Tumulduria incomperta and the case for Tommotian trilobites

The enigmatic fossil Tumulduria incomperta Missanhevsky 1969 from the basal Tommotian Stage at 'Dvortsy' on the Aldan River, Siberian Platform, is reinvestigated in light of the report (Fedorov et al. 1979, Dokl. AN SSSR 249) of trilobite remains from the same beds. Comparisons of these supposed trilobites with old and new collections from 'Dvortsy' leave no doubt that they are identical to Tumulduria incomperta. Tumulduria is represented by phosphatic plates with a crude bilateral symmetry, consisting of a central longitudinal rounded ridge flanked by flat lateral portions. They are built of growth lamellae overlapping each other along the axis of symmetry. The surface carries prominent transverse folds and lamellar terminations. There is considerable morphological variation, and the similarity of some specimens to trilobites is only superficial. Tumulduria is interpreted as a bilaterally symmetrical metazoan with a dorsal protective plate; it probably represents a short-lived group that left no descendants.     

Hyostragulum simplex n. sp. (Incertae sedis) du dévonien du massif armoricain intérêt paléobiogéographique et systématique

In the western part of the “Synclinorium median armoricain (Rade de Brest), the presence of Hyostragulum simplex n. sp. in the lower part of the Troaon formation (Upper Emsian) is a supplementary element of hercynian fauna in Brittany during lower devonian time.For the new species, characterised by the lack of a septum on the base of the corallite, a fibrous microstructure, perhaps trabecular, is shown. The systematic position of the genus is discussed but ends up with a suggestion to let it remain incertae sedis..     

Rings without a lord? Enigmatic fossils from the lower Palaeozoic of Bohemia and the Carnic Alps

Fossilized ring‐like structures with enigmatic function and taxonomic affiliation were recovered for the first time from the Upper Ordovician of the Carnic Alps and the Silurian of Bohemia. These rings, already mentioned as minor constituents in previous conodont studies (e.g. Webers 1966, p. 1; Bischoff 1973, p. 147), were reported from the Palaeozoic of several regions in Europe and North America. Originally considered as inwardly accreted adhering discs of a benthic hyolithelminth worm with a phosphatic tubular projection, they were later reinterpreted in relation to a putative crinoid epibiont or even as possible scyphozoans. Despite a long debate, neither the function of the enigmatic Palaeozoic rings nor their taxonomic affiliation has been fully clarified. The studied material, extracted by a standard technique in use for conodonts, consists of 235 elements from 16 stratigraphic levels in the Plöcken Formation (Carnic Alps, Cellon Section; Amorphognathus ordovicicus Biozone, Hirnantian, Ordovician) and in the Kopanina Formation (Bohemia, Mu?lovka Quarry; Polygnathoides siluricus Biozone, Ludfordian, Silurian). To explore whether ring size and shape changed over time, we employed a novel combination of geometric morphometric approaches for outlines with no ‘homologous’ landmarks and showed that only size appreciably varied with an increase of ca. 20%. The emerging data from this study are consistent with the interpretation of the rings as an adhering structure of a benthic organism living on a relatively uniform hard substrate.     

Strengths abd weaknesses of the reef guild concepts and quantitative data: Application to the upper Capitan-massive community (Permian), Guadalupe Mountains, New Mexico-Texas

Summary Analyses of large acatate sheet tracings, close-up photos and 105 sub-horizontal quadrat surfaces at four localities near the base of the Guadalupe Mountains Escarpment indicate that the biotic framework of the upper Capitan reef was built by about 35 species: one codiacean (Eugonophyllum sp.), 17 calcisponges, 9 bryozoans, one richthofenid brachipod, some crinoid (known only from columns), 4 Problematica and microbes. This widespread fossil community included members of the Constructor, Baffler and Binder Guilds. A re-evaluation of the Guild Concept (Fagerstrom, 1987, 1991) highlights the validity of the functional roles of the Constructor and Binder Guilds for reef construction. Members of the Baffler Guild, however, need to be revised and an interpretation of microbial micrite and cryptic biota remains controversial. Open surface phylloid algal and cryptic sponge-bryozoan dominated sub-communities were of only local importance. The upper Capitanmassive differs from its Permian conterparts in the low diversity and areal cover of the frame-building biota, low micrite content and abundant micro-frameworks, i.e, intergrown small sponges, Problematica and syndepositional cements (botryoidal and isopachous, fibrous calcite). Quantitative areal cover data were assessed at various scales. Large acetate sheets generally have low coverage of macro-biota (5.4%). By contrast, analysis of small areas of local high areal cover (selected acetate sheet quadrats, subvertical photographs, and quadrat samples: 15–21%) provide detailed insights into clustered patches forming the inital reef framework. Both data sets provide useful clues for an integrated approach to framework assessment. Mean acetate sheet data are limited by their somewhat generalized pattern, while small investigation areas may overemphasize local variation. Erect and pendant sponges with solitary, sub-cylindrical and multi-branche/clonal forms, were the predominant initial frame-builders in both open surface and cryptic habitats. Selective larval recruitment of erects sponges to firm substrates produced continous upward accretion of the initial framework. On open surfaces and in pores formed by tabular sponges and fenestrate bryozoans, erect and pendant sponges were supported in their hydrodynamically unstable growth position by encrusters, chieflyArchaeolithoporelle hidensis, Shamovella obscura, an unnamed tubular organism, and microbes. Subsequent growth ofArchaeolithoporella hidensis, microbial crusts and syndepositional cements on the outer walls of live sponges would have impeded ambient water circulation and may have led to ‘creeping sponge death by suffocation’ or complete encrustation after death. Filling of pores in the initial and encrusted reef framework by internal sediment (packstone-grainstone; derived from the framework and the back-reef shelf/platform) and voluminous syndepositional marine-phreatic cements completed the framebuilding process.     

