, Schizoria, and Strongyloria. Other described genera are: Burmirhynchia Buckman, 1917, Cymatorhynchia Buckman, 1917, Daghanirhynchia Muir-Wood, 1935, Gibbirhynchia Buckman, 1917, Globirhynchia Buckman, 1917, Kallirhynchia Buckman, 1917, Kutchirhynchia Buckman, 1917, Somalirhynchia Weir, 1925, Sphenorhynchia Buckman, 1917, and Torquirhynchia Childs, 1969.

Of Spiriferinacea, one genus, Calyptoria, is new and two genera are described: Liospiriferina Rouselle, 1977, and Spiriferina d'Orbigny, 1847. The Terebratulacea are represented by 11 new genera: Arabatia, Arabicella, Arapsopleurum, Arapsothyris, Dissoria, Ectyphoria, Pionopleurum, Pleuraloma, Stenorina, Tanyothyris, and Toxonelasma. Seventeen described terebratulaceans are Apatecosia Cooper, 1983, Avonothyris? Buckman, 1917, Bihenithyris Muir-Wood, 1935, Dolichobrochus Cooper, 1983, Dorsoplicathyris? Almeras, 1971, Glyphisaria? Cooper, 1983, Gyrosina? Cooper, 1983, Habrobrochus Cooper, 1983, Kutchithyris? Buckman, 1917, Loboidothyris? Buckman, 1917, Orthotoma Quenstedt, 1869, Plectothyris? Buckman, 1917, Pseudowattonithyris? Almeras, 1971, Somalithyris Muir-Wood, 1935; Sphaeroidothyris Buckman, 1917, Stiphrothyris? Buckman, 1917, and Striithyris Muir-Wood, 1935.

The Zeilleriacea include four new genera: Apothyris, Mycerosia, Sphriganaria and Xenorina. Described zeilleriids are Flabellothyris Eudes-Deslongchamps, 1884, Rugitela Muir-Wood, 1936, and Zeilleria Bayle, 1878. A total of 166 species are described and 25 lots are identifiable as species.

Pseudoglossothyris? sulcata Muir-Wood, 1935, from Somaliland (Somali Republic) is shown to be a zeilleriid, and the species is herein transferred to Aulacothyris. Eudesia cardioides Douvillé, 1916, is herein transferred to the new genus Sphriganaria.

The Liassic Marrat Formation abounds in spiriferinids. The Dhruma Formation (Bajocian to Callovian) is rich in rhynchonellids which dwindle in numbers in late Dhruma beds. The overlying Tuwaiq Mountain and Hanifa formations (Callovian to Kimmeridgian) are conspicuous for the large numbers of terebratulaceans which far outnumber the rhynchonellids.

Correlation with Jurassic sequences near and far is difficult because of the high degree of endemism shown by the Saudi Arabian brachiopods. Precise correlation with British and European faunas is not now possible. Relationships with the Jurassic faunas of the Sinai, Israel and East Africa in the Callovian is suggested by the presence of two species in common and generic representation shown by Daghanirhynchia, Somalirhynchia, Bihenithyris, Somalithyris, and Striithyris.

The Autochthonous North American Musk Oxen Bootherium, Symbos, and Gidleya (Mammalia: Artiodactyla: Bovidae)
Jerry N. McDonald and Clayton E. Ray
77 pages, 64 figures, 4 tables
1989 (Date of Issue: 8 June 1989)
Number 66, Smithsonian Contributions to Paleobiology
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The history of taxonomy of the autochthonous genera of North American musk oxen—Bootherium, Symbos, and Gidleya—is reviewed. The bases upon which taxonomic judgments within the group have been made are identified. These bases are reevaluated in the light of current information on patterns of ontogenesis, sexual dimorphism, postmortem alteration of skeletal remains, and spatial and temporal distribution of musk ox records. The bases used by taxonomists in the past to justify separation of this musk ox group into multiple genera and species can be explained best as indices of sexual dimorphism or postmortem weathering and abrasion. All nominal species within Bootherium, Symbos, and Gidleya are, therefore, placed in synonymy with the senior name in the group, Bootherium bombifrons (Harlan, 1825). A revised diagnosis is provided for the monotypic species.

Rodents of the Bridgerian (Middle Eocene) Elderberry Canyon Local Fauna of Eastern Nevada
Robert J. Emry and William W. Korth
14 pages, 5 figures
1989 (Date of Issue: 30 November 1989)
Number 67, Smithsonian Contributions to Paleobiology
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The Elderberry Canyon Local Fauna, presently represented by more than 40 vertebrate taxa, including at least 30 mammals, occurs in carbonate rocks assigned to the Sheep Pass Formation, near Ely, Nevada. Among the mammals, nine rodent species, representing four families, are recognized. Reithroparamys delicatissimus, and R. cf. R. huerfanensis are species previously known from the Rocky Mountain region; Microparamys sambucus and Pauromys exallos are new species assigned to genera commonly occurring in Eocene faunas elsewhere; Elymys complexus is a new genus and species of minute rodent thought to be related to Simimys and questionably assigned to the family Zapodidae; Sciuravus, Mattimys, and Knightomys are represented by material not assignable to species; and another unidentified ischyromyid completes the roster of rodents presently known in the assemblage. The fauna as a whole, and the rodents in particular, share the greatest homotaxial similarity with early Bridgerian (early middle Eocene) faunas elsewhere, allowing a confident early Bridgerian age assignment for the Elderberry Canyon Local Fauna. The composition of this rodent assemblage suggests that faunal interchange was relatively unrestricted from southern California through Nevada to the Rocky Mountain region in Bridgerian time.

Spatial Distribution of Miocene Foraminifera at Calvert Cliffs, Maryland
Martin A. Buzas and Thomas G. Gibson
35 pages, 4 figures, 14 tables
1990 (Date of Issue: 7 March 1990)
Number 68, Smithsonian Contributions to Paleobiology
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Excavations made in middle Miocene strata in the Calvert Cliffs of Maryland during construction of a nuclear power plant were used for a spatial distribution study of fossil benthic foraminifera over large bedding surfaces. Two bedding surfaces were sampled, a larger and older one involving a 400 m2 surface in the Calvert Formation, and a slightly younger one involving a 50 m2 surface in the Choptank Formation. The sampling procedure for both surfaces consisted of a 3- × 3-station grid, with 5 replicates taken at each of 9 stations. The larger surface had stations at 9.5 m centers, and the smaller surface had stations at 3.6 m centers.

The amount of variation in species proportions and densities from each bedding surface were used to determine how much confidence can be placed in the results from the usual paleontologic sampling procedure of a single sample.

Unispecies and multispecies analyses were done on the 45 samples from each of the surfaces. The study of the older surface involved 36 species, and the younger 33. Analyses indicate a remarkable degree of homogeneity of species densities and proportions in both of these beds. Species usually remain in the same rank order at all stations within each surface, indicating that any of the 45 samples gives a reasonable species composition for the surface. Species densities exhibit greater variability; the determination of confidence limits for species densities requires multiple samples to reach limits of ±50 percent accuracy.

