Internal supporting structures of some Late Ordovician graptolites from South China revealed by micro-CT

Three-dimensionally preserved pyritic internal moulds of the Late Ordovician graptolites Appendispinograptus leptothecalis and Styracograptus chiai from the Upper Ordovician Wufeng Formation of the Daheba section, Hunan Province, South China, were imaged by micro-CT. The distinctive pea-shaped pits/dimples aligned in a zigzag line arising from the second/third thecal pair on the tubarium in these species are interpreted as the surface expression of the junction between the lateral thecal walls and the thickened internal cross-bars that support the nema. The relations between different internal structures such as the nema, median septum, aboral lists, interthecal septa and cross-bars are summarized, and examples of these supporting systems in different taxa are illustrated. The presence of isolated cross-bars in different diplograptid tubarium may suggest that they have evolved as a result of constructional restrictions.     

A first report of Prolinograptus packhami Rickards and Wright, 1997 from Baltica,Poland

A well-preserved isolated graptolite Prolinograptus packhami Rickards and Wright, 1997, well known from Australia, is described for the first time in Europe. The genus has now been recognized in three continents (Australia, Europe and North America). The Australian P. packhami occurs in the Neocucullograptus inexpectatus/Neocucullograptus kozlowskii biozonal interval, while the form from the Mielnik IG-1 core, Poland, appears earlier, in the Bohemograptus praecornutus Biozone. Prolinograptus, of unclear origin, is one of the least-known genera of the subfamily Linograptinae. P. packhami, along with Neodiversograptus palmeri and Prolinograptus orangensis, form a group of very diminutive specimens, with slender stipes and thecal apertures showing varying degrees of eversion. Scanning electron microscope examination revealed the presence of some micro-ornamentation on the rhabdosome surface of the Polish Prolinograptus, similar to that known in retiolitids (Retiolitidae).     

Rhabdotubus, a Middle Cambrian rhabdopleurid hemichordate

Rhabdotubus johanssoni n.gen., n.sp., is described from the early Middle Cambrian Eccaparadoxides pinus Zone of Närke, southern Sweden. The colonies encrusted shells of inarticulate brachiopods, and occasionally trilobites, on otherwise soft substrates. The tubarium consists of repent and erect tubes. The former branch irregularly and produce a thecorhiza-like structure; the latter are erect and mostly isolated, up to 10 mm in length and widening gradually to about 1 mm width. Both repent and erect tubes are composed of fusellar bands, mostly irregularly arranged. Branching of repent tubes takes place through resorption or perforation of fusellar tissue in the parent tube. Branching of erect tubes occurs sporadically. There is no thecal dimorphism. No sclerotized stolon is present. Rhabdotubus is interpreted as the earnest known rhabdopleurid (Class Pterobranchia, Phylum Hemichordata). In general habitus it is similar to sessile graptolites of the Order Tuboidea. These similarities may well have phylogenetic significance, but further knowledge of the Tuboidea and other sessile orders of the Graptolithina is required to clarify the early evolution of graptolites.     

New observations on the fine structure of graptoblasts

Graptoblasts are redefined as widenings of the sealed terminal portions of graptolite stolothecae resulting in ovoid bodies composed of an outer fusellar layer, named here as blastotheca, and an inner secondary layer of electron dense material, named the blastocrypt. In their proximal portion a number of graptoblasts display a scar with fuselli stripped and the crassal material of the blastotheca exposed and displaying a rough and dull surface. The boundary of this area, termed the talus, is edged by a distinct escarpment, probably the margin of the preserved part of the blastotheca. The outer wall of the talus is usually abruptly terminated. Only exceptionally is the outer wall conical and gradually widening. Thus the transverse partition between the parental stolotheca and the graptoblast was produced by the blastocrypt without fusellar covering. It seems, therefore, that graptoblasts were most probably lacking any internal portion of their blastothecae which could possibly be compared with the internal parts of a dendroid auto-theca. It is concluded that graptoblasts housed a zooid that never functioned as an autozooid, but suffered arrested development and encystation. The biological role of graptoblasts as dormant bodies is discussed.     

Collagen in the pterobranch coenecium and the problem of graptolite affinities

Amino-acid analysis of the extracellular skeleton (coenecium) of two pterobranchs (Hemichordata) Rhabdopleura normani and Cephalodiscus (C.) hodgsoni shows that both contain considerable quantities of collagenous material with relatively high hydroxyproline and low hydroxylysine levels. The appearance of the fibrous material in the skeleton of pterobranchs, although collagenous, differs from standard EM characteristics of collagen. The identification of the collagenous nature of the pterobranch skeleton with the presumed presence of collagen-like material in the periderm of fossil graptolites, taken in conjunction with other data, supports the hypothesis that both groups may be closely related phylogenetically.     

Stratigraphical distribution of the Ordovician graptolite Azygograptus Nicholson & Lapworth in the Central Andean Basin (northwestern Argentina and southern Bolivia)

We analyzed new occurrences of Azygograptus lapworthi from the Cordillera Oriental, Argentina. The bearer sandstones levels, corresponding to the Acoite Formation, are overlying the deposits, in which the Didymograptellus bifidus Biozone (Lower Ordovician, upper Floian, Fl3) was previously recognized, and are overlain by younger pelitic levels yielding Xiphograptus lofuensis (Middle Ordovician, early Dapingian, Dp2). Previous records from the Central Andean Basin are also reviewed in detail and accurately correlated, allowing us to conclude that the Azygograptus lapworthi Biozone corresponds to the Middle Ordovician (lower Dapingian, Dp1). This biostratigraphic framework documents that the transition between the Lower and Middle Ordovician deposits occurs in the uppermost levels of the Acoite Formation in the Argentine Cordillera Oriental. It is additionally integrated with up to date conodont records to establish a high-resolution regional correlation, with equivalent deposits from the Puna of northwestern Argentina and Cordillera Oriental of Bolivia, and to discuss new insights for global correlation.     

The restoration of flattened fossils

Neither collapse, due to decay in a soft-bodied organism, nor compaction, due to overburden pressure, normally lead to significant lateral expansion in flattened fossils except in the case of some with rigidly mineralized skeletons. The fossils are thus analogous to a variety of two-dimensional views of a three-dimensional object. This realization provides a foundation for drawing and testing a reconstruction using either computer or manual graphic restoration methods. A complementary approach based on the photography of simple models, which is particularly useful where a complex three-dimensional morphology is under study, is described and illustrated by two examples, the Middle Cambrian arthropod Odaraia and the Upper Ordovician graptolite Direllograptus .     

Latest Ordovician graptolites from the Mandalay Region,Myanmar

The graptolite fauna of the Mandalay Region, Myanmar, is poorly known. In this paper, 13 species in the genera Avitograptus, Korenograptus, Metabolograptus, Neodiplograptus and Normalograptus are described from the uppermost Ordovician strata of the area. Two of these species, Korenograptus selectus X. Chen n. sp. and Neodiplograptus mandalayensis X. Chen n. sp., are new. On the basis of the graptolite faunas, a biostratigraphic scheme for the uppermost Ordovician (Hirnantian: Metabolograptus extraordinarius Biozone and Metabolograptus? persculptus Biozone?) strata in this area is erected.