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A revision of the Early-Middle Miocene anguine, Pseudopus laurillardi (Lartet, 1851), is presented based on a detailed anatomical analysis of one newly discovered articulated specimen and numerous disarticulated cranial and postcranial elements from several localities in Germany, as well as on the restudy of the original skeletal material of Lartet. The articulated specimen represents the first record of an articulated anguine from the Neogene. We demonstrate that the contemporaneous anguine Propseudopus fraasii ( = Pseudopus fraasi, Pseudopus moguntinus or Ophisaurus moguntinus) from Germany and elsewhere in Europe represents a junior synonym of P. laurillardi. Three species of Pseudopus can by discriminated in the Cenozoic of Eurasia: P. laurillardi (Early-Middle Miocene of Europe), P. pannonicus (Late Miocene-Middle Pleistocene of central and eastern Europe) and P. apodus (Late Pleistocene-Recent, from Eastern Europe to central Asia). Eleven morphological characters of the skull have been identified that distinguish Pseudopus laurillardi from P. pannonicus and P. apodus. Four of these characters regard the frontal and parietal bones, whereas all other characters regard the dentary and dentition. The genus Pseudopus represents the largest and most robust taxon in the subfamily Anguinae and first occurs in central Europe at the beginning of MN 4 (∼18.5 Ma). In contrast to the extant species, P. apodus, the fossils P. laurillardi and P. pannonicus had a greater ecological plasticity and lived in various types of environments. The fossil remains of these taxa are most frequently found in localities characterized by sub-humid to humid climate, which may indicate that their preferential habitats include forested environments.
Cyriacotheriidae are a family of unusual small-bodied pantodonts known from the Paleocene of the Western Interior of North America. Cyriacotheriids possess a suite of dental characters similar to that of pantodonts (e.g., molar dilambdodonty, lingual molar hypoconulids), as well as several divergent features (e.g., molarized premolars, strong molar conules) that have been interpreted as “dermopteran-like.” The unusual combination of pantodont and dermopteran-like characters, combined with a limited fossil record, has made attempts at understanding the broader relationships of Cyriacotheriidae difficult. This paper reports on a new genus and two new species of cyriacotheriids from the Paleocene of Alberta, Canada, with both species significantly older than those of the only previously described cyriacotheriid, Cyriacotherium. Collectively, the dentitions of these new taxa exhibit derived characters seen in Cyriacotherium (e.g., robust molar conules, strong molar dilambdodonty) in addition to a number of plesiomorphies seen in more basal pantodonts (e.g., conspicuous molar entoconids, deep premolar ectoflexus) and, importantly, posterior premolars that are weakly molariform and non-dilambdodont. A phylogenetic analysis of the new cyriacotheriid, basal pantodonts, dermopterans, and dermopteran-like eutherians resulted in Cyriacotheriidae nesting within a monophyletic Pantodonta. The results strengthen previous hypotheses regarding the pantodont affinities of the family, and suggest that the dermopteran-like features seen in the more derived Cyriacotherium were acquired convergently. Although the discovery of new cyriacotheriids sheds light on the evolutionary history of the family, it cannot resolve the ongoing questions of pantodont origins; nonetheless, their discovery in strata of early Paleocene age indicates that significant parts of the evolutionary history of Cyriacotheriidae, and North American pantodonts more generally, have yet to be discovered.
Light microscopy studies on new materials and museum collections of early Cambrian organic-walled microfossils, informally called acritarchs, provide the observations on phenetic features that permit a comparison to certain Modern microalgae and the recognition of various developmental stages in their life cycle. The microfossils derive from various depositional settings in Estonia, Australia, Greenland, Sweden, and Poland. The exceptionally preserved microfossils reveal the internal body within the vesicle, the endocyst, and the process of releasing the endocyst from the cyst. Vegetative cells, cysts, and endocysts are distinguished, and the hypothetical reconstruction of a complex life cycle with the alternation of sexual and asexual generations is proposed. Acritarchs from the Skiagia-plexus are cysts, and likely zygotes in the sexual generation, which periodically rested as “benthic plankton.” Some microfossils of the Leiosphaeridia-plexus that are inferred to be vegetative cells were planktonic and probably haplobiontic. These form-taxa may belong to a single biological species, or a few closely related species, and represent the developmental stages and alternating generations in a complex life cycle that is expressed by polymorphic, sphaero- and acanthomorphic acritarchs. The morphological resemblance and diagnostic cell wall ultrastructure with the trilaminar sheath structure known from earlier studies suggest that the early Cambrian microfossils are the ancestral representatives and/or early lineages to the Modern class Chlorophyceae and the orders Volvocales and Chlorococcales.
