Smithsonian Contributions to Paleobiology

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New Permian Brachiopods from West Texas
G. Arthur Cooper and Richard E. Grant
20 pages, 5 plates
1969 (Date of Issue: 14 July 1969)
Number 1, Smithsonian Contributions to Paleobiology
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Thirty-five new genera are described, 27 of them based on new species. They are classified by superfamily as follows: ENTELETACEA: Acosarina (A. dorsisulcata, new species). DAVIDSONIACEA: Goniarina (G. pyelodes, new species), Tropidelasma (T. culmenatum, new species). CHONETACEA: Chonetinetes (C. reversus, new species), Micraphelia (M. scitula, new species), Rugaria (Chonetes hessensis R. E. King), Sulcataria (Chonetina? rostrata Dunbar and Condra), Undulella (U. undulata, new species). STROPHALOSIACEA: Acritosia (A. magna, new speccies), Agelesia (Aulosteges triagonalis R. E. King). RICHTHOFENIACEA: Collumatus (C. solitarius, new species), Cyclacantharia (C. kingorum, new species), Hercosestria (H. cribrosa, new species), Hercosia (Richthofenia uddeni Böse), Sestropoma (S. cribriferum, new species). PRODUCTACEA: Anemonaria (A. inflata, new species), Dasysaria (D. undulata, new species), Oncosarina (O. spinicostata, new species), Thamnosia (T. anterospinosa, new species). LYTTONIACEA: Petasmaia (P. expansa, new species). RHYNCHONELLACEA: Amphipella (A. arcaria, new species), Bryorhynchus (Camarophoria? bisulcata Shumard) Divaricosta (D. squarrosa, new species), Petasmatherus (P. opulus, new species), Phrenophoria (P. subcarinata, new species), Pontisia (P. stehlii, new species), Strigirhynchus (Rhynchonella? indentata Shumard), Tautosia (T. fastigiata, new species). SPIRIFERACEA: Lepidospirifer (L. angulatus, new species). SPIRIFERINACEA: Sarganostega (S. transversalis, new species), Xestotrema (Spirifera pulchra Meek). RETICULARIACEA: Anomaloria (A. anomala, new species), Astegosia (Squamularia guadalupensis subquadrata Girty). DIELASMATACEA: Plectelasma (P. kingi, new species). CRYPTONELLACEA: Texasia (T. elongata, new species).

The genus Cooperina Termier, Termier and Pajaud, based on material from the Glass Mountains, Texas, was assigned by its authors to the Thecideidina. Evidence is adduced here to show that it is classified more properly among the Strophalosiacea of the Productidina and that it is unrelated to the thecideids.

A New Occurrence of Paleocene Mammals in the Evanston Formation, Southwestern Wyoming
C. Lewis Gazin
17 pages, 1 figure, 3 plates
1969 (Date of Issue: 31 December 1969)
Number 2, Smithsonian Contributions to Paleobiology
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A new fossil horizon and locality for the Evanston formation, near Little Muddy Creek in the Fossil Basin of southwestern Wyoming, has yielded remains of a mammalian faunule of middle Paleocene age. Relationships are shown to the two widely separated classic occurrences: the upper Lebo of the Montana Fort Union and the New Mexico Torrejon. A correlation is also indicated with the Battle Mountain and Rock Bench occurrences in the more local Wyoming region. The known faunule is comprised of essentially small Mammalia representing the orders Multituberculata, Insectivora, Primates, Creodonta, and Condylarthra. The material consists for the most part of isolated teeth, hence identifications are necessarily tentative in nature.

Paleozoic Perspectives: A Paleontological Tribute to G. Arthur Cooper
J. Thomas Dutro, Jr., editor
390 pages, 78 figures, 63 plates, 16 tables
1971 (Date of Issue: 22 February 1971)
Number 3, Smithsonian Contributions to Paleobiology
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This collection of papers was solicited from colleagues and students of G. Arthur Cooper as a Festchrift in recognition of his profound influence on the study of brachiopods and their biostratigraphic application to geologic problems, especially in the Paleozoic Era.

Dr. Cooper initiated a period of growth in both the research staff and the National Collections of Fossils that guided the Smithsonian Institution to its present position of leadership in paleontological research. His own superb studies of fossil and living brachiopods are unsurpassed in breadth and paleontological significance. Mainly through his efforts, the Smithsonian has acquired an outstanding reference collection of invertebrate fossils that is the envy of the scientific community.

Distribution of Planktonic Foraminifera in the Vicinity of the North Atlantic Current
Richard Cifelli and Roberta K. Smith
52 pages, 22 figures, 6 plates, 8 tables
1970 (Date of Issue: 13 April 1970)
Number 4, Smithsonian Contributions to Paleobiology
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Planktonic Foraminifera collected from the vicinity of the North Atlantic Current and the Gulf Stream during late winter-early spring and fall of 1964 are described and their distributions are recorded. Variations in faunal composition seem to be related largely to water regime dynamics and seasonal cycle. Among the fall collections, three distinctive assemblages can be recognized: a western group in the vicinity of the Gulf Stream, containing predominantly Sargasso Sea-Gulf Stream species dominated by Globigerinoides ruber; a northern group, dominated by Globigerina quinqueloba egelida, new subspecies, reflecting the influence of cold, northern waters adjacent to the North Atlantic Current; and an eastern group, dominated by Globigerina incompta, apparently developed within the limits of the North Atlantic Current. The last group seemingly represents an anomaly, as North Atlantic Current surface temperatures were relatively high at the time of collection, and dominance of a warm-water form, such as Globigerinoides ruber, might have been expected. The anomaly suggests that the North Atlantic Current is a partially closed gyre, fed by both slope waters and Gulf Stream. Temperatures are considered to be close to threshold for both cold and warm-water species.

Distributional patterns displayed by the late winter-early spring collections are compatible with the proposed model. Also, these collections, taken over a period of almost three months, reflect marked seasonal changes in faunal composition, particularly in Sargasso Sea-Gulf Stream elements.

Twenty-five species and subspecies are described. One species, Globigerina atlantisae, and one subspecies, Globigerina quinqueloba egelida, are new.

Catalog of the Illustrated Paleozoic Plant Specimens in the National Museum of Natural History
Arthur D. Watt
53 pages
1970 (Date of Issue: 17 September 1970)
Number 5, Smithsonian Contributions to Paleobiology
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A catalog of the illustrated Paleozoic fossils in the National Museum of Natural History is presented, updating United States National Museum Bulletin 53, part 2, section 3, 1907.

Functional Morphology and Biofacies Distribution of Cheilostome Bryozoa in the Danian Stage (Paleocene) of Southern Scandinavia
Alan H. Cheetham
87 pages, 29 figures, 17 plates, 10 tables
1971 (Date of Issue: 27 September 1971)
Number 6, Smithsonian Contributions to Paleobiology
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Highly diversified assemblages of cheilostome Bryozoa in the Danian Stage of southern Sweden and Denmark represent the culmination of primarily divergent evolutionary trends originating in the first appearance of the group in Early Cretaceous time. Functional relationships between colony and zooid morphology are less likely to have been obscured by vestigial structures and convergent and parallel evolution in these assemblages than in later Cenozoic faunas. The Danian assemblages, then, provide a test of the hypothesis that, in the early evolution of cheilostomes, environmentally correlated variation in the form of colonies depended functionally upon the structure of their component zooids.

Theoretically, the rigidly erect growth form should have an adaptive advantage over the presumed ancestral encrusting form, by virtue of a vastly increased potential zooid density relative to substrate occupied. A rigidly erect colony must be able to resist stresses induced by vertical loading, bending, and twisting and thus appears to require calcified walls, especially on the frontal sides of its zooids. Given the constraints imposed by the cheilostome mode of growing and calcifying zooid walls and of operating the hydrostatic system, zooid morphotypes can be relatively graded for efficiency in structural support of the colony by the degree to which their joint calcification approaches a laterally merging, continuously thickening, distally tapering skeletal mass analogous to the outer walls of an enlarging cantilever beam.

These hypothetical relationships are generally consistent with biofacies distributions of more than 50 species associated with a single middle Danian mound in southern Sweden. This mound is typical of many which accumulated, probably at depths approximating the shelf-edge, in southern Scandinavia during Danian time. It includes three biofacies: (1) the flanks, dominated by bryozoans; (2) the core, rich in octocorals with less abundant colonial scleractinians and bryozoans; and (3) transitional areas, between the two, dominated by octocorals but with abundant bryozoans. Sediments of the three biofacies contain distinctive assemblages of cheilostome species which differ in abundance rather than by presence or absence. The flanks are dominated by species inferred to have had erect colonies and the more complex zooid morphotypes. This group of species constitutes the bulk of the total fauna in weight-abundance but fewer than half the species. Species dominant in the core facies make up about half the total number of species and are inferred to have had mostly encrusting colonies with zooids of all morphotypes recognized, including the simplest. The transitional facies includes a mixture in subequal proportions of the two groups of species dominant in the other facies; however, this facies has other distinctive species in abundance and thus may represent an ecotone. Morphologically, the cheilostomes abundant in the transitional facies are intermediate in inferred zooid morphotypes and colony forms.

The relation between abundance and morphology of Danian cheilostomes suggests that attainment of the more advantageous rigidly erect colony form was functionally more probable for zooid morphotypes susceptible of heavy frontal calcification than for others. If a minimum amount of frontal calcification must have been present before the rigidly erect mode of growth could be assumed, then frontal calcification was associated originally with some other function, such as protection of the lophophore. It is possible that the various further advances in zooid morphotype could also have been made as separate prospective adaptations, but it seems more likely that some or all of them represent direct adaptive improvements for the structural support of rigidly erect colonies.