New Enaliarctos* (Pinnipedimorpha) from the Oligocene and Miocene of Oregon and the Role of “Enaliarctids” in Pinniped Phylogeny
Annalisa Berta
33 pages, 22 figures, 7 tables
1991 (Date of Issue: 16 December 1991)
Number 69, Smithsonian Contributions to Paleobiology
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Three new species of the pinnipedimorph Enaliarctos* are described from the marine late Oligocene and early Miocene (Arikareean and Hemingfordian or early Barstovian correlatives) of coastal Oregon. Enaliarctos tedfordi, new species, is based on a partial cranium from the late Oligocene Yaquina Formation. A related new species, Enaliarctos emlongi, is founded on a nearly complete cranium, jaws, and associated skeletal elements from the late Oligocene to early Miocene Nye Mudstone. A third new species, Enaliarctos barnesi, is based on a partial cranium and jaws from late Oligocene or early Miocene rocks near the contact between the Yaquina Formation and the Nye Mudstone. Another skull, from the Nye Mudstone, is referred to a previously described species, Enaliarctos mitchelli Barnes, 1979. Three of these species, E. mitchelli, E. emlongi, and E. tedfordi form a monophyletic clade, united by reduced cheek teeth cingula and short metacone of the upper carnassial. The major trend observed in Enaliarctos over 10 million years of history is an intermediate stage in the transformation to homodonty evidenced by premolarization of the upper carnassial and molars and reduction and simplification of cusps on the lower carnassial.

Cladistic analysis of 52 cranial and dental characters suggests the following phylogenetic hypotheses: (1) the subfamily “Enaliarctinae” (= “Enaliarctidae”) is paraphyletic, (2) monophyly of the genus Enaliarctos* is questioned although the status of this taxon as sister taxon to other pinnipeds is affirmed, (3) other “enaliarctid” pinnipeds, Pteronarctos and Pinnarctidion, are assigned to less inclusive pinniped clades (Pteronarctos + all other pinnipeds and Pinnarctidion + Desmatophoca, Allodesmus, and the Phocidae).

Morphology, Anatomy, and Systematics of the Cinctiporidae, New Family (Bryozoa: Stenolaemata)
Richard S. Boardman, Frank K. McKinney and Paul D. Taylor
81 pages, 137 figures
1992 (Date of Issue: 14 April 1992)
Number 70, Smithsonian Contributions to Paleobiology
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The study of thin sections, peels, and SEM micrographs of skeletons, and sections of skeletons and soft parts together, has revealed new morphology and anatomy resulting in new growth and functional interpretations in a new stenolaemate family, the Cinctiporidae. Cinctiporids are found primarily in the New Zealand region and range from upper Cretaceous to Recent. In contrast to the present classification of stenolaemates, the family includes eight species grouped into two free-walled genera, one fixed-walled genus, and one mixed free-/fixed-walled genus. This unconventional family is inferred to be monophyletic based upon a number of shared character states, zones of astogenetic change with fixed, free, and fixed/free apertures, and zones of repetition with both fixed- and free-walled zooids. These taxa are described using both external and internal skeletal morphology and soft part anatomy. Zooids of cinctiporids are typically several times larger than those of other stenolaemates, facilitating detailed observations.

In dendroid growth habits, growth rates of zooids are greatest in endozones at growing tips of branches, and in cinctiporids skeletal and fully regenerated polypide sizes are roughly proportional. Growth rates decrease greatly as young zooids reach exozones by saltation in a series of polypide cycles. In exozones, attachment organs become fixed in position, some in attachment scars in skeletal linings, resulting in regenerated polypides being fixed in position in subsequent cycles. During regenerating phases of any single cycle, polypides grow inward from attachment organs, ingesting and eliminating as they grow. Retractor muscles must function, therefore, and must slide their skeletal connections inward also as polypides increase in length. In early phases of a regeneration the developing polypides ingest and eliminate from within their living chambers. Elimination within living chambers is apparently facilited by faecal pellets passing out through the atrium and vestibule. The funiculus of stenolaemates generally ends blindly against skeletal walls preventing connection to neighboring zooids by that means as in gymnolaemates. A large funicular muscle or muscles aids in retracting polypides of the cinctiporids.

Skeletal walls calcified from one side necessarily form against pre-existing membranes. Exterior skeletal walls in stenolaemates calcify against outer cuticles, preserving the shapes of the cuticles as they are calcified, explaining growth undulations on frontal walls and intimate contacts that basal colony walls make with substrates.

A New Genus and Species of Boxfish (Tetraodontiformes: Ostraciidae) from the Oligocene of Moravia, the Second Fossil Representative of the Family
James C. Tyler and Ruzena Gregorova
20 pages, 8 figures, 1 table
1991 (Date of Issue: 16 December 1991)
Number 71, Smithsonian Contributions to Paleobiology
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A new genus and species of boxfish (Ostraciidae) is described on the basis of five specimens from the Menilitic Formation of the Lower Rupelian part of the middle Oligocene (ca. 35 MYA) of Moravia, Czechoslovakia. A sixth but incomplete specimen is recorded from the Menilitic Formation of southern Poland. The fossil record of the family previously was based only on Eolactoria sorbinii from the lower Eocene of Monte Bolca, Italy, and on isolated carapace scale plates also dating back to the Eocene. The new genus differs most significantly from all other Recent and fossil ostraciids in the greater posterior extension of the carapace onto the caudal peduncle; the lesser number of anteroposteriorly compressed centra of the vertebrae of the caudal peduncle; the greater length of the caudal fin in relatively small specimens; and perhaps by having a reduced number of caudal rays (nine rather than 10).

The new species is a typical member of the in-shore benthic Ostraciidae, but the other fishes with which it is associated in the Menilitic Formation are nearly all more off-shore mesopelagic species of Myctophidae and Gonostomatidae. To account for this, we propose that the new species represents the oceanic, pelagic, pre-settlement stage of this genus, as in the Recent Lactoria, some species of which, especially L. diaphana, remain pelagic to sizes of over 100 mm SL and can be sexually mature while pelagic before settling into their conclusive benthic state.

A Remarkable New Genus of Tetraodontiform Fish with Features of Both Balistids and Ostraciids from the Eocene of Turkmenistan
James C. Tyler and Alexandre F. Bannikov
14 pages, 4 figures, 1 table
1992 (Date of Issue: 20 December 1992)
Number 72, Smithsonian Contributions to Paleobiology
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A new genus and species of tetraodontiform fish, Eospinus daniltshenkoi, is described from the Lower Eocene of Turkmenistan (Danatinsk Formation). It is referred to the Balistoidea because it has three large dorsal-fin spines; the pelvic fin reduced to a rudimentary but prominent structure apparently composed of two partially fused spines at the posterior end of the pelvis; and enlarged scale plates that form an incomplete carapace or loosely articulated armature around much of the body. It differs from all other balistoids in having a long median spine projecting forward from the snout and another spine projecting posteriorly from the middle of each side of the body; lower jaw teeth twice as long as the upper jaw teeth; and in lacking encasing scales around the rudimentary pelvic spine. Eospinus is the first record of a balistoid fish from the Eocene with three dorsal-fin spines and the pelvic spines fused together at the end of the pelvis, as otherwise only occurs in balistids, which are first recorded from the Oligocene.

Jurassic Rhynchonellids: Internal Structures and Taxonomic Revisions
Xiao-ying Shi and Richard E. Grant
190 pages, 83 figures, 18 plates
1993 (Date of Issue: 1 November 1993)
Number 73, Smithsonian Contributions to Paleobiology
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Jurassic brachiopods of the order Rhynchonellida are classified according to modern concepts and techniques, with special attention to internal structures. They are grouped into 6 families and 16 subfamilies of which three are new: the Acanthorhynchiinae, the Cryptorhynchiinae, and the Piarorhynchiinae. Subfamilies emended or revised are the Acanthothyridinae Schuchert (1913) raised to family rank, Davanirhynchinae Ovtsharenko (1983), Dzhangirhynchinae Ovtsharenko (1983), Erymnariinae Cooper (1959), Indorhynchiinae Ovtsharenko (1975), Septocrurellinae Ager, Childs, and Pearson (1972), and Striirhynchiinae Kamyshan (1968). New genera are Aalenirhynchia (type-species Rhynchonella subdecorata Davidson, 1853), Bradfordirhynchia (type-species Cryptorhynchia bradfordensis Buckman, 1918), and Sharpirhynchia (type-species Kallirhynchia sharpi Muir-Wood, 1938). A new subgenus is Burmirhynchia (Hopkinsirhynchia) (type-species Burmirhynchia hopkinsi Davidson, 1854). The only new species is Pycnoria depressa. Eleven genera are revised, and many are transferred among the subfamilies; lectotypes are designated where needed.

