Comparison of molecular and paleontological data in diatoms suggests a major gap in the fossil record

Diatoms, unicellular eukaryotic algae with a siliceous skeleton, offer the rare advantage of displaying both an extensive fossil record and numerous extant species, thus providing the opportunity of confronting molecular and paleontological data in a protist group. A portion of the 28S ribosomal RNA was sequenced from 5 diatoms, the divergence times of which are well known. The nucleotide substitution rate was estimated in these unicellular eukaryotes and compared with the rate of multicellular eukaryotes, using a broad data base comprising metazoans and metaphytes. When using fossil record derived divergence times, our results show that the nucleotide substitution rate is about 5 times faster in diatoms than in chordates. But, when using the relative rate test, it is observed that, over a long time period, the nucleotide substitution rate may in fact have been slightly slower in diatoms than in chordates. For this contradiction, two possible explanations are proposed: (i) a failure of the relative rate test, (ii) a gap in the pre-Jurassic diatom fossil record. We have checked that our results concerning the relative rate test were valid. Thus, the second hypothesis, which implies pre-Jurassic diatom evolution, in fact already suggested by some non-molecular evidences, is favoured. Decoupling of morphological differentiation from genetic speciation also appears to have occurred and may account in part for the underestimation of the dates of recent cladogenesis events.     

The fossil record of North American mammals: evidence for a Paleocene evolutionary radiation

Paleontologists long have argued that the most important evolutionary radiation of mammals occurred during the early Cenozoic, if not that all eutherians originated from a single common post-Cretaceous ancestor. Nonetheless, several recent molecular analyses claim to show that because several interordinal splits occurred during the Cretaceous, a major therian radiation was then underway. This claim conflicts with statistical evidence from the well-sampled latest Cretaceous and Cenozoic North American fossil record. Paleofaunal data confirm that there were fewer mammalian species during the latest Cretaceous than during any interval of the Cenozoic, and that a massive diversification took place during the early Paleocene, immediately after a mass extinction. Measurement data show that Cretaceous mammals were on average small and occupied a narrow range of body sizes; after the Cretaceous-Tertiary mass extinction, there was a rapid and permanent shift in the mean. The fact that there was an early Cenozoic mammalian radiation is entirely compatible with the existence of a few Cretaceous splits among modern mammal lineages.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

Autoradiographic demonstration of specific binding sites for E. coli enterotoxin in various epithelia of the North American opossum

Summary In the North American opossum, heat-stable specific binding sites for E. coli enterotoxin are observed (i) in epithelial cells lining the small intestine, colon, gall bladder, cystic duct, common bile duct and trachea, and (ii) in epithelial cells forming the duodenal (Brunner's) glands, liver, kidneys (metanephros, mesonephros) and testis, as demonstrated by autoradiography. Enterotoxin-specific binding sites in the intestinal tract are only found in intestinal epithelial cells with the highest concentration in the microvillus border. Enterotoxin-specific binding sites also occur in epithelial cells comprising the secretory tubules and ducts of the duodenal glands. In the kidneys (metanephros and mesonephros), enterotoxin-specific binding sites are confirmed primarily to the proximal tubules, whereas in the testis they are localized in seminiferous tubules. In the liver, enterotoxin-specific binding sites are confined primarily to hepatocytes. E. coli enterotoxin caused a 7-fold increase of cGMP in the liver and a 30-fold increase in the duodenal glands. The liver responded in about half of the animals studied, whereas the duodenal glands gave a consistent response in each case. Likewise, the duodenal glands consistently showed strong labelling for ¹²⁵I-enterotoxin, whereas receptor labelling of hepatocytes was inconsistent in nearly half the incubations and corresponds to the observed cGMP measurements.Supported by a Weldon Springs Grant, University of Missouri and by funds from the Medical Research Service, Department of Veteran's Affairs     