A New Cenozoic Deep-Sea Genus, Abyssocythere (Crustacea: Ostracoda: Trachyleberididae), with Descriptions of Five New Species
Richard H. Benson
25 pages, 12 figures, 3 plates, 1 table
1971 (Date of Issue: 11 August 1971)
Number 7, Smithsonian Contributions to Paleobiology
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The new genus Abyssocythere Benson has been erected to receive five new species and one described species of psychrospheric ostracode ranging in age from the Paleocene to the Recent. These species include Abyssocythere casca Benson, new species (Indian Ocean), herein designated the type species, A. pannucea Benson, new species (eastern Pacific), A. japonica Benson, new species (western Pacific), A. atlantica Benson, new species (Atlantic), A. australis (Southern Ocean), and A. trinidadensis (van den Bold) from the Caribbean region. Modern species are typical of the deep-sea floor and are common to depths below 2,000 meters. Fossils have been found in deep-water Paleocene and Miocene strata in Trinidad and in deep-sea cores in Pleistocene sediments. These species are thought to have descended from a yet unknown form of the complex of species assigned to the Cretaceous genus Cythereis (sensu lato). There seems to be an evolution of finer surface features of the carapace and an increase in size throughout the Cenozoic.

Mode of Growth and Functional Morphology of Autozooids in Some Recent and Paleozoic Tubular Bryozoa
Richard S. Boardman
51 pages, 6 figures, 11 plates
1971 (Date of Issue: 23 August 1971)
Number 8, Smithsonian Contributions to Paleobiology
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Membranous structures reflecting functional organs are recognizable in a relatively few tubular Bryozoa of Paleozoic age belonging largely to the order Trepostomata. Some skeletal structures also seem to reflect functional organs in a generalized way. Thin sections, including both hard and soft parts, of several genera of Recent tubular Bryozoa of the order Cyclostomata provide a first approximation to the shape, size, and position of cuticular or membranous structures in autozooids that might be preserved under exceptional conditions in fossils. Potentially preservable cuticular or membranous structures include: (1) outward opening funnel-shaped terminal-vestibular membranes and sphincter muscle regions; (2) flask- or sac-shaped membranous sacs; and (3) the spherical-to-formless sex organs and brown bodies.

Most of the diaphragms common to trepostome autozooecia presumably formed floors for living chambers of successive functioning bodies in the degeneration-regeneration cycle. The position of some skeletal intrazooecial structures within living chambers must have been lateral to functioning organs. Mural spines that have a definite distributional pattern might represent calcified attachment points for ligaments or muscles. Skeletal cystiphragms, hemiphragms, ring septa, and autozooecial wall thickenings all seem to be lateral features which provided significant modifications to the shape and size of the autozooidal living chamber. These and other skeletal structures appear to have been developed by zooids growing with colony-wide cyclic coordination so that skeletal structures commonly display a constant relative spacing or size correlation in the growth sequence of a colony. Hemiphragms, cystoidal diaphragms, ring septa, and skeletal cystiphragms and funnel-cystiphragms in some species are perhaps more comparable in cycle with basal diaphragms of autozooecia, suggesting that their distribution might have been controlled largely by degeneration-regeneration cycles. Closely tabulated mesopores seem to provide an expression of the most frequent colony-wide cycles in many species and can be correlated one-to-one with some mural spines and skeletal cystiphragms. Perhaps these most closely spaced structures reflect an increase in length of soft parts during a single functional stage of the degeneration-regeneration cycle.

Some monticuliporid and diaphragmed trepostomes contain a second type of cystiphragm that forms small flask-shaped chambers filled with brown deposits that suggest a concentration of organic material during the life of the colony. These chambers do not preclude retractable lophophores but almost certainly the inflexible necks restrict significantly the room for passage of membranous structures. Because of this restriction and the scattered or thinly cyclic distribution of flask-shaped chambers known from only a few species, a primary food-gathering function does not seem feasible for them. Possibly, these restricted chambers had a reproductive function, conceivably comparable to the male zooids with reduced numbers of tentacles reported in a few species of cheilostome Bryozoa. Regardless of function, if the flask-shaped chambers and their inferred organs were zooids, they represent intrazooecial polymorphism, contrasting morphologically with the alternating and consistently present living chambers that presumably contained food-gathering organs. The shape, size, and position of food-gathering organs seem more likely then to be reflected by intrazooecial structures that are repeated regularly in autozooecia, such as basal diaphragms, cystiphragms, hemisepta, ring septa, and annular thickenings of zooecial walls.

Catalog of Type Specimens of Invertebrate Fossils: Conodonta
Frederick J. Collier, compiler
256 pages
1971 (Date of Issue: 23 September 1971)
Number 9, Smithsonian Contributions to Paleobiology
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Type specimens of the conodonts in the national collection are listed alphabetically by generic and specific name. Geographic, stratigraphic, bibliographic and other pertinent information concerning each specimen is included as an initial record entry. Additional entries for each specimen list references and binomen changes subsequent to the original isolation of the specimen in the literature. Name changes are also cross-indexed in the initial record list. Two appendices list occurrence of species by stratigraphy (system, series and formation) and geography (country and state).

Tertiary and Mesozoic Echinoids of Saudi Arabia
Porter M. Kier
242 pages, 50 figures, 67 plates
1972 (Date of Issue: 14 June 1972)
Number 10, Smithsonian Contributions to Paleobiology
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The Mesozoic and Tertiary echinoids are described from Saudi Arabia. Fifty-one species, thirty-four of them new, occur in beds ranging from the Lower Jurassic to the Miocene. Two species are present in the Lower Jurassic (Toarcian) Marrat Formation, one is new: Acrosalenia marratensis. The Middle Jurassic (Bajocian-Bathonian) middle Dhruma Formation yielded eleven species, all new: Acrosalenia arabica, Acrosalenia dhrumaensis, Heterosalenia dhrumaensis, Pseudocidaris depressa, Polycyphus arabicus, Pseudosalenia magniprocta, Farquharsonia crenulata, Leioechinus namus (type species of new genus of the family Stomechinidae), Plesiechinus altus, Bothryopneustes arabica, and Bothryopneustes dhrumaensis. Thirteen species are described from the Late Jurassic (Callovian) upper Dhruma Formation, including twelve new species: Acrosalenia bowersi, Pseudocidaris romani, Pseudocidaris raratuberculata, Hypodiadema nanituberculata, Heterosalenia brocki, Heterosalenia ornata, Leioechinus amplus, Polycyphus parvituberculatus, Holectypus phelani, Pygurus (Pygurus) arabicus, Bothryopneustes kauffmani, and Bothryopneustes inflata. One species, Bothryopneustes orientalis Fourtau, occurs in the Callovian Tuwaiq Mountain Limestone. Eleven species are reported from the Late Cretaceous (Campanian or Maestrichtian) Aruma Formation, five of which are new. Rhynchopygus arumaensis, Proraster granti, Iraniaster bowersi, Iraniaster affinimorgani, and Iraniaster affinidouvillei. The sympatric pairing of two species of Iraniaster corresponds to a pairing of another two species of this genus in the Senonian of Iran. This pairing has been reported in living spatangoids. The Early Cretaceous Yamama Formation yielded the new species Pygurus (Pygurus) yamamaensis. Beds of Eocene or Oligocene age yielded three echinoid species, one of them new: Agassizia arabica. Nine species occur in the Miocene Dam Formation, three are new: Schizechinus pentagonus, Fibularia damensis, and Agassizia powersi. These Miocene echinoids are quite similar to species now living in the littoral zone.

The distribution and affinities of the echinoid species indicate faunal provinces in the Jurassic largely confined to Saudi Arabia, and in the Cretaceous confined to Saudi Arabia and Iran with some connections to North Africa but not to India. The Miocene distribution differs in being a part of a fauna occurring along the present borders of the Persian Gulf and the Arabian Sea of Saudi Arabia, Iran, West Pakistan, and northwestern India.

Homeomorphy in Recent Deep-Sea Brachiopods
G. Arthur Cooper
25 pages, 5 figures, 4 plates
1972 (Date of Issue: 10 March 1972)
Number 11, Smithsonian Contributions to Paleobiology
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A collection of brachiopods from the Baja California Abyssal Plain forms a deep-sea assemblage unusual in that it contains three genera that are unrelated but externally almost identical; i.e., they are homeomorphs. One is Neorhynchia, an impunctate rhynchonellid; the second, a punctate terebratulid with short loop, is called Abyssothyris; and the third is referred to a new genus, Notorygmia, related to Macandrevia. A discussion of homeomorphy is followed by the systematics of the genera and species involved.

The Bradleya Problem, with Descriptions of Two New Psychrospheric Ostracode Genera, Agrenocythere and Poseidonamicus (Ostracoda: Crustacea)
Richard H. Benson
138 pages, 67 figures, 14 plates, 4 tables
1972 (Date of Issue: 30 October 1972)
Number 12, Smithsonian Contributions to Paleobiology
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The “Bradleya problem” is concerned with the discovery and definition of a group of fossil and Recent reticulate ostracodes, several of which are common to Cenozoic deep-sea sediments in many parts of the world ocean floor. These species have often been misunderstood and taxonomically confused with genera characteristic of the study of shallow-water forms. The present study attempts to resolve some of these misunderstandings by designation of several important type-specimens, description of new evidence and the proposal of a new classification based on the concept of the evolution of a reticulum in response to environmental change. A method of pattern analysis is used to define elements of the reticulum subject to evolutionary change.

