Ocular structures in an Ordovician agnostid trilobite

Ocular structures have recently been discovered on the cephalon of an Ordovician agnostid trilobite, Oculagnostus frici (Holub, 1908). They are represented mainly by obvious palpebral lobes situated at the postero-lateral part of the genae. In addition, in one specimen there is a dome-shaped structure below the palpebral lobe. This may well be the remains of an eye, but details of its structure are indeterminate. In the other two cephala to hand there is a notch or cavity beneath the palpebral lobe, indicating that the visual surface may have been encompassed by a circum-ocular suture such that the lentiferous region normally dropped out during ecdysis. The origin of the ocular structures in Oculagnostus frici is briefly discussed. Oculagnostus n. gen. is described.     

Exoskeletal structure in the Ordovician trilobite Flexicalymene

Scanning Electron Microscope study of the exoskeletal ultrastructure of secondarily phosphatized material of Flexicalymene sp. from the Upper Ordovician Maquoketa Shale, Iowa, USA, shows that the exocuticle, comprising 20% of the total exoskeletal thickness, is composed of horizontal laminar units between 0.2 and 1 μm thick. These units consist of primarily mineralized organic fibres which form horizontal laminae interconnected by inter-laminae. The endocuticle is considerably more mineralized than the exocuticle, and its original organic structure cannot be observed in untreated preparations. Etching with chromium sulphate reveals: (1) horizontal organic laminar units, 0.2 to 2 μm thick, and (2) pore canals with non-twisted walls about 0.3 μm in diameter. Exuvia cannot be distinguished from the exoskeletons of dead animals. The exoskeletal ultrastructure in trilobites agrees essentially with that in crustaceans.     

An opportunistic Upper Ordovician trilobite assemblage from Missouri

The trilobite Ampyxina bellatula (Savage) occurs in monospecific accumulations within dolomitic strata of the shaly Upper Ordovician Maquoketa Group of northeastern Missouri. A dolomitic stratum containing 97 complete individuals and considerable amounts of pyrite exhibited no associated benthic megafauna. The strata appear to be equivalent to the Scales Formation of the Maquoketa Group of northern Illinois and eastern Iowa which have been interpreted as having been deposited in a muddy environment with severe circulation restrictions. The presence of the monospecific trilobite association in what otherwise would be an unfossiliferous stratigraphic sequence is interpreted as being the result of opportunistic strategy by this trilobite in a resource-rich unstable environmental setting. The association was apparently cat-astrophically buried, thus preserving the age structure of the holaspid population. A size-distribution histogram of the holaspid population indicates that reproduction may have occurred in periodic bursts with lesser reproductive events occurring during the intervening periods. Holaspid survivorship information indicates that mortality was low during the early holaspid stage and increased with age. Anomalous single bed trilobite accumulations of various age rocks have been documented by numerous authors. These may in part be analogous to the Ampyxina occurrence in the Maquoketa Group of Missouri, and be attributable to opportunistic strategies.     

Evolution in a cline: the trilobite Triarthrus along an Ordovician depth gradient

Head proportions in the trilobite Triarthrus change by small but significant amounts with both lime and environment along a 'fossil' depth gradient in the Trenton Group of central New York. The putative zone species T. becki (Denmarkian Stage) and T. eatoni (post-Denmarkian; = T. becki , new synonymy) intermingle throughout the roughly two million year long sequence studied. The two represent extreme morphs in a graded spatio-temporal cline, not distinct species. Transgression as much as evolution accounts for their stratigraphic zonation. Clinal variation is expressed in the dependence of anatomical measures on the position of samples on a downslope transect parallel to bentonite beds, and on ordination score for community samples, a measure of environment that applies over time. Anatomy changes with environment in basically the same way it changes with time, and the amount of change over two million years in a given environment roughly equals the amount of change with environment at a given time. Compensated to a standard environment, anatomical measures follow a continuous trend that suggests an evolutionary random walk with non-random, directional drift. The case of Triarthrus illustrates that directional anatomical variation can have important components that are not strictly temporal and evolutionary in the normal sense.     