Patterns of evolutionary transformation in the petrosal bone and some basicranial features in marsupial mammals, with special reference to didelphids

Twelve petrosal and four nonpetrosal characters were coded for representatives of all 15 extant genera of Didelphidae and for 16 additional genera of marsupials representing all extant orders. Three basal metatherians were used as outgroup comparison. Histological sections of a subset of the data were examined. An intermediate position of the hiatus Fallopii supports the monophyly of Didelphidae. Several basicranial regions support different clades within the Didelphidae that recent molecular work has identified, including a sister group relationship of Caluromys and Caluromysiops , the monophyly of large opossums, a Lestodelphys-Thylamys clade, and a Lestodelphys-Thylamys-Gracilinanus-Marmosops clade. Glironia lacks petrosal and jaw synapomorphies of Caluromys and Caluromysiops. The transverse canal, a synapomorphy of the crown-group Marsupialia, opens as a single foramen anterior to the carotid foramen in most marsupials or as numerous foramina in the pterygoid fossa in diprotodontians. It is either intramural (most marsupials) or simply endocranial (most diprotodontians excluding koalas and wombats). Loss of a deep sulcus in the anterior pole of the promontorium for the internal carotid artery and a rostral tympanic process of the petrosal also characterize the groundplan of the crown group Marsupialia. Pouch-young wombats show a groove in the anterior pole of the petrosal for the internal carotid artery. The absence of a prootic canal foramen in the tympanic side of the petrosal of adults supports the monophyly of Australidelphia. Some pouch-young marsupials possess a prootic canal that is later lost in ontogeny. A rather flat promontorium and a crest running medio-distally in the middle of the promontorium characterize Macropodidae.     

Preface: The phylogeny, fossil record and ecological diversity of ostracod crustaceans

After more than two centuries of research, more than 65,000 living and fossil ostracod species have been described and studied, yet much remains to be learned about this ancient, widespread and diverse group of bivalved arthropods. Their higher classification and phylogeny are subjects of vigorous debate, as is their position in the broader picture of crustacean phylogeny. At the same time, major advances in our understanding of ostracod lineages and their relationships are resulting from the application of innovative approaches and techniques. This preface provides a contextual overview of the 15 contributions to this volume, which resulted from the 14th International Symposium on Ostracoda (ISO2001) held in 2001at Shizuoka, Japan. As such it provides a cross-section of topics at the forefront of research on the evolution and diversity of Ostracoda, and indicates directions for future work.     

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine 'collectorship' since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities ( Otogornis , Gobipteryx ); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.     

Phylogeny and adaptive diversity of rodents of the family Ctenomyidae (Caviomorpha): delimiting lineages and genera in the fossil record

Differentiation of genera of the modern (Late Miocene to Recent) South American rodent family Ctenomyidae would have been linked to the acquisition of disparate adaptations to digging and life underground. In accordance with this hypothesis, the delimitation of lineages and genera in the ctenomyid fossil record is evaluated here following an adaptation-rooted criterion that involves both an assessment of the monophyly and of the adaptive profiles of recognized clades. The application of such a criterion, including morphofunctional information, delimited four cohesive lineages among crown ctenomyids (i.e. euhypsodont species of the Late Miocene to Recent): Eucelophorus (Early Pliocene–Middle Pleistocene), Xenodontomys-Actenomys (Late Miocene–Pliocene), Praectenomys (Pliocene) and Ctenomys (including Paractenomys ; Pliocene–Recent); in addition, the results supported the status of Xenodontomys as a paraphyletic ancestor of Actenomys . The cladogenesis that gave rise to the crown group would have occurred immediately after the acquisition of euhypsodonty in a Xenodontomys simpsoni -like ancestor during the Late Miocene. This putative ancestor would have had fossorial habits and moderate digging specializations, an adaptive profile maintained in Xenodontomys-Actenomys . Eucelophorus and Ctenomys would have independently evolved subterranean habits at least since the Pliocene. Although the earliest history of the only living representative, Ctenomys , is known only fragmentarily, remains from Esquina Blanca (Uquía Formation), in north-western Argentina, suggest a minimum age of around 3.5 Ma (Early–Late Pliocene) for the differentiation of the genus. This date agrees with recent molecular estimates.     