Over 40 reticulate species, which would have at one time been regarded as Bradleya, were examined; only 14 of these are assigned and belong to Bradleya. Two new genera, Agrenocythere and Poseidonamicus, are described for the reception of the others, and these are placed in the new subfamily, Bradleyinae, and placed with Thaerocytherinae Hazel in a new family (Thaerocytheridae Hazel). Twenty-seven of these species are described, including Bradleya arata (Brady), B. dictyon (Brady), B. normani (Brady), Agrenocythere radula (Brady), A. pliocenica (Sequenza), and A. hazelae (van den Bold). The diagnostic characteristics of the related genera Cletocythereis, Oertliella, Jugosocythereis, and Hermanites are discussed and illustrated.

It is concluded that the psychrospheric species Agrenocythere pliocenica, which has been reported from outcrops in Italy and a long core from the Tyrrhenian Sea floor, is most closely related to A. hazelae, which became geographically widespread during the Miocene. Bradleya, Jugosocythereis, Agrenocythere, and Cletocythereis, now genera in separate families, are all thought to have been derived from a common stock of Cretaceous age.

Upper Miocene Echinoids from the Yorktown Formation of Virginia and Their Environmental Significance
Porter M. Kier
41 pages, 7 figures, 10 plates
1972 (Date of Issue: 10 April 1972)
Number 13, Smithsonian Contributions to Paleobiology
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Five echinoid species are described from the upper Miocene part of the Yorktown Formation of Virginia: Echinocardium orthonotum (Conrad), Arbacia imporcera (Conrad), Psammechinus philanthropus (Conrad), Mellita aclinensis Kier, and Spatangus glenni Cooke. The assemblage probably lived in shallow, warm-temperate waters, E. orthonotum deeply buried near shore, S. glenni shallowly buried offshore, and M. aclinensis with its test just covered near shore. Arbacia improcera and P. philanthropus presumably lived together intertidally and near shore, P. philanthropus living in holes in the indurated sediments or on the sand with its test covered with debris, whereas A. improcera probably was easily visible with nothing covering its test. Specimens formerly referred to E. orthonotum from the middle Miocene Choptank Formation from Maryland are referred to E. marylandiense, new species. Echinocardium gothicum (Ravenel), from the Bear Bluff Formation of South Carolina, is considered a junior subjective synonym of E. orthonotum.

Permian Brachiopods of West Texas, I
G. Arthur Cooper and Richard E. Grant
231 pages, 39 figures, 23 plates
1972 (Date of Issue: 29 December 1972)
Number 14, Smithsonian Contributions to Paleobiology
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The first of a projected six-part monograph on the brachiopods of the reference area for the North American Permian in the Glass, Guadalupe, Diablo, Delaware, Hueco, and Chinati Mountains of West Texas and adjacent New Mexico, this introductory volume recounts the history of geological work in the area, the development of the stratigraphic framework in the Wolfcamp, Leonard, and Guadalupe Series, and the basis for age assignments. It also explains field and laboratory techniques for collecting and preparing silicified fossils by means of acid, and it presents detailed measurements and lithic descriptions of the stratigraphic units in each mountain range in terms of the current nomenclature. The paleoecologic implications of the various rock and fossil types are interpreted, and the problems concerning large scale conglomerates, bioherms, and shell heaps are considered. The faunal composition of each stratigraphic unit in each mountain range is set forth as documentation for a local zonation of the brachiopods, intra-regional correlations, and age determinations with reference to the worldwide time scale for the Permian. There are brief accounts of each locality from which fossils were obtained. The full locality listing and the literature cited for the entire monograph are included in the present volume. Plates and line drawings illustrate the techniques of collecting and preparing fossils and the nature of certain stratigraphic units and lithic types; they diagrammatically depict numerous cross sections and correlations. Detailed maps indicate the exact positions of collections of fossils in the Glass Mountains. Taxonomic descriptions will appear in subsequent volumes.

Permian Brachiopods of West Texas, II
G. Arthur Cooper and Richard E. Grant
561 pages, 1 figure, 168 plates
1974 (Date of Issue: 16 April 1974)
Number 15, Smithsonian Contributions to Paleobiology
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The second of a six-part monograph on brachiopods from the reference area for the North American Permian in mountain ranges of West Texas and adjacent New Mexico, this volume presents a historical resumé of the ordinal classification of the Brachiopoda, definitions of morphological terms, techniques of measuring specimens, and remarks on the naming of species. The major part of the work consists of descriptions and illustrations of genera and species in the inarticulate superfamilies Discinacea and Craniacea and the articulate superfamilies Eichwaldiacea, Orthotetacea, Derbyiacea, and Lyttoniacea.

New Brachiopoda from the Indian Ocean
G. Arthur Cooper
45 pages, 1 figure, 8 plates
1973 (Date of Issue: 14 February 1973)
Number 16, Smithsonian Contributions to Paleobiology
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The collections described in this paper add significantly to the brachiopod fauna. Most of the specimens were collected on the following cruises of the Woods Hole Oceanographic research vessel Anton Bruun while participating in the biological program of the International Indian Ocean Expedition: Cruise 1, east side of the Indian Ocean and Andaman Sea; Cruise 4B, off the Gulf of Cutch in the Arabian Sea; Cruises 7 and 8, in the Mozambique Channel and off the east coast of Africa; and Cruise 9, off the east coast of the Somali Republic. Of the fifteen species in the Anton Bruun collection, nine are described as new. Three of the new species represent new genera. In addition to these a fourth new genus is based on Rhynchonella valdiviae Helmcke from the southern Indian Ocean, and a rare new species of Argyrotheca is described from the Red Sea. Species described but poorly figured by W. H. Dall also are discussed and illustrated in this paper. Available pertinent geographical and ecological data are recorded and discussed. Although Mediterranean and Atlantic brachiopod elements have been known from the Indian Ocean, the Anton Bruun collections produced two genera hitherto found only in Pacific waters.

Vema's Brachiopods (Recent)
G. Arthur Cooper
51 pages, 5 figures, 9 plates
1973 (Date of Issue: 23 February 1973)
Number 17, Smithsonian Contributions to Paleobiology
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Brachiopods dredged on the worldwide exploratory cruises of R/V Vema, of the Lamont-Doherty Geological Observatory of Columbia University, are important in expanding knowledge of brachiopod distribution and taxonomy. Thirty-two species are identified, of which six are new. Twenty-one genera are represented, one of which is new. The species of five genera and the genera of two lots could not be determined. The majority of the brachiopods were taken from waters deeper than 100 fathoms. One specimen of Abyssothyris was dredged from the greatest depth (6179 meters=20,267 feet) from which a brachiopod has been taken.

Stratigraphy and Preliminary Biostratigraphy of the Flagstaff Rim Area, Natrona County, Wyoming
Robert J. Emry
43 pages, 19 figures
1973 (Date of Issue: 17 July 1973)
Number 18, Smithsonian Contributions to Paleobiology
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About 750 feet of sediments of the early Oligocene (Chadronian) White River Formation are exposed along Flagstaff Rim in south-central Natrona County, Wyoming. About 4,000 specimens of fossil vertebrates have been collected from these outcrops. The White River Formation unconformably overlies rocks ranging in age from Precambrian to medial or late Eocene. The lithology of the White River Formation is predominantly claystone and conglomerate in the lower part of the section, changing to predominantly tuffaceous siltstone and conglomeratic channel sandstones in the upper part. Four stratigraphic sections are described. A geologic map of about 40 square miles illustrates the areal limits of the White River Formation and its relationships to underlying and overlying formations. Several distinct and easily recognizable volcanic ash beds occur at intervals within the White River sequence. These serve as convenient markers for precise stratigraphic zonation of fossils and have also provided minerals for potassium-argon dating. Dates obtained range from 35.7 to 31.6 million years.

A boulder conglomerate unit, previously considered to be the basal unit of the White River Formation and/or part of the Wind River Formation is shown to be a distinct, and probably unnamed, unit, and should not be assigned to either of these formations. It unconformably overlies the Wind River Formation and is separated from the White River Formation by an erosional disconformity with several hundred feet of relief. This information allows new interpretations of the structure of the area and adds a previously unrecognized episode of deposition and erosion to the history of the area.

The most common fossil in the White River sequence is the artiodactyl genus Leptomeryx, which is represented by two morphologically distinct lineages. One lineage is provisionally divided into two and the other into three size groups that are believed to represent different species. The local stratigraphic ranges of the different groups do not overlap. In each lineage, the size increases higher in the section. None of the groups are definitely assigned to named species, pending studies to determine the validity and limits of the named species.

Preliminary analysis of other elements of the fauna shows that there is recognizable change through time within individual lineages and that the faunal composition as a whole changes through time, within the local sequence. When the entire fauna is analyzed in detail, it should be possible to establish local range zones of the fossil species and, by their use, to gain greater temporal resolution within Chadronian time than has previously been possible.