Paleobiology of Climactichnites, an Enigmatic Late Cambrian Fossil
Ellis L. Yochelson and Mikhail A. Fedonkin
74 pages, 58 figures
1993 (Date of Issue: 16 April 1993)
Number 74, Smithsonian Contributions to Paleobiology
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Climactichnites wilsoni Logan, 1860, is redescribed from field investigations and specimens in various museums. Climactichnites youngi Todd, 1882, and C. fosteri Todd, 1882, are placed in synonymy. The species is known only from its trail, consisting of raised bars and impressed furrows, bounded by two parallel, raised lateral ridges. This structure is interpreted as being formed from damp sand, redistributed and molded by the animal. At a few localities, an oval impression occurs at the origin of the trail; a new locality for this rare feature was found in Wisconsin. In Quebec a trail crossing over itself was found; this phenomenon was known from only one other locality.

All occurrences of Climactichnites are in the Late Cambrian; specimens are known from New York, Quebec, Ontario, Wisconsin, and Missouri. This fossil is probably confined to the Dresbachian, the earliest stage of the tri-part Late Cambrian. All examples of trails are in sandstones; these are interpreted as sand flats that were just above water during low tide in the shallow epicontinental sea.

A variety of animals have been proposed as the trail maker; they include several different kinds of arthropods, mollusks, and “worms.” Each proposal has weak points and none of the suggested animal groups has appropriate morphology to produce the marking. The animal is reconstructed as relatively low, broad, and about twice as long as wide. It is hypothesized that the tough body integument secreted mucus which facilitated movement and aided in preservation of the trail. A large flap covered most of the outside body and extended laterally over the muscular foot. Free edges of the flap on either side of the body compressed and molded damp sand into parallel bounding ridges. Respiratory organs may have been present below the flap edges, kept moist by being partially enclosed. By moving the edges of these lateral flaps, the organism may have been able to swim when in water.

Most trails of Climactichnites are interpreted as a consequence of feeding activity. If so, food was taken in through a circular mouth located anteriorward on the ventral surface; this morphologic feature is inferred from circular markings seen on a few bars. As reconstructed, the sand under the animal was compressed anteriorly and laterally; the animal then brushed particles forward into a small dune-like bar, probably by the action of cilia on its ventral surface. Sand dwelling microorganisms displaced by brushing were concentrated centrally, obviating ingestion of large amounts of sediment. What sediment was taken in was released at irregular intervals through a posterior anus; medial marking on parts of some trails are interpreted as fecal strings. The complex clamping and brushing behavior, implies a well-developed nervous system.

No body fossils are known in the Vendian or in the Paleozoic that could have constructed this form of trail. The proposed method of feeding, if correctly interpreted, is unique. Thus the trail of Climactichnites may constitute the work of an otherwise unknown phylum in the animal kingdom.

Two New Genera and Species of Oligocene Spikefishes (Tetraodontiformes: Triacanthodidae), the First Fossils of the Hollardiinae and Triacanthodinae
James C. Tyler, Anna Jerzmanska, Alexandre F. Bannikov and Jacek Swidnicki
27 pages, 20 figures, 3 tables
1993 (Date of Issue: 14 July 1993)
Number 75, Smithsonian Contributions to Paleobiology
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Two new genera and species of spikefishes from the Menilitic Formation (late Tethys Sea) of the Upper Oligocene of Poland represent the first fossils of the two subfamilies of the tetraodontiform family Triacanthodidae. One of the new genera, Prohollardia, has a dome-like supraoccipital, the epiotics separated medially on the dorsal surface of the skull, the epiotics articulated anteriorly with the frontals, and a shaft-like posterior process of the pelvis, which are diagnostic features of the Hollardiinae. The other, Carpathospinosus, has a flattened supraoccipital with only a small crest anteromedially, the epiotics in contact medially on the dorsal surface of the skull, the epiotics separated from the frontals by the sphenotic, and a broad basin-like posterior process of the pelvis, which are diagnostic features of the Triacanthodinae. Some of these features of the Triacanthodinae are shown to be derived.

The separation of the two subfamilies of Triacanthodidae took place no less than about 29 to 24 MYA.

In an addendum, the Oligocene fish from Romania that was described in the dactylopteriform family Cephalacanthidae (Dactylopteridae) as Cephalacanthus trispinosus Ciobanu (1977) is referred to the Triacanthidae (the anatomically derived sistergroup of the Triacanthodidae) as a member of the triplespine genus Acanthopleurus Agassiz (1842). The single specimen is a juvenile and at least closely related to A. serratus Agassiz (1842) and A. collettei Tyler (1980), both from the Oligocene of Switzerland, and possibly identical to one or the other.

Brachiopods Near the Permian-Triassic Boundary in South China
Guirong Xu and Richard E. Grant
68 pages, 54 figures, 7 tables
1994 (Date of Issue: 24 March 1994)
Number 76, Smithsonian Contributions to Paleobiology
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Sixty-eight genera and 164 species in the Changxingian Stage and 12 genera and 20 species in the lower Griesbachian Stage are recorded on the basis of brachiopod fossils collected from 32 sections in South China and from review of the Chinese literature. Of these, 24 genera and 34 species are described here, including three new genera (Fanichonetes, Prelissorhynchia, and Rectambitus) and 24 new species (Acosarina strophiria, Enteletes asymmatrosis, Peltichia schizoloides, Derbyia pannuciella, Perigeyerella altilosina, Chonetinella cursothonia, C. volitanliopsis, Fanichonetes campigia, Cathaysia spiriferoides, Uncinunellina multicostifera, Prelissorhynchia triplicatioid, Cyrolexis antearcus, Cyrolexis beccojectus, Cartorhium xikouensis, C. twifurcifer, Callispirina rotundella, Araxathyris subpentagulata, A. beipeiensis, Spirigerella discsella, S. ovaloides, Squamularia formilla, Hustedia orbicostata, Rostranteris ptychiventria, and Notothyris bifoldes).

The Cathaysia chonetoides-Chonetinella substrophomenoides assemblage zone and the Cathaysia sinuata-Waagenites barusiensis assemblage zone represent respectively faunas of the lower Changxingian and the upper Changxingian in clastic lithofacies; whereas the Peltichia zigzag-Prelissorhynchia triplicatioid assemblage zone and the Spirigerella discusella-Acosarina minuta assemblage zone represent faunas in limestone lithofacies. The Crurithyris pusilla-Lingula subcircularis assemblage zone and Permian-type brachiopods are present in the lower Griesbachian.

The Changxingian brachiopod fauna can be correlated with the Dorashamian fauna of Armenia; the brachiopod faunas of the Ali Bashi Formation, North-West Iran; unit 7 of the Hambast Formation, Central Iran; and the upper part of the Bellerophon Formation of the Southern Alps.

The genera Cathaysia, Peltichia, and Prelissorhynchia are especially characteristic of the Cathaysia Tethyan Subprovince. In contrast, the West Tethyan Subprovince is characterized by the genera Costiferina, Ombonia, Comelicania, and many other species. Four brachiopod ecofacies are recognized in the Changxingian of South China: (1) antibiohermal dwellers; (2) calcareous substratum dwellers; (3) biohermal dwellers; and (4) ubiquitous substrate dwellers. In the lower Griesbachian, the brachiopod fauna of Lingula and Crurithyris spreads across the entire Tethys and is called the Circum-Pangaea brachiopod fauna.