The seven species of extant sea turtles show a diversity of diets and feeding specializations. Some of these species represent distinctive ecomorphs that can be recognized by osteological characters and therefore can be identified in fossil taxa. Specifically, modifications to the feeding apparatus for shearing or crushing (durophagy) are easily recognizable in the cranium and jaw. New sea turtle fossils from the Miocene of Peru, described as a new genus and species (Pacifichelys urbinai n. gen. and n. sp.), correspond to the durophagous ecomorph. This new taxon is closely related to a recently described sea turtle from the middle Miocene of California, USA (Pacifichelys hutchisoni n. comb.), providing additional information on the osteological characters of this lineage. A phylogenetic analysis of Pacifichelys and other pan-chelonioid sea turtle lineages shows that at least seven lineages independently evolved feeding specialized for shearing or crushing. The iterative evolution of these morphologies is plausibly linked to ecological factors such as the development of seagrass communities and the opening of niches through extinction that occurred from the Cretaceous to the Miocene.
The Lower Ferron Sandstone Member of the Mancos Shale in southeastern Utah preserves a chondrichthyan assemblage of at least 13 taxa that include: Hybodus sp., Ptychodus cf. P. mammillarisAgassiz, 1843, Ptychodus whippleiMarcou, 1858, cf. Chiloscyllium sp., Scapanorhynchus raphiodon (Agassiz, 1843), Cretodus crassidens (Dixon, 1850), cf. Leptostyrax sp., cf. Cretalamna appendiculata (Agassiz, 1835), Squalicorax sp., Pseudohypolophus mcnultyi (Thurmond, 1971), Protoplatyrhina hopiiWilliamson, Kirkland and Lucas, 1993, Ischyrhiza schneideri (Slaughter and Steiner, 1968), and Ptychotrygon triangularis (Reuss, 1844). Although this assemblage is typical of other Turonian chondrichthyan faunas in North America, fossil teeth are preserved in two unique facies associations that consist of arenitic sandstones with mud interclasts and rounded chert, feldspar, and quartz pebbles. The coarser beds within these facies associations are previously interpreted to represent storm events and turbidity flows associated with a sea level lowstand. Chondrichthyan teeth occurring within these coarser beds are indicative of extensive transport and reworking and attest to the durable nature of chondrichthyan teeth for biostratigraphic and paleoecological interpretations. Similar studies of chondrichthyan teeth in shelf marine settings may also provide new insights for facies interpretations related to sequence stratigraphy and regional stratigraphic correlations.
The ontogeny of Belosaepia ungulaGabb, 1860 from the Crockett Formation (Bartonian stage, Eocene) of Texas is documented for growth from embryo to old age. During the last stage of life, much skeletal resorption occurred, resulting in a major change in form of the skeleton. The animal produced a large skeleton (to 180 mm in length and 50 mm in diameter) with endogastric coiling, oblique septa and a very large siphuncle. The skeleton has a guard with a solid posterior prong, a posteroventral corona plate and a noded dorsal shield. The ventral margin of the skeleton consists of a thin flattened deck containing strongly recurved septa, conotheca and a secondary prismatic shell layer. New terms are defined for features of the skeleton not previously described. The microstructure of the ventral deck and the presence of a rod structure between the prong and callus are described for the first time. Chamberlets similar to those in living Sepia cuttlebones are present between closely spaced septa and they vary from walled units on lateral margins to pillar form in mid-ventor. The siphuncle is secondarily thickened within the dorsal interior, producing a siphuncle band. The skeleton was produced by a deep-bodied animal of demersal life habits. The species B. uncinata, B. harrisi and B. alabamensis voltzi proposed by Palmer (1937) are synonymised with B. ungula. The species B. veatchii and B. saccaria of Palmer (1937) are considered to be valid species, but B. alabamensis proposed by Palmer (1937) is synonymised with B. veatchii. Descriptions of belosaepiid species must be based only on specimens of adult size that have not been affected by resorption.