Permian Brachiopods of West Texas, III (Part I - Text)
G. Arthur Cooper and Richard E. Grant
1127 pages, 311 plates
1975 (Date of Issue: 29 December 1975)
Number 19 (Part I - Text), Smithsonian Contributions to Paleobiology
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The third of a six-part monograph on the Permian brachiopods of the Glass, Guadalupe and other mountain ranges of West Texas, this volume contains systematic descriptions of genera and species in the suborders Productidina and Chonetidina. The Productidina, which constitute about 45 percent of the brachiopod specimens in the collections from West Texas, are divided into the superfamilies Strophalosiacea, Aulostegacea, Richthofeniacea, and Productacea. The Chonetidina, less numerous, contain the single superfamily Chonetacea.

Permian Brachiopods of West Texas, III (Part II - Plates)
G. Arthur Cooper and Richard E. Grant
1127 pages, 311 plates
1975 (Date of Issue: 29 December 1975)
Number 19 (Part II - Plates), Smithsonian Contributions to Paleobiology
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The third of a six-part monograph on the Permian brachiopods of the Glass, Guadalupe and other mountain ranges of West Texas, this volume contains systematic descriptions of genera and species in the suborders Productidina and Chonetidina. The Productidina, which constitute about 45 percent of the brachiopod specimens in the collections from West Texas, are divided into the superfamilies Strophalosiacea, Aulostegacea, Richthofeniacea, and Productacea. The Chonetidina, less numerous, contain the single superfamily Chonetacea.

Ultrastructural Studies on Graptolites, 1: The Periderm and Its Derivatives in the Dendroidea and in Mastigograptus
Adam Urbanek and Kenneth M. Towe
48 pages, 2 figures, 30 plates, 2 tables
1974 (Date of Issue: 15 May 1974)
Number 20, Smithsonian Contributions to Paleobiology
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Organic skeletons of two dendroid graptolites and an aberrant sessile graptolite (Mastigograptus sp.), all Ordovician in age, were isolated chemically from the matrix and used subsequently for ultrastructural studies with the transmission electron microscope. Peridermal material of all forms investigated proved to be unusually well preserved as far as ultrastructural features are concerned, and it reveals a variety of fabrics and patterns classified into two structural categories: fabric and tissue. Following Kozlowski (1949) the term tissue is retained for larger structural components of the periderm, defined from a morphogenetic point of view. To distinguish the fusellar and cortical tissue, directly observed structures are classified as fabrics defined by the form of their unit elements and their spatial interrelations. Fusellar, cortical, sheet, and crassal fabrics were distinguished and characterized. Fusellar and cortical fabrics are both fibrillar but differ in fibril diameter. Fibrils are branched and wavy, or straight, producing a mesh, or showing a parallel arrangement and packed into layers. Sheet fabric is an electron dense, homogenous or densely reticulated material delimiting particular layers within the cortical tissue or producing an external pellicle on the fuselli. Crassal fabric is an electron dense and featureless material found in the sheath of stolons and as a secondary deposit inside the thecae in Mastigograptus sp. A given peridermal tissue is thus composed of more than one fabric, but with a predominance of either a fusellar or cortical one.

In addition to an earlier observation (Towe and Urbanek, 1972) that cortical fibrils are collagen-like, arguments are presented in favor of the opinion that fusellar fibrils may also represent the collagen group of fibrous proteins. Frequently observed passages of a single fusellar fibril within the body of a fusellus, into a fibril of an outer lamella of the same fusellus, showing a typically cortical arrangement of the fibrils, seems to indicate the same chemical nature of the fibrous components in both fabrics. Differences in their physical organization are here ascribed to the changes in the composition of the matrix.

The unexpected presence of a cortical coating on the inner thecal walls in Dictyonema sp. is explained through delayed growth of lateral thecae in the triad and very early deposition of cortical tissue over the outer surface of the young autotheca (autocortex). This wall is later overgrown by lateral thecae to become an inner thecal wall. A common cortical envelope secreted later over the outer surface of all thecae is called the rhabdocortex. The presence of a cortical component on the inner surface of the inner thecal walls in Acanthograptus sp. suggests, that at least in some dendroids also, the soft tissues inside the thecal cavity were capable of secretion of the cortical tissue. This changes the classical scheme of cortical tissue formation as proposed by Kozlowski (1949).

The fusellar tissue in Mastigograptus sp. reveals unusual, erratic distribution of fuselli, which are present only on certain places in the thecal wall. Over large areas these are substituted by a peculiar layer of electron dense, homogenous material termed here the crassal fabric. This is interpreted as a presumed secondary resorption and subsequent substitution of the fusellum by the crassal layer. The basal disc of Mastigograptus sp. is composed of a material resembling that in cortical fabric but with fewer fibrils embedded in abundant matrix. The lower layer of the basal disc is provided with numerous filaments made of delicate fibrils. The sclerotized sheaths of stolons recognized in Acanthograptus sp. are made mainly of the crassal fabric.

Permian Brachiopods of West Texas, IV (text)
G. Arthur Cooper and Richard E. Grant
685 pages, 1 figure, 160 plates
1976 (Date of Issue: 12 February 1976)
Number 21 (text), Smithsonian Contributions to Paleobiology
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The fourth of a six-part monograph on the Permian brachiopods from several mountain ranges in western Texas, especially the Glass Mountains of Brewster County, this volume contains descriptions of genera and species in the orders Rhynchonellida and Spiriferida. The Rhynchonellida contain 30 genera in the superfamily Rhynchonellacea and 3 genera in the superfamily Stenoscismatacea. The Spiriferida contain 2 genera of Cyrtiacea, 4 of Athyridacea, 9 of Spiriferacea, and 6 of Reticulariacea.

Permian Brachiopods of West Texas, IV (plates)
G. Arthur Cooper and Richard E. Grant
685 pages, 1 figure, 160 plates
1976 (Date of Issue: 12 February 1976)
Number 21 (plates), Smithsonian Contributions to Paleobiology
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The fourth of a six-part monograph on the Permian brachiopods from several mountain ranges in western Texas, especially the Glass Mountains of Brewster County, this volume contains descriptions of genera and species in the orders Rhynchonellida and Spiriferida. The Rhynchonellida contain 30 genera in the superfamily Rhynchonellacea and 3 genera in the superfamily Stenoscismatacea. The Spiriferida contain 2 genera of Cyrtiacea, 4 of Athyridacea, 9 of Spiriferacea, and 6 of Reticulariacea.

Ultrastructural Studies on Graptolites, 2: The Periderm and Its Derivatives in the Graptoloidea
Adam Urbanek and Kenneth M. Towe
48 pages, 3 figures, 24 plates, 1 table
1975 (Date of Issue: 16 May 1975)
Number 22, Smithsonian Contributions to Paleobiology
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The ultrastructure of the organic periderm in Didymograptus sp. (Lower Ordovician), Pristiograptus dubius (Suess), and in the retiolitid Holoretiolites mancki (Münch) (both Upper Silurian), isolated by chemical treatment, has been studied with the transmission electron microscope. In the first two graptoloids, with continuous peridermal walls in the rhabdosome, the periderm is composed of the major fusellar and cortical components which are essentially the same as those recognized earlier in the dendroid graptolites (Urbanek and Towe, 1974). In addition, both graptoloids examined show important differences in the fabric ultrastructure of some homologous parts of the rhabdosome such as the nema in Didymograptus sp., the virgula in P. dubius, as well as the prosicula in both. These occurrences of sharp differences in the submicroscopic structure of homologous parts of rhabdosomes within the Graptoloidea are suggestive of rather substantial phylogenetic changes in the history of the group at the ultrastructural level.

In addition to the peridermal materials previously recognized in the dendroids, certain parts of the rhabdosomes in graptoloids are constructed of a peculiar fabric termed the virgular fabric. It has been recognized so far in the virgula of Pristiograptus dubius and in the lists of the peridemal framework (clathrium) of Holoretiolites mancki, where it is the only component of the skeleton. It is composed of layers made of lucent fibrils with a unique substructure, embedded in an electron dense and homogeneous matrix, and separated by thin layers formed by this matrix alone. The biochemical relationships of these fibrils with extant fibrous materials are uncertain but a correlation with certain collagens has been suggested.

The structural relationships of the outer cortical deposit of thecae examined in Didymograptus sp. and in Pristiograptus dubius are indicative of different modes of secretion of the cortex in both. In Didymograptus sp. the cortex is formed through an accumulation of the overlapping outer lamellae of fuselli, while in Pristiograptus dubius the layers of the cortex are laid down over the surface of the fuselli as independent units. Moreover, other observations seem to indicate that in Didymograptus sp. the cortex has been formed in a somewhat different way at certain places on the rhabdosome. There does not seem to be any single, uniform pattern of secretion of the cortex in the Graptoloidea.