Massive extinction of brachiopod faunas occurred at the close of the Changxingian, with only a few Permian-types surviving into the early Griesbachian, and they completely vanished after the early Griesbachian except for harbingers of Mesozoic brachiopods.

Ontogenetic Morphometrics of Some Late Cretaceous Trochospiral Planktonic Foraminifera from the Austral Realm
Brian T. Huber
85 pages, 37 figures, 10 plates, 15 tables
1994 (Date of Issue: 11 January 1994)
Number 77, Smithsonian Contributions to Paleobiology
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Biometric analysis of ontogenetic changes in test morphology is employed to determine the taxonomic status of several trochospiral planktonic foraminiferal species from southern high latitude Upper Cretaceous sediments. Ontogenetic morphometric data obtained from specimens of Hedbergella sliteri Huber, Archaeoglobigerina australis Huber (both micromorph and normal-sized populations), and Archaeoglobigerina mateola Huber are compared with topotype populations of Hedbergella holmdelensis Olsson, H. monmouthensis (Olsson), Costellagerina pilula (Belford), and Rugoglobigerina rugosa (Plummer). Southern South Atlantic specimens of Archaeoglobigerina bosquensis Pessagno and Archaeoglobigerina cretacea (d'Orbigny) are also analyzed for comparison. Numerous biometric data, measured from exterior observations of whole tests, contact microradiographs, and Scanning Electron Microscope (SEM) micrograph images of serially dissected foraminifera, are used to characterize developmental changes in morphology of the planktonic foraminiferal species. The most useful variables for discriminating taxonomic differences are discussed for each method.

Results indicate that the ontogenetic morphometric approach to study of planktonic foraminifera can be effectively used to resolve problems in taxonomic classification, particularly for species that appear homeomorphic in exterior view. This approach was particularly useful for demonstrating that the ontogenetic morphologies of A. australis, C. pilula, and R. rugosa are very different and, therefore, previous assignment of A. australis morphotypes to various species of Rugoglobigerina and Costellagerina were incorrect. This study also demonstrates that the growth morphology of the new high latitude species H. sliteri significantly differs from H. holmdelensis and H. monmouthensis, thus confirming recognition of H. sliteri as a valid taxon. However, taxonomic uncertainty persists for some high latitude morphotypes that have external characteristics similar to R. rugosa (e.g., faint umbilical apertures, faint costellae that are meridionally aligned, presence of tegilla), but ontogenetic morphologies more similar to A. australis.

Morphologic changes during ontogeny, including changes in (1) shell pore characteristics (pore diameter, pore density, and porosity), (2) rates of increase in cross-sectional chamber area, (3) apertural position, (4) chamber surface ornamentation, and (5) umbilical diameter, were used to recognize ontogenetic stages in the foraminiferal shells. These include the prolocular, juvenile, neanic, and adult stages. The growth patterns of H. holmdelensis, H. monmouthensis, H. sliteri, and C. pilula are very uniform and do not show discernable transitions from the juvenile to neanic and adult stages. All four ontogenetic stages were recognized in A. australis, A. bosquensis, A. mateola, A. cretacea, and R. rugosa, although the abruptness of the transitions and the chamber number where these transitions occur are variable within and between species. Recognition of these growth stages enables taxonomic identification of pre-adult morphologies that occur in smaller size fractions. This is particularly useful since these smaller forms have dominated in unstable environments such as the highly seasonal circum-antarctic oceans.

New Specimens of the Pinnipediform Pteronarctos from the Miocene of Oregon
Annalisa Berta
30 pages, 21 figures
1994 (Date of Issue: 15 February 1994)
Number 78, Smithsonian Contributions to Paleobiology
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New fossils of the pinnipediform Pteronarctos from the Miocene Astoria Formation of coastal Oregon provide more information on the cranial anatomy, interrelationships, and higher level phylogeny of this taxon. Additional skulls are referred to a previously described species, P. goedertae (Barnes, 1989). A previously named species, Pteronarctos piersoni (Barnes, 1990) is recognized as the junior synonym of P. goedertae. The genus Pacificotaria (Barnes, 1992) is synonymized with Pteronarctos. The genus Pteronarctos is removed from the “Enaliarctinae” (= “Enaliarctidae”) and recognized as the sister group of the Pinnipedia (including Otariidae, Odobenidae, Allodesmus, Desmatophoca, Pinnarctidion, and Phocidae).

Ontogeny, Intraspecific Variation, and Systematics of the Late Cambrian Trilobite Dikelocephalus
Nigel C. Hughes
89 pages, 47 figures, 11 plates, 27 tables
1994 (Date of Issue: 29 November 1994)
Number 79, Smithsonian Contributions to Paleobiology
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Biometric analyses of well-localized specimens of the trilobite Dikelocephalus from the St. Lawrence Formation (Upper Cambrian), northern Mississippi Valley, suggest that all specimens belong to a single, highly variable morphospecies, D. minnesotensis. A complex pattern of ontogenetically-related and ontogeny-independent variation produced a mosaic of morphotypes, which show greater diversity than previously recorded within trilobite species. There is considerable variation within collections made from single beds. Variations of characters among collections are mosaic, and are clinal in some cases. Patterns of variation within Dikelocephalus cannot be related to lithofacies occurrence. There are no obvious temporal variations in D. minnesotensis within the St. Lawrence Formation, but some Dikelocephalus from the underlying Tunnel City Group may belong to a different taxon. The validity of this early taxon is questionable due to a lack of available material. The mosaic pattern of variation in Dikelocephalus mimics that documented at higher taxonomic levels in primitive libristomate trilobites, and helps explain difficulties in providing a workable taxonomy of primitive trilobites. Results caution proposition of evolutionary scenarios that do not take account of intraspecific variation. The recovery of dorsal shields of Dikelocephalus permits the first detailed reconstruction of the entire exoskeleton. The systematics of the genus is revised and twenty-five species are suppressed as junior synonyms of D. minnesotensis.

Bibliography and Index of the Sirenia and Desmostylia
Daryl Paul Domning
611 pages
1996 (Date of Issue: 25 July 1996)
Number 80, Smithsonian Contributions to Paleobiology
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The significant published literature on the neobiology, paleobiology, and ethnobiology of the mammalian orders Sirenia and Desmostylia is exhaustively cataloged in approximately 4590 main entries alphabetized by author. Both technical and popular works are included, and many entries are annotated. The earliest work cited is a letter by N. Syllacio published in 1494 or 1495, describing Columbus's second voyage to the New World. The effective closing date of the bibliography was 1 May 1994.

Six appendices list serial publications devoted to Sirenia, additional sources for history of sirenology and sirenian conservation, coins and postage stamps depicting sirenians, a comprehensive classification and synonymy of sirenians and desmostylians, a summary of the nomenclature of the Recent species of sirenians, and an alphabetical list of the species-group names that have been applied to sirenians and desmostylians.

An extensive index is provided, employing 1059 subject headings and cross references; the subject headings include all Linnaean names and combinations ever employed for sirenians and desmostylians, as well as names of all reported sirenian food plants and parasites. More than 40% of the main entries are fully indexed, and many others are partially indexed, yielding a total of over 13,950 index entries. Each complete index entry includes author and date of the work cited, a brief annotation describing the content of the work as it pertains to the indexed subject, and a page reference for the material pertaining to that subject.