Recently collected specimens of the extinct tayassuine peccary “Cynorca” occidentale (and another indeterminant tayassuid) are described from new excavations along the southern reaches of the Panama Canal. Fossil peccaries were previously unknown from Panama, and these new tayassuid specimens therefore add to the extinct mammalian biodiversity in this region. “Cynorca” occidentale occurs in situ in the Centenario Fauna (new name) from both the upper part of the Culebra Formation and overlying Cucaracha Formation, thus encompassing a stratigraphic interval that includes both of these formations and the previously described and more restricted Gaillard Cut Local Fauna. “Cynorca” occidentale is a primitive member of the clade that gives rise to modern tayassuines in the New World. Diagnostic characters for “C.” occidentale include a retained primitive M1, reduced M3, and shallow mandible, and this species is small relative to most other extinct and modern tayassuine peccaries. Based on the closest biostratigraphic comparisions (Maryland, Florida, Texas, and California), the presence of “C.” occidentale indicates an interval of uncertain duration within the early Hemingfordian (He1) to early Barstovian (Ba 1) land mammal ages (early to middle Miocene) for the Centenario Fauna, between about 19 and 14.8 million years ago. Based on what is known of the modern ecology of tayassuines and previous paleoecological interpretations for Panama, “C.” occidentale likely occupied a variety of environments, ranging from forested to open country habitat mosaics and fed on the diverse array of available plants.
African wild dogs (Lycaon pictus) occupy an ecological niche characterized by hypercarnivory and cursorial hunting. Previous interpretations drawn from a limited, mostly Eurasian fossil record suggest that the evolutionary shift to cursorial hunting preceded the emergence of hypercarnivory in the Lycaon lineage. Here we describe 1.9–1.0 ma fossils from two South African sites representing a putative ancestor of the wild dog. The holotype is a nearly complete maxilla from Coopers Cave, and another specimen tentatively assigned to the new taxon, from Gladysvale, is the most nearly complete mammalian skeleton ever described from the Sterkfontein Valley, Gauteng, South Africa. The canid represented by these fossils is larger and more robust than are any of the other fossil or extant sub-Saharan canids. Unlike other purported L. pictus ancestors, it has distinct accessory cusps on its premolars and anterior accessory cuspids on its lower premolars–a trait unique to Lycaon among living canids. However, another hallmark autapomorphy of L. pictus, the tetradactyl manus, is not found in the new species; the Gladysvale skeleton includes a large first metacarpal. Thus, the anatomy of this new early member of the Lycaon branch suggests that, contrary to previous hypotheses, dietary specialization appears to have preceded cursorial hunting in the evolution of the Lycaon lineage. We assign these specimens to the taxon Lycaon sekowei n. sp.
The order Asaphida has been characterized by the possession of a globular protaspis and a ventral median suture. The superfamily Dikelocephaloidea was included in the Asaphida, although there has been no reliable ontogenetic information for this superfamily, and thus whether dikelocephaloids had a protaspis of globular shape has remained unclear. The ontogenetic study of the dikelocephaloid trilobite, Asioptychaspis subglobosa, from Korea reveals that the protaspis of A. subglobosa is not of globular morphology but of general benthic adult-like morphology, heralding that the Dikelocephaloidea may not be closely related to other trilobite groups within the Order Asaphida. This protaspid morphology is clearly distinguished from the highly globular protaspis of the Remopleuridioidea, which has been considered the sister group to the Dikelocephaloidea. In addition, the ontogenetic development of the free cheeks shows that A. subglobosa possessed anteriorly yoked free cheeks during the early phase of development but formed a ventral median suture by splitting the yoked cheeks in the later phase of ontogeny. This contrasts with the previously suggested mode of ventral median suture formation of the Order Asaphida. This alternative mode of ventral median suture formation of A. subglobosa, along with non-asaphoid protaspid morphology, warrants removal of the Dikelocephaloidea from the current Order Asaphida. This study further demonstrates that the ventral median suture did not evolve only once in the evolutionary history of trilobites. With the addition of the mode here documented for Asioptychaspis, the ventral median suture is now known to have developed independently and in a different fashion at least three times in the history of Cambrian trilobites.