Paleornithology of St. Helena Island, South Atlantic Ocean
Storrs L. Olson
49 pages, 10 figures, 6 plates, 8 tables
1975 (Date of Issue: 20 June 1975)
Number 23, Smithsonian Contributions to Paleobiology
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The present avifauna of St. Helena is a very depauperate one, many species of birds having been extirpated since man's discovery of the island in 1502. The great extent of this extinction was confirmed by a study of over 4600 specimens of fossil and subfossil bird bones, representing 21 species, collected from rich deposits on the island. These deposits vary in age and fall roughly into three groups, the oldest of which extends well back into the Pleistocene, the youngest of which is very recent, and the third is intermediate. The deposits yielded the remains of the following species, six of which are here described as new: Procellariidae—Pterodroma rupinarum, new species, Bulweria bifax, new species, Puffinus pacificoides, new species, P. griseus, P. Iherminieri; Oceanitidae—Pelagodroma marina, Oceanodroma castro; Phaethontidae—Phaethon aethereus; Sulidae—Sula sula, S. dactylatra; Fregatidae—Fregata ariel trinitatis, F. minor; Rallidae—Atlantisia podarces, Porzana astrictocarpus; Charadriidae—Charadrius sanctaehelenae; Laridae—Larus sp., Gygis alba, Sterna fuscata; Columbidae—Dysmoropelia dekarchiskos, new genus and species; Cuculidae—Nannococcyx psix, new genus and species; Upupidae—Upupa antaios, new species. Pterodroma rupinarum and Puffinus pacificoides belong to species-groups that presently occur in the Indo-Pacific but not in the Atlantic; no gadfly-petrel of the size of Bulweria bifax is found in the Atlantic today. Puffinus griseus and Larus sp. are thought merely to have been vagrants on St. Helena. The pigeon and the hoopoe were both large and probably flightless. Differences in the composition and relative abundance of species between the deposits of different age indicate that the marine environment at St. Helena became progressively more tropical late in the Pleistocene. This resulted in great decreases in, or even extinction of, some of the species of Procellariiformes and in the appearance later in the fossil record of more purely tropical seabirds such as boobies, frigatebirds, and Sooty Terns.

Permian Brachiopods of West Texas, V
G. Arthur Cooper and Richard E. Grant
551 pages, 1 figure, 118 plates
1976 (Date of Issue: 15 October 1976)
Number 24, Smithsonian Contributions to Paleobiology
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The fifth of a six-part monograph on the brachiopods and Permian stratigraphy of the Glass Mountains and other ranges in western Texas and adjacent areas, this volume completes the systematic and descriptive part of the monograph with a discussion of the punctate groups. The Order Rhipidomellida contains 1 genus in the superfamily Rhipidomellacea, 4 in the Enteletacea, 1 in the Rhynchoporcea, 11 in the Spiriferinacea, and 2 in the Retziacea. The greatest number of species in this volume belong to the Order Terebratulida, contained in 19 genera.

Revised Tertiary Stratigraphy and Paleontology of the Western Beaver Divide, Fremont County, Wyoming
Robert J. Emry
20 pages, 6 figures
1975 (Date of Issue: 23 October 1975)
Number 25, Smithsonian Contributions to Paleobiology
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In the western Beaver Divide area in west-central Wyoming, a lens of coarse Tertiary volcanic conglomerate and tuff disconformably overlies Uintan rocks of the Wagon Bed Formation. The coarse volcaniclastic rocks were previously regarded as a facies of the Beaver Divide Conglomerate Member of the Chadronian White River Formation, although fossil mammals from the volcaniclastic unit are species known otherwise only from Uintan rocks. Reexamination of field relations has shown that the White River Formation disconformably overlies the volcaniclastic unit. The “lower Uinta C” temporal equivalence indicated by the fossils from the volcaniclastic unit is no longer anomalous; the underlying Wagon Bed Formation has fossil mammals indicating “Uinta B” equivalence, and the overlying White River Formation has a fairly diverse Chadronian fauna. The Uintan volcaniclastic unit is assigned to the Wiggins Formation. The Beaver Divide Conglomerate Member is restricted to conglomerate in the lower part of the White River Formation. It is composed predominantly of clasts of locally derived Precambrian crystalline rocks.

Mammalian Faunal Zones of the Bridger Middle Eocene
C. Lewis Gazin
25 pages
1976 (Date of Issue: 20 January 1976)
Number 26, Smithsonian Contributions to Paleobiology
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The zoning arrangement of the Bridger Middle Eocene as defined by W. D. Matthew in his 1909 monograph on the Carnivora and Insectivora of the Bridger Basin included a series of stratigraphic units lettered from A to E. The type section is in the western part of the basin but correlation of the sequence in the eastern part of the basin erred in that a very large area shown by Matthew as C, or upper Bridger, is actually B, or lower Bridger. As a consequence many of the mammalian remains collected in the eastern part of the basin were attributed to the wrong horizon. This was discovered in my faunal studies and verified by Wilmot Bradley's mapping of the Sage Creek White Layer, which is the base of Bridger C or upper Bridger.

A faunal list of the Mammalia recognized in the Bridger is given with type localities and their horizons, so far as known, and the number of specimens in the National Museum of Natural History collections from each of the two divisions, lower and upper. Following this a discussion of species is given in which the evidence for any species being restricted to one or the other of the stratigraphic divisions is cited, or such information demonstrating its occurrence in both levels, if this is not indicated by the National Museum of Natural History collections (under the catalog numbers of the old United States National Museum).

The Annotated Bibliography includes references to all papers in which recognized new mammalian families, genera, and species included in the Bridger faunas are described. Also included are papers in which stratigraphic and additional or detailed information on Bridger mammals is provided, with notations as to extent of coverage, and possible errors of detail or interpretation in certain cases.

Collected Papers in Avian Paleontology Honoring the 90th Birthday of Alexander Wetmore
Storrs L. Olson, editor
211 pages, 91 figures, 38 tables
1976 (Date of Issue: 21 May 1976)
Number 27, Smithsonian Contributions to Paleobiology
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Eighteen papers covering diverse aspects of avian paleontology—from the earliest known bird to extinct species found in Indian middens—are collected here to honor the 90th birthday of Alexander Wetmore. These are preceded by an appraisal of the current state of avian paleontology and of Alexander Wetmore's influence on it, including a bibliography of his publications in this field. John H. Ostrom analyzes the hypothetical steps in the origin of flight between Archaeopteryx and modern birds. Philip D. Gingerich confirms that Ichthyornis and Hesperornis did indeed bear teeth, that the palate in Hesperornis is paleognathous, and that these Cretaceous toothed birds appear to occupy a position intermediate between dinosaurs and modern birds. Larry D. Martin and James Tate, Jr. describe the skeleton of the Cretaceous diving bird Baptornis advenus and conclude that the Baptornithidae belong in the Hesperornithiformes, but are less specialized than Hesperornis. Pierce Brodkorb describes the first known Cretaceous land bird as forming a new order possibly ancestral to the Coraciiformes and Piciformes. E. N. Kurochkin summarizes the distribution and paleoecology of the Paleogene birds of Asia, with particular emphasis on the evolution of the gruiform families Eogruidae and Ergilornithidae. Pat Vickers Rich and David J. Bohaska describe the earliest known owl from Paleocene deposits in Colorado. Alan Feduccia transfers the Eocene genus Neanis from the Passeriformes to the Piciformes and he and Larry D. Martin go on to refer this and four other genera to a new family of Piciformes, concluding that these were the dominant perching land birds of the Eocene of North America. Storrs L. Olson describes a new species of Todidae from the Oligocene of Wyoming and refers the genus Protornis from the Oligocene of Switzerland to the Momotidae, concluding that the New World Coraciiformes originated in the Old World. Charles T. Collins describes two new species of the Eo-Oligocene genus Aegialornis and presents evidence that the Aegialornithidae should be referred to the Caprimulgiformes rather than to the Apodiformes, although they might be ancestral to the swifts. In the following paper he shows that the earliest known true swifts (Apodidae) are three nominal forms from the Lower Miocene of France which prove to be but a single species of Cypseloides, a modern genus belonging to a primitive subfamily now restricted to the New World. Stuart L. Warter describes a new osprey from the Miocene of California to provide the earliest certain occurrence of the family Pandionidae and he treats functional aspects of the evolution of the wing in Pandion. Hildegarde Howard describes a new species of flightless mancalline auk, also from the Miocene of California, which is temporally and morphologically intermediate between Praemancalla lagunensis and the species of Mancalla. Robert W. Storer analyzes Pleistocene fossils of pied-billed grebes, synonymizing Podilymbus magnus Shufeldt with modern P. podiceps and describing a new species from peninsular Florida. Kenneth E. Campbell, Jr., lists 53 species of birds, including new species of Buteo and Oreopholus, from a Pleistocene deposit in southwestern Ecuador and compares this with a fauna of similar age from northwestern Peru, both of which indicate more humid conditions in the past. Oscar Arredondo summarizes aspects of the morphology, evolution, and ecology of the gigantic owls, eagles, and vultures recently discovered in Pleistocene deposits in Cuba. Joel Cracraft analyzes variation in the moas of New Zealand, reduces the number of species recognized to 13, and suggests that several “species pairs” represent examples of sexual size dimorphism. G. Victor Morejohn reports remains of the extinct flightless duck Chendytes lawi, previously known only from Pleistocene deposits, from Indian middens in northern California and concludes that the species became extinct through human agency less than 3800 years ago.

Phoca wymani and Other Tertiary Seals (Mammalia: Phocidae) Described from the Eastern Seaboard of North America
Clayton E. Ray
36 pages, 3 figures, 11 plates
1976 (Date of Issue: 14 May 1976)
Number 28, Smithsonian Contributions to Paleobiology
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Fossil seal remains from Richmond, Virginia, first reported by Wyman in 1850, and named Phoca wymani by Leidy in 1853, have been neglected and unjustifiably regarded as cetacean by most subsequent authors. Recently recognized parts of the holotype and other material, in part recently collected in Richmond, show that the species is a monachine seal, here called Monotherium? wymani (Leidy, 1853a). It is derived from Miocene beds that are definitely older than the Yorktown Formation and probably correlative with the Calvert Formation of Maryland. Thus Monotherium? wymani is probably the oldest known monachine. Other evidence of fossil phocids in eastern North America is reviewed.