Phylogenetic Revision of the Fish Families Luvaridae and †Kushlukiidae (Acanthuroidei), with a New Genus and Two New Species of Eocene Luvarids
Alexandre F. Bannikov and James C. Tyler
45 pages, 20 figures
1995 (Date of Issue: 18 May 1995)
Number 81, Smithsonian Contributions to Paleobiology
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Ten synapomorphies are described that support the proposed sister group relationship of the Eocene Kushlukiidae and the Eocene to Recent Luvaridae as the superfamilial clade Luvaroidea at the node between the siganid and zanclid+acanthurid clades of acanthuroid fishes. The Kushlukiidae previously have been known only on the basis of Kushlukia permira Danilchenko from the Eocene of Turkmenistan, but another species of that age is shown to have been present in India; the latter is not given a new specific name because the two specimens of it are only fragments. In addition to the well known Recent species Luvarus imperialis, three fossil species have been referred to the Luvaridae: Proluvarus necopinatus Danilchenko from the Eocene of Turkmenistan is here recognied as a valid species of Luvarus, with Proluvarus becoming a junior synonym of Luvarus; Eoluvarus bondei Sahni and Choudhary from the Eocene of India is shown to be not a luvarid but, rather, a member of the fossil perciform family Exelliidae, the affinities of which family are poorly understood; Luvarus praeimperialis Arambourg from the Oligocene of Iran is shown to be not a luvarid but, rather, a representative of the new genus Aluvarus of such uncertain affinity that we simply place it incertae sedis among the percomorphs.

Among the materials used by Danilchenko in the description of Proluvarus necopinatus only the holotype and the five other largest specimens (about 326-495 mm SL) represent that species. All of the other specimens are smaller than about 215 mm SL and represent two new species of a new genus of luvarid, Avitoluvarus dianae and A. mariannae.

Numerous derived features are used to define both the Kushlukiidae and Luvaridae. Within the Luvaridae, the preponderance of derived features is found in Luvarus, whereas Avitoluvarus has only two unequivocal synapomorphies.

The Luvaroidea, therefore, are represented by the Eocene Kushlukiidae with one genus and two species (Kushlukia permira and K. sp.) and the Luvaridae with one Eocene genus with two species (Avitoluvarus dianae and A. mariannae) and one genus, Luvarus, with one Eocene species, L. necopinatus, and one Recent species, L. imperialis.

New Superfamily and Three New Families of Tetraodontiform Fishes from the Upper Cretaceous: The Earliest and Most Morphologically Primitive Plectognaths
James C. Tyler and Lorenzo Sorbini
59 pages, 25 figures, 2 tables
1996 (Date of Issue: 15 April 1996)
Number 82, Smithsonian Contributions to Paleobiology
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Fishes of the order Tetraodontiformes previously have been known with assurance from as early as the Lower Eocene, about 55 MYA. Two Upper Cretaceous taxa, Protriacanthus d'Erasmo (1946) from Comen, Slovenia, about 90 MYA, and Plectocretacicus Sorbini (1979) from Hakel, Lebanon, about 95 MYA, were referred by their authors to the tetraodontiforms, respectively in the vicinity of triacanthoids and ostracioids. Both of these taxa were originally based on single specimens in which insufficient details of critical osteological features were exposed, and their familial and ordinal placement were open to question. Based on additional specimens now available, including acid preparations of several specimens of both species, we believe that they represent two new families (Protriacanthidae and Plectocretacicidae) of tetraodontiforms with numerous features that are more primitive than previously reported for the order. We describe a third taxon, the new genus Cretatriacanthus (and new family Cretatriacanthidae), based on a single specimen from the Upper Cretaceous of Nardò, Italy, about 70 MYA. It shares most of the numerous primitive features of Protriacanthus and Plectocretacicus.

Although many of their features are primitive, all three of these Upper Cretaceous taxa possess the most salient derived features of tetraodontiforms (e.g., reduced number of vertebrae, no anal-fin spines, reduced number of pelvic-fin rays, absence of certain skull bones, and, when present, long posterior process of pelvis with its halves in close contact or fused). The three Upper Cretaceous taxa share four derived features (absence of teeth, modified scales around base of pelvic spine, presence of subocular shelf, and diminutive size) that unite them in a clade herein recognized as the superfamily Plectocretacicoidea, whereas all other tetraodontiforms (those from the Lower Eocene to present) are united by seven derived features. Within the Plectocretacicoidea, five derived features support the sister-group relationship of Protriacanthus and Plectocretacicus.

Each of the three Upper Cretaceous taxa has a mosaic of primitive and specialized features, with some of the latter being independently derived relative to similar features of triacanthoids, balistoids, and ostracioids. The evidence indicates that the Plectocretacicoidea (presently known from 70 to 95 MYA) are the morphologically primitive sister group of all other tetraodontiforms (presently known from 55 MYA to Recent).

Oligocene Echinoids from North Carolina
Porter M. Kier
37 pages, 6 figures, 11 plates, 2 tables
1997 (Date of Issue: 6 May 1997)
Number 83, Smithsonian Contributions to Paleobiology
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Oligocene echinoids are rare, which makes important this material from three quarries in North Carolina. Three species occur in the state quarry at Pollocksville: Psammechinus carolinensis, new species, Maretia carolinensis, new species, and Agassizia sp. At the New Bern quarry occur Rhyncholampas gouldii (Bouvé) newbernensis, new subspecies, Agassizia mossomi Cooke, Psammechinus carolinensis, new species, Dixieus dixie (Cooke), Clypeaster rogersi (Morton), and Maretia sp. Periarchus lyelli (Conrad) is found in underlying Eocene beds. The Belgrade quarry fauna consists of Arbia aldrichi (Clark), Gagaria mossomi (Cooke), Echinocyamus wilsoni, new species, and Agassizia mossomi Cooke. These echinoids indicate a middle to late Oligocene age for the Trent Formation (= River Bend) and Belgrade Formation (= River Bend).

Relationships of the Fossil and Recent Genera of Rabbitfishes (Acanthuroidei: Siganidae)
James C. Tyler and Alexandre F. Bannikov
35 pages, 21 figures, 1 table
1997 (Date of Issue: 2 September 1997)
Number 84, Smithsonian Contributions to Paleobiology
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Four genera of fossil siganid fishes of early Eocene to early Oligocene age are recognized in addition to the single Recent genus. The osteological features of these five genera are described and illustrated. A phylogenetic analysis utilizing PAUP indicates that the genera have the following phyletic sequence convention: Ruffoichthys Sorbini (two species from the middle Eocene of Italy)—Eosiganus, new genus (one new species from the middle Eocene of Russia)—Siganopygaeus Danilchenko (one species from the early Eocene of Turkmenistan)—Protosiganus Whitley (one species from the early Oligocene of Switzerland)—Siganus Forsskål (27 Recent species in the Indo-Pacific).

Atlas of Paleocene Planktonic Foraminifera
Richard K. Olsson, Christoph Hemleben, William A. Berggren and Brian T. Huber, editors
252 pages, 37 figures, 71 plates
1999 (Date of Issue: 3 March 1999)
Number 85, Smithsonian Contributions to Paleobiology
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Sixty-seven species of Paleocene planktonic foraminifera are described and illustrated, including three species of Eoglobigerina, four species of Parasubbotina, five species of Subbotina, two species of Hedbergella, 10 species of Globanomalina, six species of Acarinina, 12 species of Morozovella, three species of Igorina, four species of Praemurica, one species of Guembelitria, one species of Globoconusa, three species of Parvularugoglobigerina, two species of Woodringina, six species of Chiloguembelina, one species of Rectoguembelina, and four species of Zeauvigerina. Taxonomic classification of normal perforate taxa are organized according to wall texture. Spinose cancellate genera include Eoglobigerina, Parasubbotina, and Subbotina; cancellate nonspinose genera include Igorinina and Praemurica; smooth-walled genera include Hedbergella and Globanomalina; and muricate genera include Acarinina and Morozovella. Taxonomic classification of microperforate taxa (including Guembelitria, Globoconusa, Parvularugoglobigerina, Woodringina, Chiloguembelina, Rectoguembelina, and Zeauvigerina) are organized according to test morphology.