Cone pigments in a North American marsupial, the opossum (Didelphis virginiana)

Only two of the four cone opsin gene families found in vertebrates are represented in contemporary eutherian and marsupial species. Recent genetic studies of two species of South American marsupial detected the presence of representatives from two of the classes of cone opsin genes and the structures of these genes predicted cone pigments with respective peaks in the ultraviolet and long-wavelength portions of the spectrum. The Virginia opossum (Didelphis virginiana), a profoundly nocturnal animal, is the only marsupial species found in North America. The prospects for cone-based vision in this species were examined through recordings of the electroretinogram (ERG), a commonly examined retinal response to photic stimulation. Recorded under flickering-light conditions that elicit signals from cone photoreceptors, the spectral sensitivity of the opossum eye is well accounted for by contributions from the presence of a single cone pigment having peak absorption at 561–562 nm. A series of additional experiments that employed various chromatic adaptation paradigms were conducted in a search for possible contributions from a second (short-wavelength sensitive) cone pigment. We found no evidence that such a mechanism contributes to the ERG in this marsupial.     

First record of a Late Proterozoic faunal assemblage in the North American Cordillera

Hofmann, Hans J. 1981 12 15: First record of a Late Proterozoic faunal assemblage in the North American Cordillera. Lethaia, Vol. 14, pp. 303–310. Oslo. ISSN 0024–1164. Megafossils are reported from an unnamed Late Proterozoic formation in the Sekwi Brook area of the Mackenzie Mountains of northwestern Canada. The biota includes the metazoans Inkylovia sp. and Sekwia excentrica Hofmann n, gen., n. sp., the trace fossils Gordia sp., Gordiu? sp., and Torrowangea sp., as well as structures of problematic affinities. The metazoans support a latest Proterozoic age for the containing beds. 0 Early Metazoa, body fossils, trace fossils, Problematica, Late Proterozoic, North American Cordillera, Markenzie Mountains, Canada.     

Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record

Schisandraceae are traditionally subdivided in two genera, Schisandra and Kadsura, based on differences in the organisation of the floral receptacle, the carpels, and the presence or absence of a ``pseudostigma'. Recently, phylogenetic analyses utilizing ITS sequence data and morphological data resulted in incongruent tree topologies, with the morphological trees suggesting monophyly of the two genera, whereas ITS trees did not resolve Schisandra and Kadsura as monophyletic clades. In the present paper we study seed morphology and leaf epidermal features of 22 species of Schisandraceae in order to provide additional data for a morphological data matrix. Seed morphological characters are highly homoplastic and do not yield further evidence for monophyly of the two genera. Instead, a number of characters appear to support sister group relationships between taxa within the genera, such as, for instance, for K. coccinea and K. scandens, both of which have large seeds along with a multi-layered mesotesta. Considering leaf epidermal characteristics, species of Kadsura were found to be consistently amphistomatic, whereas species of Schisandra are always hypostomatic. Phylogenetic analysis using the extended data matrix resulted in weakly supported Kadsura and Schisandra clades with five and four synapomorphies indicating monophyly of Kadsura and Schisandra, respectively. Fossils ascribed to Schisandraceae date back to the Late Cretaceous. These are tri-and hexacolpate pollen types displaying a combination of features found in modern Schisandraceae and partly also in Illiciaceae. Leaf remains from this period are poorly preserved and difficult to ascribe to Schisandraceae because of the lack of synapomorphies for the family. In the Early Cainozoic, leaf and seed remains from North America and Europe unambiguously belong to the family. Seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, while fossils from Europe show more similarities to modern Asian species.     