A Vector Approach to Size and Shape Comparisons among Zooids in Cheilostome Bryozoans
Alan H. Cheetham and Douglas M. Lorenz
55 pages, 37 figures, 19 tables
1976 (Date of Issue: 8 July 1976)
Number 29, Smithsonian Contributions to Paleobiology
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Although zooid size and shape have long been used in comparative studies of cheilostome bryozoans, procedures for measuring these properties have been little investigated. Predominatly intussusceptive growth of buds suggests a method of comparing zooid outlines based on (1) correspondence of principal growth direction (proximal-distal axis) and (2) size and shape properties expressing differential growth about this axis.

Vector properties of a wide variety of autozooidal outlines (in frontal view) were studied by principal components. Size (area within the outline) accounts for more than one-third of the variation and tends to vary less within colonies than shape, even in severely disturbed budding patterns. The portion of shape independent of size is divisible into three components. Each of the first two components accounts for about one-fourth of the total variation, the third for less than five percent. One shape component is associated with asymmetry of outline, as measured both by departure of the mean vector direction from the proximal-distal axis and by inequality of vector lengths on either side of the axis. The amount of asymmetry is small, can be either antisymmetry or fluctuating asymmetry, and varies greatly within colonies apparently with microenvironmental effects on budding patterns. The second shape component is associated with elongation (concentration of vector lengths near the mean growth direction) and distal inflation (proportion of area distal to the midpoint of the proximaldistal axis). These two variables seem less affected by microenvironment than is asymmetry. The third component accounts for only the small part of variation in elongation and distal inflation that is not positively correlated. Variation in this component suggests that distal inflation is slightly more sensitive to microenvironment than is elongation. Estimates of intrapopulation variation in one fossil species suggest that size and that part of elongation varying in opposition to distal inflation are sufficiently consistent within single populations, under the same conditions of ontogeny, astogeny, and polymorphism, to form a basis for taxonomic discrimination. Within the range of colony means for each of these two properties among the variety of outlines examined, at least three and possibly four potentially taxonomically distinct intervals can be recognized. The number of measurements per colony needed to detect differences between these intervals is surprisingly small.

Triassic Echinoids
Porter M. Kier
88 pages, 16 figures, 21 plates, 2 tables
1977 (Date of Issue: 28 January 1977)
Number 30, Smithsonian Contributions to Paleobiology
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Although 142 species of Triassic echinoids have been reported, only 24 are based on sufficient material to permit reliable generic identification. These species are redescribed and illustrated. Twelve of them are from the St. Cassian Beds in the Italian Dolomites. A large collection of specimens from these beds is described, from which are erected two new species, Megaporocidaris mariana and Levicidaris zardinia, and five new genera: Zardinechinus, a miocidarid, Paurocidaris and Leurocidaris, cidarids, and the psychocidarids, Levicidaris and Megaporocidaris. A new pedinoid, Hemipedina hudsoni, is described from the Norian of Arabia. During the Early and Middle Triassic, only flexible miocidarids were present. Apparently all Mesozoic echinoids are descended from them. The first cidarids occur in the Late Triassic (Karnian) together with the first psychocidarids. The first certain pedinoid appears in the Norian. Finally at the close of the Triassic, in the Rhaetian, the first hemicidaroid appears.

Ecology and Systematics of Foraminifera in Two Thalassia Habitats, Jamaica, West Indies
Martin A. Buzas, Roberta K. Smith and Kenneth A. Beem
139 pages, 38 figures, 8 plates, 34 tables
1977 (Date of Issue: 11 July 1977)
Number 31, Smithsonian Contributions to Paleobiology
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Homogeneous Thalassia beds in back-reef flat (less than 1 m) and Discovery Bay (about 3 m) were sampled for 12 successive months in Jamaica, West Indies. Living foraminifera were enumerated in each of four monthly replicates consisting of 20 ml of sediment. At the sampling times, water temperature, sediment temperature, salinity, oxygen saturation, water pH, sediment pH, sediment median, sediment sorting, turbidity, particulate organic carbon, Thalassia weight, and weight percent silt plus clay were measured.

In all, 18,644 individuals belonging to 143 species were picked, sorted, and identified. The back-reef flat habitat contained 7,745 individuals belonging to 115 species, while the Discovery Bay contained 10,899 individuals belonging to 117 species. Fisher's log-series fits the distribution of species abundances at both habitats well. The number of species, information function, and equitability are usually greater at Discovery Bay for individual 20 ml samples.

A general linear model consisting of parameters for station differences, overall periodicity, interaction of station differences and overall periodicity, and environmental variables was constructed. The densities of the 19 most abundant species were statistically analyzed individually (univariate) and simultaneously (multivariate).

Univariate analyses indicate six species have significant station differences (95% level) and seven exhibit periodicity. The environmental variables are not significant for any of the species. Multivariate analyses indicate a significant difference between stations and an overall periodicity. As in the univariate analyses, environmental variables are not significant. The results suggest that in tropical habitats changes in species densities are regulated biotically.

The new species Ammonia jacksoni, Elphidium norvangi, Fissurina goreaui, Discorbinella minuta, Glabratella altispira and G. compressa are described. Taxonomic remarks are presented for most of the species.

Permian Brachiopods of West Texas, VI
G. Arthur Cooper and Richard E. Grant
210 pages, 3 tables
1977 (Date of Issue: 6 October 1977)
Number 32, Smithsonian Contributions to Paleobiology
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The sixth and final part of a monograph of Permian strata and faunas of West Texas and adjacent parts of New Mexico, this volume consists primarily of faunal lists and a taxonomic index of the previous five parts. A list of brachiopods grouped according to R. E. King's localities and compiled from King's data by the authors is followed by the author's compilation of lists of brachiopods found at the localities designated by the United States Geological Survey, the American Museum of Natural History, Kansas University, and the National Museum of Natural History. The list pertaining to the last three groups of localities carries a code indicating the approximate number of specimens of each taxon in the museum collections. A list of the authors' collection of ammonites is included, followed by lists of their fusulinids; these are appended to this monograph on brachiopods because they have important implications for dating and correlation. Also included are Corrigenda (with reference to the previously published parts) and suggestions for future related research.

Evolution of Oblitacythereis from Paleocosta (Ostracoda: Trachyleberididae) during the Cenozoic in the Mediterranean and Atlantic
Richard H. Benson
47 pages, 11 figures, 4 plates
1977 (Date of Issue: 29 August 1977)
Number 33, Smithsonian Contributions to Paleobiology
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A new ostracode genus Oblitacythereis containing two new subgenera (the nominotypical form and the older Paleoblitacythereis) containing two new species (O. (O.) mediterranea and O. (P.) luandaensis) and one old species (new designation, O. (P.) ruggierii (Russo)) have been demonstrated to have descended from a common ancestral stock (new genus Paleocosta) of the genus Costa, the nominate form of the tribe Costini.

This genus contains heavily costate species whose history has been one of invasion of the greater depths of Tethys, which became thermospheric in the middle Miocene. Species of subgenus Paleoblitacythereis became adapted to upper slope and warm basinal habitats and underwent considerable modification of its carapace structure. When Tethys became extinct as a marine environment at the end of the Miocene, subgenus Paleoblitacythereis was eradicated in the Mediterranean region but survived in the Atlantic, where it lives today. Its descendant subgenus Oblitacythereis invaded the newly formed Mediterranean in the Early Pliocene, structurally modified to live in cooler water.

The history of Oblitacythereis was traceable because of a detailed analysis of structural and form homology, substantiated by quantitative Theta-Rho test.

Index of Living and Fossil Echinoids 1924-1970
Porter M. Kier and Mary Hurd Lawson
182 pages
1978 (Date of Issue: 10 February 1978)
Number 34, Smithsonian Contributions to Paleobiology
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All new taxa of fossil and living echinoids described from 1924 to 1970 are listed with their age, geographic and stratigraphic occurrence, and bibliographic citation.

A Lower Eocene Frigatebird from the Green River Formation of Wyoming (Pelecaniformes: Fregatidae)
Storrs L. Olson
33 pages, 31 figures
1977 (Date of Issue: 7 October 1977)
Number 35, Smithsonian Contributions to Paleobiology
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A new subfamily, Limnofregatinae, is erected for the first known Tertiary specimens of the pelecaniform family Fregatidae. These are described as a new genus and species, Limnofregata azygosternon. The holotype is a nearly complete skeleton with feather impressions. Two additional specimens, consisting of a partial skeleton and a fragment of ulna, are referred to the same species. All are from freshwater lake deposits in the Lower Eocene Green River Formation of Wyoming, which are roughly 50 million years old. The principal differences between Limnofregata and modern frigatebirds (Fregata) are seen in its shorter and less hooked rostrum, proportionately shorter wings and longer hindlimb, in the lack of fusion in the pectoral girdle, and in the lack of extensive pneumatization of the skeleton. The fossilform is, nevertheless, well advanced along the lines of modern frigatebirds and nothing in its morphology seems to preclude its being ancestral to Fregata. The fossil provides additional support for placing the suborder Fregatae between the more primitive tropicbirds (suborder Phaethontes) and the members of the suborder Pelecani. Frigatebirds in the early Tertiary evidently occupied different habitats or a wider range of habitats than the modern forms and have only subsequently been restricted to a purely oceanic environment.