Scanning electron microscope (SEM) images of type species described by Morozova in the collections of the Geological Institute, Academy of Sciences (GAN), Moscow, and the type material described by Subbotina in the collections of the All Union Petroleum Scientific Research Geological Prospecting Institute (VNIGRI), St. Petersburg, are shown on Plates 8-12. Twelve species described by Morozova, nine species described by Subbotina, and one species described by Bykova are illustrated. In addition, SEM images of 28 holotypes and two paratypes from the Smithsonian Institution collections are shown on Plates 13-17, and the lectotype for Globigerina compressa Plummer, 1926, and the neotype for Globorotalia monmouthensis Olsson, 1961, are designated and illustrated with SEM images.

Paleobiogeographic maps showing the global distribution of 29 commonly occurring Paleocene taxa are included in the atlas, as well as figures showing the stratigraphic ranges of species by genus and stratigraphic first and last appearances. The biostratigraphic framework used in the atlas is the revised biostratigraphy given in Berggren et al., 1995, which is summarized in the atlas. Wall texture and morphological relationships between species and genera form the basis of phylogenetic interpretations. This is discussed in the section “Wall Texture, Classification, and Phylogeny” and is referenced to Plates 1-7.

Reflections on the Morphology, Anatomy, Evolution, and Classification of the Class Stenolaemata (Bryozoa)
Richard S. Boardman
59 pages, 129 figures
1998 (Date of Issue: 26 August 1998)
Number 86, Smithsonian Contributions to Paleobiology
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Thin sections of 37 stenolamate species with soft parts within the skeletons unexpectedly reveal 10 variations in polypide anatomy and related methods of tentacle protrusion. Six of the variations modify attachment organs, and four occur in species that lack such organs. Attachment organs are required for the majority of “progressive” degeneration-regeneration polypide cycles that advance new polypides as the skeletons grow, so that functionally constant protrusion distances are maintained. In most species lacking attachment organs, polypides retract and regenerate in the bottoms of their living chambers so polypide cycles are “stationary” and protrustion distances increase as the skeletons grow in length.

Internal studies indicate that skeletal microstructure and polypide anatomy in Recent stenolaemates are generally qualitatively uniform within colonies and within species. Unexpectedly, skeletal microstructure and polypide anatomy often do not correlate with each other or with the skeletal structures of the five informal taxa (horneroids, tubuliporines, fasciculates, disporelloids, and heteroporoids) used to group Recent species. These groupings have been established historically using only the relatively few external characters available. Internally, four of the five groupings have more than one kind of attachment organ, and three of the five have polypides both with and without attachment organs. Furthermore, organs that apparently occur in relatively few taxa occur in more than one grouping. Skeletally, more than one microstructure occurs in four of the five groupings. Noncorrelations of internal character states are so numerous that it seems almost any polypide can occur in almost any skeleton.

Genetically controlled taxonomic characters of fossil stenolaemates that seem promising in ellucidating relationships are skeletal microstructure, skeletal structures that reflect polypide anatomy and function, such as nutrient exchange systems, and mode of growth. Species in which laminae in the vertical zooidal walls grew inward from zooidal boundaries under outer body cavities dominated throughout the Paleozoic and Triassic and that inward-growing microstructure has continued to the present. Communication pores and frontal walls developed and disappeared in a meager number of species that became extinct in the Paleozoic. Species that grew laminae outwardly from zooidal boundaries, or grew crystallites at right angles to zooidal growth, apparently appeared first in the Jurassic. Three Paleozoic orders are inferred to have survived into the Jurassic, and, with the new Mesozoic clades, to have independently evolved communication pores and frontal walls. The three surviving Paleozoic orders are inferred to have interchanged primitive characteristics with the advanced features of newly evolved Mesozoic clades to produce the detailed noncorrelations described here in Recent species. Post-Triassic stenolaemates, therefore, are inferred to be polyphyletic at the order level.

With just 37 Recent species available, it is not clear whether polythetically described taxa ultimately can be distinguished in a manner that adequately reflects phylogenetic patterns in younger stenolaemates. Differing, but functionally similar, parts might have been so pervasive across clades that the resulting noncorrelations have produced a mosaic distribution of combined primitive and advanced character states that makes any sort of natural classification unlikely.

Ostracoda from the Late Permian of Greece (Thaumatocyprididae and Polycopidae)
I. G. Sohn and Louis S. Kornicker
34 pages, 20 figures, 1 map, 2 tables
1998 (Date of Issue: 26 August 1998)
Number 87, Smithsonian Contributions to Paleobiology
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The ornamentation of the outer surface of the carapace of both living and fossil thaumatocyprids is compared and discussed. Seven new species of Thaumatomma Kornicker and Sohn, 1976, one species of Thaumatomma in open nomenclature, and a new species of Polycope from the Late Permian of the islands of Hydra and Salamis, Greece, are described and illustrated. A key to the species of Thaumatomma is presented.

Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods
Peter J. Wagner
152 pages, 37 figures, 3 tables
2002 (Date of Issue: 30 January 2002)
Number 88, Smithsonian Contributions to Paleobiology
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In order to explore the phylogenetic relationships among early gastropods, cladistic analyses were conducted of nearly 300 “archaeogastropod” species known from the latest Cambrian through the Silurian. The study includes an extended outgroup analysis of Cambrian molluscs. The resulting estimates of gastropod phylogeny differ not only from traditional ideas about early gastropod relationships, but also from most alternative notions. Outgroup analyses suggest that gastropods had ancestors among the Tergomya (= Monoplacophora of many workers) of the Middle or Late Cambrian. Putative gastropods from older strata (e.g., the Pelagiellida and early Onychochilidae) apparently are not closely related to gastropods. The hypothesized ancestor of gastropods possessed dextral-coiling, septation, a deep sinus, and a peripheral band. An anal slit is commonly described as a synapomorphy of gastropods that many clades subsequently lost; however, this study suggests that the slit is a rare, highly derived, and polyphyletic character among early Paleozoic species, and that the ancestors of most “advanced” clades (e.g., the Apogastropoda) never had slits.

This study suggests that two major subclades evolved by the earliest Ordovician. The diagnoses and definitions of these two subclades best correspond to the traditional diagnoses and definitions of the Euomphalina and Murchisoniina. The Pleurotomarioidea is not a paraphyletic ancestral taxon as typically suggested, but instead it is a polyphyletic assemblage derived multiple times from “euomphalinae” and “murchisoniinae” species. The Bellerophontina is at least diphyletic, as the taxon includes both the ancestors of “archaeogastropods” and a clade of planispiral species that is secondarily derived from “archaeogastropods.” Macluritoids sensu stricto represent a restricted subclade of the “euomphalinae”; other supposed macluritoids evolved among different euomphalinae subclades or are not gastropods. Early Paleozoic species previously classified as caenogastropods (i.e., the Loxonematoidea and Subulitoidea) represent separate murchisoniinae subclades, with some putative members of the Subulitoidea derived within the Loxonematoidea. Early Paleozoic species assigned to the Trochoidea also represent several subclades, with most of those clades having evolved from the “euomphalinae.”

An extensive taxonomic revision is presented, which removes all early Paleozoic taxa from the Pleurotomariina and broadly expands the definitions of the Euomphalina and Murchisoniina.