Dispersal and diversity in the earliest North American sauropodomorph dinosaurs, with a description of a new taxon

Sauropodomorph dinosaurs originated in the Southern Hemisphere in the Middle or Late Triassic and are commonly portrayed as spreading rapidly to all corners of Pangaea as part of a uniform Late Triassic to Early Jurassic cosmopolitan dinosaur fauna. Under this model, dispersal allegedly inhibited dinosaurian diversification, while vicariance and local extinction enhanced it. However, apomorphy-based analyses of the known fossil record indicate that sauropodomorphs were absent in North America until the Early Jurassic, reframing the temporal context of their arrival. We describe a new taxon from the Kayenta Formation of Arizona that comprises the third diagnosable sauropodomorph from the Early Jurassic of North America. We analysed its relationships to test whether sauropodomorphs reached North America in a single sweepstakes event or in separate dispersals. Our finding of separate arrivals by all three taxa suggests dispersal as a chief factor in dinosaurian diversification during at least the early Mesozoic. It questions whether a 'cosmopolitan' dinosaur fauna ever existed, and corroborates that vicariance, extinction and dispersal did not operate uniformly in time or under uniform conditions during the Mesozoic. Their relative importance is best measured in narrow time slices and circumscribed geographical regions.     

Phylogenetic analysis of life-history adaptations in parasitic cowbirds

Parasitic cowbirds lay eggs in the nests of other species anddupe them into caring for their young. Unlike other brood parasites,cowbirds have not developed egg mimicry or bizarre chick morphology.However, most of them parasitize a large number of hosts. Severalfeatures of cowbirds have been proposed as more general adaptationsto brood parasitism. In this study, we used a recent molecularphylogeny as a historical framework to test the possible adaptationsof the parasitic cowbird, including egg size, eggshell thicknessand energy content of the eggs, length of the incubation period,and growth pattern of cowbird nestlings. We used a recentlydeveloped extension of independent contrasts to test whetherthe five cowbird species deviate from general allometric equations.We generated prediction intervals for a nonparasite that evolvedin the place of the cowbirds. By using these prediction intervals,we found that parasitic cowbirds had not reduced weight or energycontent of their eggs, nor their incubation period over evolutionarytime. Cowbird chicks and those of nonparasitic relatives hadsimilar growth pattern. The only characteristic that separatedparasitic cowbirds from their nonparasitic relatives was anincrease in eggshell thickness. All these findings were robustand resisted the use of three models of character evolution.The fact that most traits exhibited by cowbirds were inheritedfrom a nonparasitic ancestor does not rule out that they areadvantageous for parasitism. Future research should focus onsuch traits of cowbird relatives and on how these traits preadapteda particular lineage to become parasites.     

A preliminary phylogeny of the Pterasteridae (Echinodermata,Asteroidea) and the first fossil record: Late Cretaceous of Germany and Belgium

The Pterasteridae comprises a diversified group of extant largely deep-sea starfishes. Generic diagnoses have been based classically on soft tissue characters and skeletal architecture. A preliminary phylogeny of sixteen extant species is here worked out by cladistic analysis. The resulting tree suggests monophyly of extant genera and the validity of dissociated plates for identification of genera. Fossil remains of Pterasteridae are here described for the first time. By comparison with extant species, all the skeletal remains from the lower Upper Campanian of Belgium and from the lower Maastrichtian of Germany are tentatively assigned to the genusPteraster. The fossil record of starfishes is poor, but the present Late Cretaceous pterasterids provide one more piece of evidence of the high diversity of starfishes during the Mesozoic. Known Late Cretaceous and Paleogene fossils are broadly similar, which suggests the end-Cretaceous extinction event did not cause major turnover in asteroid faunal composition. As suggested for other starfish groups, both the fossil record of deep-sea Pterasteridae in shelf settings and tree topology imply an onshore-offshore evolutionary trend.