Taxonomy and Paleoecology of Early Miocene Benthic Foraminifera of Northern New Zealand and the North Tasman Sea
Bruce W. Hayward and Martin A. Buzas
154 pages, 26 figures, 28 plates, 4 tables
1979 (Date of Issue: 3 August 1979)
Number 36, Smithsonian Contributions to Paleobiology
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Data from 51 samples of early Miocene benthic foraminifera (200-300 individuals per sample) from west Northland, New Zealand (Waitakere and Waitemata Groups), together with those from four samples from the north Tasman sea (Deep Sea Drilling Project 206), are analysed by multidimensional scaling and cluster analysis. The samples are grouped in terms of species abundances into six thanatotopes, which are interpreted as follows: A, dominated by robust Amphistegina madagascariensis, 10%-37% planktonics, inner neritic; B, Cibicides-Cibicidoides dominant, 10%-55% planktonics, outer neritic; C, Gyroidina, Euuvigerina, Astrononion, Lenticulina most abundant, 32%-87% planktonics, upper bathyal; D, Cassidulina-Bolivina-Cibicides dominant, 16%-99.5% planktonics, upper and midbathyal; E, Globocassidulina-Epistominella dominant, 99.5% planktonics, lower bathyal; F, Quinqueloculina dominant, 11%-32% planktonics, inner and midneritic.

Using these thanatotope interpretations in conjunction with their stratigraphic and geographic distributons, a model of the early Miocene paleogeography of west Northland is deduced, refining traditional models for the area. A central mid-bathyal basin (Waitemata Basin), bounded in the southwest by a pile of volcanic sediments (Waitakere volcanic pile), built up to an island surrounded by neritic and upper bathyal slopes. In the northwest (Kaipara area) volcanics erupted through a neritic shelf. This shelf became shallower and partly terrestrial during latter parts of the early Miocene. Upper bathyal slopes existed around the southern edge of the northwestern shelf. Submarine canyons cut through these slopes channeling shelf sediment into the bathyal basin. Several ungrouped, greatly mixed samples (interbedded with basin sediments) contain individuals from neritic and bathyal thanatotopes and are interpreted as having been mixed during transportation down through the canyons into the basin in the form of subaqueous sediment gravity flows.

No change in depth from the present lower bathyal is inferred to have occurred in the vicinity of DSDP 206 (north Tasman Sea) since the early Miocene.

All 378 identified species are listed together with their synonomies; many are described and 194 species are figured. Besides a number of first records for New Zealand, three new species—Elphidium gibsoni, Elphidium kanoum, and Eoeponidella scotti—are described.

Tertiary and Cretaceous Brachiopods from Cuba and the Caribbean
G. Arthur Cooper
45 pages, 2 figures, 7 plates, 1 table
1979 (Date of Issue: 30 January 1979)
Number 37, Smithsonian Contributions to Paleobiology
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Thirty-nine taxa of fossil brachiopods are described, figured, and discussed. Three come from Cretaceous rocks of Cuba and the remainder were found in Tertiary sediments of Cuba and other parts of the Caribbean region. These range in age from Eocene to Pliocene. Fourteen genera are identified of which two are new: one from the Cretaceous and the other from the Eocene, both from Cuba.

Thirty species are recognized among the fossil genera: Cruralina, 1 (new); Terebratulina, 1 (new); Tichosina, 2 new; Tichosina?, 3 (1 new); Stenosarina, 1 (new); Gryphus, 4 (2 new); Gryphus?, 1; Dyscritothyris, 1 (new genus and species); Argyrotheca, 12 (11 new); Cistellarcula, 1 (new); Hercothyris, 2 (new genus and 2 new species); Lacazella, 1. Representatives of the following genera are not identified specifically: Cryptopora, Rugia, Terebratulina, Platidia, Argyrotheca, Thecidellina.

Tertiary and Quaternary Brachiopods from the Southwest Pacific
G. Arthur Cooper
23 pages, 4 figures, 2 plates
1978 (Date of Issue: 15 December 1978)
Number 38, Smithsonian Contributions to Paleobiology
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Brachiopods from Tertiary and Quaternary sediments in the islands of the South Pacific are rarities. They are important in helping us to understand the geographic and geologic distribution and evolution of parts of the phylum. This paper describes genera and species from Fiji, Java, and the New Hebrides. All except two of the genera live in the South Pacific today but are rare, absent, or not yet taken from the waters surrounding Fiji and the New Hebrides: Craniscus?, Cryptopora, Basiliola, Terebratulina, Abyssothyris, Dallithyris?, Argyrotheca, Platidia, Frenulina, Dallina, and Thecidellina. The exceptions are: an extinct new genus, here named Dicrosia, and Lacazella, a genus common in the Mediterranean, less common in the Caribbean, but not now known to be living in the Pacific.

The Echinoids of the Middle Eocene Warley Hill Formation, Santee Limestone, and Castle Hayne Limestone of North and South Carolina
Porter M. Kier
102 pages, 26 figures, 22 plates, 4 tables
1980 (Date of Issue: 6 November 1980)
Number 39, Smithsonian Contributions to Paleobiology
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The echinoids are described from the middle Eocene Warley Hill Formation, Santee Limestone, and Castle Hayne Limestone of North and South Carolina. Twenty-seven species are present including the following new taxa: Eurhodia baumi, Eurhodia rugosa ideali, Eurhodia rugosa depressa, Eupatagus wilsoni, Eupatagus lawsonae, Linthia harmatuki, Agassizia wilmingtonica Cooke inflata, and Protoscutella mississippiensis (Twitchell) rosehillensis. Three zones are identified: the earliest characterized by Protoscutella mississippiensis (Twitchell) and Santeelampas oviformis (Conrad), a “middle zone” with Linthia harmatuki and the youngest species of Protoscutella, and a “late zone” with large numbers of Periarchus lyelli (Conrad) and Echinolampas appendiculata Emmons. The “early zone” is considered early middle Eocene, the “middle zone” middle Eocene and the “late zone” probably late middle Eocene. The three species of Protoscutella appear to represent an evolutionary series—P. mississippiensis (Twitchell) to P. conradi (Cotteau) to P. plana (Conrad)—characterized by the shifting of the periproct nearer to the peristome.

The echinoids lived in well-aerated sediments in a tropical sea.

Glyptodonts of North America
David D. Gillette and Clayton E. Ray
255 pages, 97 figures, 70 tables
1981 (Date of Issue: 21 December 1981)
Number 40, Smithsonian Contributions to Paleobiology
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All known North American glyptodonts belong in the genus Glyptotherium Osborn, 1903 (Family Glyptodontidae, Subfamily Glyptodontinae). Junior synonyms are Brachyostracon Brown, 1912; Boreostracon Simpson, 1929; Xenoglyptodon Meade, 1953; and all assignments of North American specimens to Glyptodon Owen, 1838. The ancestral species is Glyptotherium texanum from the Early Pleistocene Tusker (Arizona) and Blanco (Texas) local faunas of the Blancan Land Mammal Age; G. texanum is smaller and lacks many of the exaggerated features of the descendant species. The descendant species are G. arizonae (Blancan? and Irvingtonian); G. floridanum (Rancholabrean); and two species known from isolated localities in Mexico, G. cylindricum and G. mexicanum. The taxonomic validity of G. mexicanum is questionable.

The geographic distribution and faunal associations of Glyptotherium clearly indicate tropical or subtropical habitats. North American glyptodonts exhibit extreme tendencies toward hypsodonty and homodonty in the dentition, and they lack both incisiform and caniniform teeth. They probably fed on soft vegetation near permanent bodies of water. Graviportal limb proportions and details of the gross osteology suggest slow and cumbersome locomotion, which probably precluded occupation of upland habitats.

A substantial expansion in the number of specimens available for study has extensively improved our knowledge of the gross osteology of Glyptotherium, especially for G. texanum and G. arizonae.

New Brachiopods from the Southern Hemisphere and Cryptopora from Oregon (Recent)
G. Arthur Cooper
43 pages, 4 figures, 7 plates
1982 (Date of Issue: 29 July 1982)
Number 41, Smithsonian Contributions to Paleobiology
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The Recent brachiopods described herein were collected during dredging operations of several research vessels: the United States R/V Eltanin (alias Islas Orcadas while on loan to Argentina) in Antarctica, in the subantarctic waters of the Atlantic around South Georgia and the South Sandwich Islands, and in the Scotia Sea; R/V Hero dredging in Antarctic waters; R/V Anton Bruun operating along the west coast of South America; the initial cruises of R/V Rafale and Thierry operating in the Gulf of Guinea off the coast of West Africa.

Seventeen genera are described, including 19 species and 10 lots not identified as to species; Pelagodiscus atlanticus (King), Discinisca laevis (Sowerby), Discinisca species, Cryptopora hesperis, new species, the first report of this genus in the northern Pacific, Terebratulina kiiensis Dall and Pilsbry, Terebratulina species, Abyssothyris wyvillei (Davidson), A. cf. elongata Cooper, Abyssothyris? species, Liothyrella delsolari, new species, L. expansa, new species, L. fosteri, new species, L. georgiana Foster, L. hendleri, new species, L. notorcadensis (Jackson), L.? vema Cooper, Platidia species, Argyrotheca species, Megathiris species, Pantellaria monstruosa (Scacchi), Macandrevia americana Dall, Macandrevia species, Notorygmia species, Terebratella? species, Syntomaria curiosa, new genus and species, Dyscritosia secreta, new genus and species, Neothyris parva, new species. The loop development of the new genera is described and illustrated in detail.

Most of the localities from which the specimens were taken add geographic and bathymetric information new for these genera.