Avian Paleontology at the Close of the 20th Century: Proceedings of the 4th International Meeting of the Society of Avian Paleontology and Evolution, Washington D.C., 4-7 June 1996
Storrs L. Olson, editor
344 pages, 169 figures, 49 tables
1999 (Date of Issue: 14 December 1999)
Number 89, Smithsonian Contributions to Paleobiology
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The 32 papers collected herein reflect the great diversity and interest that the study of fossil birds has generated in recent years. The first seven papers (Mourer-Chauviré et al., Worthy and Jouventin, Seguí and Alcover, Steadman and Hilgartner, Millener, Worthy, Pavia) relate to late Quaternary birds from islands, where human intervention in the last few thousand years has caused many heretofore unrecorded extinctions. Three papers on Quaternary avifaunas of continental Europe deal with distributional changes and cultural use of birds by humans in Siberia (Potapova and Panteleyev), the utility of patterns of seabird distribution in determining former marine climatic conditions (Tyrberg), and temporal changes in morphology of ptarmigans (Lagopus) through the late Pleistocene (Stewart). Three papers deal with late Cenozoic raptors (Campbell et al., Tambussi and Noriega, Emslie and Czaplewski). New genera from Paleogene deposits are described by Boles and Ivison, Karhu, and Peters. Five papers deal with ancient waterfowl. Alvarenga describes the first fossil screamer (Anhimidae) from the Oligocene of Brazil. Olson provides the first fossil records of the Anseranatidae, with the description of a new species from the early Eocene of England, which is referred to Anatalavis from the Paleocene/Cretaceous of New Jersey. Ericson provides the means to distiguish Eocene fossils of the duck-like Presbyornis from the flamingo-like Juncitarsus and gives new records of the latter. Benson shows that the Paleocene Presbyornis isoni once ranged from Maryland to North Dakota, and he gives records of other Paleocene birds from North Dakota. Hope names a new, larger species of Graculavus, extending the range of the genus from New Jersey to the Cretaceous of Wyoming.

The early history and evolution of birds receives great attention. Dzerzhinsky expands upon the significance of cranial morphology in paleognathous birds. Kurochkin relates the early Cretaceous genus Ambiortus to the Chinese Otogornis, which are supposed to be on a line with modern birds, as opposed to the Enantiornithes. Bochenski uses paleogeography to suggest that the Enantiornithes must antedate Archaeopteryx. Zhou and Martin show that the manus of Archaeopteryx is more bird-like than previously realized. Martin and Stewart use bird teeth to argue against dinosaurian origins for Aves, whereas Elzanowski diverges on various aspects of dinosaurian cranial morphology and that of early birds that may have evolutionary significance. Witmer, Chiappe, and Goslow present summaries of three sessions of a roundtable discussion on avian origins, early evolution of birds, and the origins of flight, which was held on June 7, the last day of the meeting, and which covered much controversial territory.

Geology and Paleontology of the Lee Creek Mine, North Carolina, III
Clayton E. Ray and David J. Bohaska, editors
365 pages, 127 figures, 45 plates, 32 tables
2001 (Date of Issue: 11 May 2001)
Number 90, Smithsonian Contributions to Paleobiology
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This volume on the geology and paleontology of the Lee Creek Mine is the third of four to be dedicated to the late Remington Kellogg. It includes a prodromus and six papers on nonmammalian vertebrate paleontology. The prodromus continues the historical theme of the introductions to volumes I and II, reviewing and resuscitating additional early reports of Atlantic Coastal Plain fossils. Harry L. Fierstine identifies five species of the billfish family Istiophoridae from some 500 bones collected in the Yorktown Formation. These include the only record of Makaira purdyi Fierstine, the first fossil record of the genus Tetrapturus, specifically T. albidus Poey, the second fossil record of Istiophorus platypterus (Shaw and Nodder) and Makaira indica (Cuvier), and the first fossil record of I. platypterus, M. indica, M. nigricans Lacépède, and T. albidus from fossil deposits bordering the Atlantic Ocean. Robert W. Purdy and five coauthors identify 104 taxa from 52 families of cartilaginous and bony fishes from the Pungo River and Yorktown formations. The 10 teleosts and 44 selachians from the Pungo River Formation indicate correlation with the Burdigalian and Langhian stages. The 37 cartilaginous and 40 bony fishes, mostly from the Sunken Meadow member of the Yorktown Formation, are compatible with assignment to the early Pliocene planktonic foraminiferal zones N18 or N19. The Pungo River fish fauna is dominated by warm water taxa; the Yorktown fauna includes warm and cool water species. These changes are attributed to increased upwelling waters in Yorktown time. The abundant fossils provide the basis for several changes in selachian taxonomy and for two new species of bony fishes. George R. Zug records 11 taxa of turtles from the Yorktown Formation: a sideneck (Bothremys); six sea turtles (Caretta, ?Chelonia, Lepidochelys, Procolpochelys, Psephophorus, Syllomus); a softshell turtle (trionychid); two pond turtles (probably Pseudemys and Trachemys); and a giant tortoise (Geochelone). Albert C. Myrick, Jr., records the crocodylian Thecachampsa antiqua (Leidy) on the basis of fragmentary float material from the Pungo River or Yorktown Formation, or both. Robert W. Storer describes a new species of grebe of the genus Podiceps from the Yorktown Formation. Storrs L. Olson and Pamela C. Rasmussen record some 112 species of birds from the Pungo River and Yorktown formations. Apart from an undetermined number of shearwaters, only a few species are thought to come from the Pungo River Formation. The marine species from the Yorktown Formation include three loons, two grebes, five albatrosses, at least 16 shearwaters and petrels, one pelican, two pseudodontorns, three gannets, two cormorants, 9-11 auks and puffins, one skua, three jaegers, five gulls, two terns, and 20 ducks, geese, and swans. The less common land and shore birds are represented by 29 species, including three cranes, one rail, two oystercatchers, one plover, four scolopacids, one flamingo, one ibis, one heron, three storks, one condor, five accipitrids, one osprey, one phasianid, one turkey, one pigeon, and one crow. The fauna is dominated by a radiation of auks of the genus Alca. The early Pliocene fauna is very modern in aspect, suggesting that most modern lineages of birds were already in existence.

Myodocopid Ostracoda from the Late Permian of Greece and a Basic Classification for Paleozoic and Mesozoic Myodocopida
Louis S. Kornicker and I. G. Sohn
33 pages, 22 figures, 1 table
2000 (Date of Issue: 1 May 2000)
Number 91, Smithsonian Contributions to Paleobiology
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Four new genera and six new species are described from the top of the Episkopi Formation (Dorashamian) on the island of Hydra, Greece: Cypridinelliforma rex (new species), Nodophilomedes phoenix (new genus, new species), Swainella bex (new genus, new species), Triadocypris pax (new species), Siveterella pax (new genus, new species), Siveterella flex (new species), and Sylvesterella (new genus), based on specimens in the collection from Greece. Supplementary descriptions are presented of Philomedes rankiniana (Jones and Kirkby, 1867) and Eocypridina radiata (Jones and Kirkby, 1874).

A basic classification proposed for Paleozoic and Mesozoic Myodocopida includes a new suborder, three new superfamilies, and three new families.

Triassic Gastropods of the Southern Qinling Mountains, China
Jinnan Tong and Douglas H. Erwin
47 pages, 11 figures, 6 plates, 5 tables
2001 (Date of Issue: 7 November 2001)
Number 92, Smithsonian Contributions to Paleobiology
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Forty-eight species in 27 genera of gastropods, including 14 new species and one new genus, are described from early- to middle-Triassic (Scythian- to Ladinian-aged) rocks from the southern Qinling Mountains of Gansu and Sichuan provinces, China. This report expands the knowledge of the biogeographic distribution of gastropods during the recovery from the end-Permian mass extinction. The new taxa include Tongweispira sichuanensis, new genus and new species, and the following new species: Ananias guojiashanensis, Worthenia extendia, Gosseletina? dangchangensis, Zygites laevigatus, Trochotoma (Discotoma) gansuensis, Cheilotomona acutocarinata, Naticopsis (Dicosmos) compressus, Naticopsis (Discosmos) sichuanensis, Naticopsis? ribletella, Neritopsis planoplicatus, Platychilina sinensis, Platychilina obliqua, and Omphaloptycha gansuensis.