Mobergellans (Problematica) from the Cambrian of Greenland, Siberia and Kazakhstan

Mobergellans are disc-shaped problematic fossils of phosphatic composition bearing prominent muscle attachment structures on the presumed internal surface. They are found exclusively in sediments of Early Cambrian age, mainly from Laurentia, Baltica and Siberia. Type material ofDiscinella brastadi Poulsen, 1932 from North-East Greenland is re-studied and synonomised withDiscinella micans (Billings, 1871). Specimens from Siberia described asD. brastadi are re-assigned toMobergella hexactina n. sp., and specimens from Siberia described asMobergella radiolata Bengtson, 1968 are re-assigned toM. sibirica n. sp.Aktugaia triangula Mis-Sarzhevsky, 1976 from Kazakhstan is redescribed, and its relationship to other mobergellans is discussed.      

The nature of Radiocyathidsand their relationship to Receptaculitids and Archaeocyathids

The morphological analysis of the structural plan of the Radiocyathids skeleton implies a closer relationship to Receptaculitids than to Archaeocyathids. As a working hypothesis, they may possibly form a link between part or all of these two groups and use as a test for the validity of the new kingdom Archaeata proposed by I. T. Zhuravleva & E. I. Miagkova.     

Problematica old and new

Problematica are taxa that defy robust phylogenetic placement. Traditionally the term was restricted to fossil forms, but it is clear that extant taxa may be just as difficult to place, whether using morphological or molecular (nucleotide, gene or genomic) markers for phylogeny reconstruction. We discuss the kinds and causes of Problematica within the Metazoa, as well as criteria for their recognition and possible solutions. The inclusive set of Problematica changes depending upon the nature and quality of (homologous) data available, the methods of phylogeny reconstruction and the sister taxa inferred by their placement or displacement. We address Problematica in the context of pre-cladistic phylogenetics, numerical morphological cladistics and molecular phylogenetics, and focus on general biological and methodological implications of Problematica, rather than presenting a review of individual taxa. Rather than excluding Problematica from phylogeny reconstruction, as has often been preferred, we conclude that the study of Problematica is crucial for both the resolution of metazoan phylogeny and the proper inference of body plan evolution.     

Three-dimensional anatomy of the Tully monster casts doubt on its presumed vertebrate affinities

Tullimonstrum gregarium, also known as the Tully monster, is a well-known phylogenetic enigma, fossils of which have been found only in the Mazon Creek Lagerstätte. The affinities of Tullimonstrum have been debated since its discovery in 1966, because its peculiar morphology with stalked eyes and a proboscis cannot easily be compared with any known animal morphotypes. Recently, the possibility that Tullimonstrum was a vertebrate has attracted much attention, and it has been postulated that Tullimonstrum might fill a gap in the fossil record of early vertebrates, providing important insights into vertebrate evolutionary history. With the hope of resolving this debate, we collected 3D surface data from 153 specimens of Tullimonstrum using a high-resolution laser 3D scanner and conducted x-ray micro-computed tomographic (μCT) analysis of stylets in the proboscis. Our investigation of the resulting comprehensive 3D morphological dataset revealed that structures previously regarded as myomeres, tri-lobed brain, tectal cartilages and fin rays are not comparable with those of vertebrates. These results raise further doubts about its vertebrate affinities, and suggest that Tullimonstrum may have been either a non-vertebrate chordate or a protostome.     

A myriapod-like arthropod from the Upper Cambrian of East Siberia

Although the fossil record of biramous arthropods commences in the Lower Cambrian, unequivocal uniramous arthropods do not appear until the Upper Silurian, in association with terrestrial biotas. Here we report an Upper Cambrian marine arthropod from East Siberia that possesses some significant myriapodan features. The new arthropod,Xanthomyria spinosa n. gen., n. sp., closely resembles examples of archipolypodans from the Late Palaeozoic. If this resemblance genuinely represents myriapod affinities, this would be the first convincing myriapod from the Cambrian. Suggestions of an early branching point of the myriapods from other arthropods would be consistent with this. Conversely, an as yet poorly known clade of multi-segmented arthropods may exist in the Cambrian, with no close affinities to the myriapods.