Living and Fossil Brachiopod Genera 1775-1979: Lists and Bibliography
Rex A. Doescher
238 pages
1981 (Date of Issue: 19 November 1981)
Number 42, Smithsonian Contributions to Paleobiology
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All genera and subgenera of fossil and living brachiopods described from 1775 to 1979 are listed with comments on their current nomenclatural status. A computerized list presents all valid brachiopod genera (and subgenera), including subjective synonyms, with additional information regarding type-species, superfamily classification, and range (by period) of each genus. A bibliography supplements both lists.

Brachiopoda from the Southern Indian Ocean (Recent)
G. Arthur Cooper
93 pages, 30 figures, 14 plates, 1 table
1981 (Date of Issue: 21 December 1981)
Number 43, Smithsonian Contributions to Paleobiology
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Specimens collected from 120 stations around and between the subantarctic islands: Marion, Prince Edward, Crozet, Kerguelen, Heard, Amsterdam, and St. Paul by the M/S Marion Dufresne with the support of Terres Australes et Antarctiques Francaises, Paris, greatly increase our knowledge of the brachiopoda of the Indian Ocean. Eighteen species are recognized that include 16 genera, 2 of them new: Pemphixina and Xenobrochus; 11 new species: Basiliola arnaudi; Eucalathis magna, E. costellata, E. rotundata, Xenobrochus australis, X. anomalus; Dallithyris? dubia; Dyscolia? radiata, Platidia marionensis; Ecnomiosa inexpectata, and Thecidellina minuta. Seven hitherto described species are recorded: Pelagodiscus atlanticus (King); Pemphixina pyxidata (Davidson); Liothyrella moseleyi (Davidson); Megerlina davidsoni (Vélain); Megerlia gigantea (Deshayes); Aerothyris kerguelenensis (Davidson), and A. aff. A. macquariensis (Thomson). Six genera, which could not be identified specifically, were also taken: Crania, Basiliola, Tegulorhynchia, Eucalathis, Liothyrella, and Aerothyris. Genera recognized in the Indian Ocean for the first time are: Tegulorhynchía, Basiliola, and Ecnomiosa, the last hitherto known only from the Gulf of Mexico and Caribbean Sea.

Brachiopoda from the Gulf of Gascogne, France (Recent)
G. Arthur Cooper
35 pages, 5 figures, 3 plates
1981 (Date of Issue: 17 December 1981)
Number 44, Smithsonian Contributions to Paleobiology
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Brachiopods collected during the operations of Biogas and Polygas, Gulf of Gascogne Abyssal Survey, and Thalassa, Gulf of Gascogne Bathyal Survey of the Centre Océanologique de Bretagne, are recorded and data concerning them noted. Most of the specimens taken in Biogas and Polygas are from depths ranging from 1010 meters to 4459 meters. The specimens taken by Thalassa are mostly from waters less than 1000 meters in depth. Twenty-one species in 14 genera are recognized, six of them, including a new species, hitherto not recorded from the Gulf of Gascogne, are starred: *Pelagodiscus atlanticus (King), Crania anomala (Müller), C. anomala turbinata (Poli), *Cryptopora gnomon Jeffreys, Hispanirhynchia cornea (Fischer), Eucalathis ergastica Fischer and Oehlert, E. tuberata (Jeffreys), Terebratulina retusa (Linné), T. retusa emarginata (Risso), Gryphus vitreus (Born), *G.? cooperi d'Hondt, Dallithyris? aff. D.? sphenoidea (Jeffreys), Platidia anomioides (Scacchi and Philippi), Megerlia truncata (Linné), M. echinata (Fischer and Oehlert), Pantellaria monstruosa (Scacchi), Macandrevia cranium (Müller), *M. novangliae Dall, *Fallax dalliniformis Atkins, Dallina septigera (Lovén), and *D. parva, new species. Five described species not previously recorded from the Gulf of Gascogne are: Pelagodiscus atlanticus (King), a world-wide abyssal form; Cryptopora gnomon (Jeffreys), widespread in the northern Atlantic; Macandrevia novangliae Dall, hitherto only known from deep water off the coast of New England, United States; Fallax dalliniformis Atkins, newly discovered in the approaches to the English Channel; and Gryphus? cooperi d'Hondt recently described. The first four were taken in deep water in the Biogas operations; the fifth was taken by Thalassa. Specimens taken by Thalassa are mostly small or immature forms that are usually difficult to identify generically. Specimens of the rare genus Eucalathis appeared in several lots taken by Thalassa as well as the new species, Dallina parva. Not included in the above lists is a fossil brachiopod, dredged by Thalassa, that is assigned with a query to the Cretaceous genus Meonia.

Textural Observations on Some Living Species of Planktonic Foraminifera
Richard Cifelli
45 pages, 15 plates
1982 (Date of Issue: 18 March 1982)
Number 45, Smithsonian Contributions to Paleobiology
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Wall textures of 11 species of spinose and quasi-spinose planktonic foraminifera collected in plankton tows from the North and Equatorial Atlantic have been studied with the scanning electron microscope. Illustrations and descriptions of these textures are given herein. During the ontogeny of the individual the chamber wall thickens and the texture is modified in various forms. Thickening occurs by the addition of sheet-like laminations and clustering of crystallites that generally develop into ridges among the pores. Patterns of textural development vary among species, but there are some common trends.

A Review of the Extinct Wolverine, Plesiogulo (Carnivora: Mustelidae), from North America
Jessica A. Harrison
27 pages, 16 figures, 7 tables
1981 (Date of Issue: 21 December 1981)
Number 46, Smithsonian Contributions to Paleobiology
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There are two fossil species of Plesiogulo in North America: Plesiogulo marshalli, to which most of the fossil material is herein referred, and P. lindsayi, new species. Both species are restricted to the late Hemphillian. A formal diagnosis is offered for P. marshalli. Remains of Plesiogulo are relatively rare, possibly due to low densities in extinct populations. The only known juvenile specimens of Plesiogulo are from the Edson Local Fauna, where three juveniles together with a single mature individual probably represent a female with a litter of cubs. Plesiogulo migrated to the New World some time between 7.0 and 6.5 million years ago. This taxon, generally interpreted as an inhabitant of forest or woodland, was probably equally well adapted to the open plains.

Descriptive and Comparative Osteology of the Oldest Fossil Squirrel, Protosciurus (Rodentia: Sciuridae)
Robert J. Emry and Richard W. Thorington, Jr.
35 pages, 16 figures, 3 tables
1982 (Date of Issue: 12 July 1982)
Number 47, Smithsonian Contributions to Paleobiology
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The early history of the Sciuridae is not well known, squirrels being generally poorly represented in the Tertiary fossil record. A nearly complete skeleton, recently discovered in early Oligocene deposits of Wyoming, represents what may be the oldest fossil squirrel known. For the first time, this early squirrel can be compared fully with its extant relatives. The specimen, assigned to Protosciurus jeffersoni, retains the primitive protrogomorphous zygomasseteric structure, as in other known Protosciurus, but the masseteric fossa of the mandible is farther forward than in most nonsciurid protrogomorphs. The auditory region of the skull has derived squirrel characters, but it is in the postcranial skeleton where similarities to extant squirrels are most apparent. Except for minor differences in joint construction, the skeleton is strikingly similar to that of Sciurus niger, the living fox squirrel. It differs from extant ground squirrels in the more gracile proportions of its long bones and asymmetry of foot construction. This early member of the squirrel family was clearly an arboreal squirrel, with morphology, and presumably habits, very similar to those of extant Sciurinae.

Fossil Vertebrates from the Bahamas
Storrs L. Olson, editor
65 pages, 12 figures, 12 tables
1982 (Date of Issue: 5 August 1982)
Number 48, Smithsonian Contributions to Paleobiology
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The three papers in this volume summarize the previous literature on fossil vertebrates from the Bahamas, provide revisions of the previously described fossil specimens, include identifications of newly collected material, and discuss changes in the late Pleistocene environment of the Bahaman archipelago. Olson and Pregill review the history of fossil exploration in the Bahamas, describe the known fossil localities, and briefly discuss the depauperate mammalian fauna. Pregill reviews the Pleistocene herpetofauna of New Providence Island, which is similar to that found on the island today, the only extinct taxa being a tortoise (Geochelone), a crocodile (Crocodylus), an iguana (Cyclura), and a gecko of the genus Aristelliger (previously misidentified as Tarentola). Taphonomy of the New Providence deposits and the zoogeographical patterns of the herpetofauna are discussed in relation to arid climatic conditions of the Wisconsinan glacial period. It is suggested that the establishment of a north-south rainfall gradient within the Bahamas has caused more extinctions in the wetter northern islands, whereas a more diverse herpetofauna persists in the drier southern islands. Olson and Hilgartner review the fossil record of birds from the Bahamas and propose the following changes in nomenclature: Calohierax quadratus = Buteo sp., Burhinus nanus = Burhinus bistriatus nanus, Glaucidium dickinsoni = Athene cunicularia, Otus providentiae = Athene cunicularia, Bathoceleus hyphalus = Melanerpes superciliaris, Corvus wetmorei = Corvus nasicus. About 50% of the fossil avifauna of New Providence no longer occurs there and 40% is extinct in the Bahamas. Species composition indicates that the Bahamas in the late Pleistocene were drier and had more open savanna-like and broadleaf scrub habitats. Subsequent increases in rainfall caused habitat changes that resulted in extinction. The implications of this for modern ecological theories are discussed.

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