Cenozoic Mammals of Land and Sea: Tributes to the Career of Clayton E. Ray
Robert J. Emry, ed.
v, 372 p. : ill., maps
Number 93, Smithsonian Contributions to Paleobiology
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This is a volume of collected papers published to honor the career of Clayton E. Ray, now Curator Emeritus in the Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, and Curator of Late Cenozoic Mammals and of Fossil Marine Mammals in the same department for more than 30 years before his retirement in 1994. The volume includes a preface, a biography and bibliography of Clayton E. Ray, and 19 papers devoted principally to Pleistocene mammals and to fossil marine mammals. Gary Morgan describes late Pleistocene mammalian faunas from several sites in southernmost Florida and discusses the Neotropical influence in Florida's Pleistocene faunas. Richard H. Tedford describes the basicranium of the Pleistocene giant wombat Phascolonus gigas Owen and discusses its significance in marsupial phylogenetic reconstruction. Gerardo De Iuliis and A. Gordon Edmund describe Vassallia maxima Castellanos, the only pre-Pleistocene pampathere known in which a skull and mandible are associated with osteoderms; the range of osteoderm variation in one associated individual allows them to synonymize other taxa that had been based on osteoderm differences. Paul W. Parmalee and Russell Wm. Graham report additional records of the giant beaver, Castoroides, from the mid-South. Frederick Grady, Joaquin Arroyo-Cabrales, and E. Ray Garton report the northernmost known occurrence of vampire bats in the Pleistocene of eastern North America. H. Gregory McDonald reports the second known occurrence of the badger Taxidea taxus in the Pleistocene of Kentucky and discusses the paleoecological implications of the occurrence. Jerry N. McDonald and George E. Lammers describe Bison antiquus from Ontario and discuss the evolution of bison in the Holocene of North America. Daryl P. Domning presents a new analysis and interpretation of the terrestrial posture in desmostylians. Thomas A. Demere and Annalisa Berta describe new material and present a phylogenetic analysis of the Miocene pinniped Desmatophoca oregonensis from Oregon. Irina A. Koretsky and Dan Grigorescu describe and evaluate the systematic position of the fossil monk seal Pontophoca sarmatica from the Miocene of eastern Europe. Irina A. Koretsky and Peter Holec describe a new, primitive, phocid pinniped from the early middle Miocene of Slovakia and discuss its bearing on the phylogeny and classification of pinnipeds. Irina A. Koretsky and Albert E. Sanders report remains of the oldest known phocid pinniped from the late Oligocene of South Carolina. R. Ewan Fordyce describes and discusses a bizarre archaic Oligocene dolphin from the eastern North Pacific, on which he bases a new species, genus, and subfamily. Christian de Muizon, Daryl P. Domning, and Darlene R. Ketten describe and discuss the paleobiology and behavior of an unusual walrus-convergent delphinoid cetacean from the early Pliocene of Peru. Susan D. Dawson and Michael D. Gottfried report paleopathologic conditions in a Miocene odontocete cetacean. Albert E. Sanders and Lawrence G. Barnes contribute two papers, both describing and analyzing new, primitive, cetotheriid mysticete cetaceans from the late Oligocene of South Carolina. James W. Westgate and Frank C. Whitmore, Jr., describe a new species of bowhead whale from the Pliocene Yorktown Formation in Virginia. James G. Mead and Rosemary G. Dagit present an account of the search for the 1880s manuscript of J.A. Allen's unpublished monograph on the mammalian orders Cete and Sirenia; the manuscript was not found but the 12 plates that were prepared for it are published herein.

Middle Proterozoic (1.5 Ga) Horodyskia moniliformis Yochelson and Fedonkin, the Oldest Known Tissue-Grade Colonial Eucaryote
Mikhail A. Fedonkin and Ellis L. Yochelson
29 pages, 19 figures
2002 (Date of Issue: 29 January 2002)
Number 94, Smithsonian Contributions to Paleobiology
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“Problematic bedding-plane markings” discovered by the late R.J. Horodyski from the Appekunny Formation in Glacier National Park, Montana, and dated at approximately 1.5 giga-annum (Ga), were never formally named. We are convinced the specimens are biogenic and have placed them within Linnaean nomenclature as Horodyskia moniliformis Yochelson and Fedonkin. An apt description of the locally abundant fossils is “string of beads.” On each string, beads are of nearly uniform size and spacing; proportionally, bead size and spacing remain almost constant, regardless of string length or size of individual beads. They may not be related to any other known fossil, and their position within highest levels of the taxonomic hieararchy is enigmatic. We judge they were multicellular, tissue-grade, colonial eucaryotes. Similar strings have been reported from Western Australia, but nowhere else. The general geologic setting in Montana, details of sedimentation, and taphonomy suggest the organisms were benthonic, growing upward about 1 cm through episodically deposited eolian dust. During life, specimens were stiff and relatively strong, but show no evidence of a mineralized skeleton. They lived in poorly oxygenated water with the body progressively subjected to anaerobic conditions. Their energy source is obscure; their mode of growth and several features of interpreted environment lead us to speculate that Horodyskia likely lived primarily by ingesting chemosynthetic bacteria rather than by photosynthesis. This notion should be tested by searching red, fine-grained, subaqueous arenites of approximately the same age throughout the world for additional occurrences.

New Materials of Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001, and Implications for the Morphology of the Noasauridae (Theropoda: Ceratosauria)
Matthew T. Carrano, Mark A. Loewen, and Joseph J. W. Sertich
viii + 53 pages
2011 (Date of Issue: 18 January 2011)
Number 95, Smithsonian Contributions to Paleobiology
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Osteology of the noasaurid theropod Masiakasaurus knopfleri Sampson et al., 2001, is now two-thirds complete. We describe Masiakasaurus knopfleri in detail on the basis of examination of new specimens and emphasis on previously unknown elements. The skull is anteroposteriorly long but low in height, unlike the foreshortened abelisaurid condition. Premaxillary teeth are procumbent, like those of the dentary. Frontal bones are flat and unornamented, but the lacrimal and postorbital exhibit surface texturing. The braincase resembles that of abelisaurids but is more highly pneumatized. The neck is curved anteriorly but horizontal posteriorly, and it transitions to the trunk without significant proportional changes. Centrum pneumaticity appears confined to the neck and anterior trunk. The sacrum includes six vertebrae, and the expanded transverse processes of caudal vertebrae may articulate with caudal ribs. The scapulocoracoid is large and broad. The ilium is both anteroposteriorly long and dorsoventrally deep, and it bears pegs for articulation with sockets on the pubis and ischium, as in other ceratosaurs. The nearly complete pes shows no particular locomotor specializations and allows reinterpretation of the “raptorial” pedal ungual of Noasaurus as a manual element. These new specimens also illuminate the morphology of other noasaurids, especially those from the Lameta Formation.

In addition to Madagascar, noasaurids are known from Europe, India, South America, and Africa, spanning at least Aptian–Albian through Maastrichtian time. The new materials of Masiakasaurus increase character resolution within Abelisauroidea, identifying many formerly equivocal features as synapomorphies of the nodes Noasauridae, Abelisauridae, or Abelisauroidea. Unfortunately, the fragmentary nature of nearly all other noasaurids obviates any meaningful ingroup resolution, and as a result no particular evolutionary or biogeographic scenarios for the clade can presently be supported (or rejected) with confidence.

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