The external morphology and its variability, myocardinal apparatus, and shell structure of both valves of the radiolitid rudist Eoradiolites plicatus (Conrad) are described, based on the examination of about one hundred well-preserved, fairly complete specimens that are free of sediment matrix, found in a single bed of an ammonite-bearing Albian succession (‘Knemiceras beds’, Zebbag Formation) in central Tunisia. This study provides the most nearly complete knowledge of any Eoradiolites species to date. With this background, all previous records of E. plicatus, as well as the taxonomic status of the other Albian species of the genus, are revised considering both original material, when available, and published papers. As a result, the systematics of Eoradiolites is simplified.
A large (maximum length 80 mm), tubular, corset-like problematic fossil from the early Cambrian (Cambrian Series 2, Stage 3) Sirius Passet Lagerstätte of North Greenland is interpreted as the lorica of an ancestral loriciferan. In addition to the double circlet of 7 plates composing the lorica, Sirilorica carlsbergi new genus, new species also preserves up to six multicuspidate cuticular denticles that are similar in shape to the pharyngeal teeth of priapulid worms, although their location is suggestive of scalids. Whilst traditionally placed as a sister group of priapulid worms within Vinctiplicata (Scalidophora), recent molecular sequence data suggest that loriciferans might be more closely related to nematomorphs. The limited morphological information available from Sirilorica is consistent with this interpretation, placing the Sirius Passet fossil within the total-group of Loricifera, within the Loricifera Nematomorpha clade.
A specimen of the aquatic reptile Champsosaurus sp. from the Paleocene Black Peaks Formation in southwestern Texas is the southernmost yet known. The fragmentary specimen exhibits some unusual features, such as a great anterior extent of the quadratojugal on the lower temporal arch, and cannot be attributed with confidence to any of the named species. Champsosaurus appears to have been tolerant of temperate climates and had a northern latitudinal range exceeding that of crocodylians. It seems likely that the brief southward extension in range of Champsosaurus during early Paleocene time resulted from a decrease in mean annual temperature, comparable to over 10° of paleolatitude.
An enigmatic microproblematicum, TuxekanellaRiding and Soja (1993), has been identified in the shallow water carbonate succession of the Late Silurian Skala formation (Podolia, Ukraine). The newly found specimens suggest most probably the algal nature of this enigmatic microfossil. Tuxekanella is known only from two extremely distant regions at the moment: the Alexander terrane in Alaska and Podolia. The possibility of their paleobiogeographical link is discussed in the context of paleotectonic models, assuming the peri-Gondwanan origin of the Alexander Terrane.
The Sirius Passet biota of North Greenland is one of the oldest Cambrian lagerstätten, and although it is dominated by non-mineralized arthropods and lobopods, anomalocaridids have never been identified. Based on a single specimen, we herein describe for the first time an appendage with possible anomalocaridid affinities as suggested by an overall gross morphology similar to that of the frontal appendage of Anomalocaris from other localitites. Tamisiocaris borealis n. gen. and n. sp. has an elongated appendage with paired spines along one margin, and differs from the frontal appendage of Anomalocaris in that segment boundaries are absent and ventral spines are relatively long and spineless. These differences may be taphonomic, but the entire surface of the appendage is covered in a fine fabric, making it unlikely that this appendage was originally segmented or sclerotized. The taxon is tentatively placed within Radiodonta, but this systematic placement cannot be confirmed while complete body specimens are lacking.
We describe a Pennsylvanian (Late Carboniferous) Parapostibulla edrioasteroid that competed for living space with an encrusting cryptostome bryozoan on the brachiopod Composita. The colonial bryozoan grew faster than the solitary edrioasteroid, and the bryozoan increased its colony size by slowly encircling the edrioasteroid. Neither organism was able to overgrow the other, even where the margins became thick, and each had to accommodate the presence and possible defenses of the other. Evidence for defensive measures by edrioasteroids has not previously been reported, but this interaction does emphasize that edrioasteroids could not move away from severe crowding or overgrowth by other organisms.
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