The life habits of the Ordovician illaenine trilobite Bumastoides

The Ordovician illaenine Bumastoides exhibits a distinctive effaced and strongly convex morphology. Orientation of the visual field, the extreme convexity of the cephalon and the nature of the thoracic articulation support an interpretation of Bumastoides as an infaunal trilobite that was poorly suited to epifaunal crawling. The genus may have been sedentary; spending most of its post-larval life cycle within a burrow. Suspension feeding would be a viable existence for a sedentary trilobite such as Bumastoides. Maintenance of a burrow is essential for respiration and would require a firm, cohesive substrate. The infaunal niche had become occupied by trilobites by at least the Late Cambrian and continued to be exploited through the Ordovician, Silurian and. possibly, into the Devonian. Convergence has led to the appearance of the effaced, strongly convex morphotype in a number of unrelated families, including the Illaenidae, Asaphidac, Aulacopleuridae, Plethopeltidae and Scutelluidae. A high numerical abundance of illaenid trilobites, such as Illaenus and Bumastoides , is characteristic of the illaenid–chcirurid association, which persisted from the early Ordovician until at least the Late Silurian. This association has been recorded from shelf-edge and on-shelf carbonate buildups and shallow subtidal level bottom environments. It appears to be confined to firm substrates.     

A complete styginid trilobite from the Ordovician of Sweden

Styginidae are a small group of trilobite species, which are usually rare and most of which are incompletely known. Here we describe a complete specimen of Raymondaspis grandigena n.sp. from the Middle Ordovician (Darriwilian, upper Arenig) of Sweden. Among the group it has an unusual combination of exceptionally large genal spines, a thorax with notably short pleural spines in the anterior tergites, and a wide concave pygidial margin. The hypostome is documented for the first time in situ in a styginid, and its attachment can be best described as semi-impendent.     

First cladistic analysis of the trilobite family Olenidae from the Furongian and Ordovician

The Olenidae stands out for its abundance and biostratigraphical importance, especially in the Lower Palaeozoic rocks of northwestern Argentina. Their phylogenetic relationships have been traditionally determined stratigraphically and by direct morphological comparison. This study reports the first formal phylogenetic analysis of olenids. Eighty‐six characters (24 quantitative and 62 qualitative) were coded for 65 taxa (58 olenids). Quantitative characters were treated both as discrete and as continuous variables. To explore the best way of character coding for this group, continuous characters were coded as: median, log‐median, normalized and rescaled. Maximum parsimony and implied weighting were used as optimality criteria. A phylogenetic hypothesis more consistent with traditional taxonomy was reconstructed with both quantitative and qualitative partitions. All the trees obtained with quantitative characters coded as continuous and rescaled are better resolved, and those topologies were more similar among them. This treatment also reflects more effectively the behaviour of the original variables. Olenidae is not a monophyletic clade: Andrarina costata and Aphelaspis australis are included within the ingroup, as sister clade of Olenus gibbosus. Also, the results suggest that members of the Hypermecaspidinae constitute a new family within the Order Olenida. The traditional taxonomic scheme at subfamily level is partially supported. Triarthrinae and ‘pelturinds’ are recovered as monophyletic clades, but Oleninae is polyphyletic. This study proves, through a formal cladistic analysis, that characters disregarded by traditional taxonomy can be uncovered. Finally, this is the first step towards achieving a classification of the Olenidae taking into account the evolutionary process involved in its diversification history.     

Rapid microglial activation induced by traumatic brain injury is independent of blood brain barrier disruption

Following CNS injury, microglia respond and transform into reactive species exhibiting characteristic morphological changes that have been termed "activated" or "ameboid" microglia. In an attempt to establish that microglial reactions induced immediately after injury are caused by intrinsic mechanisms rather than infiltration of blood and its constituents, oxygenized Ringer's solution was perfused into the cerebral circulation of rats so that the circulating blood could be eliminated prior to injury induction. Under artificial respiration, a catheter was inserted from the cardiac apex into the ascending aorta, and oxygenized Ringer's solution was immediately perfused with a pulsatile blood pump, resulting in wash out of the circulating blood from the brain within 1 min. Subsequently, a cortical contusion was induced in the unilateral parietal cortex using a controlled cortical impact (CCI) device. At 5 min following the injury, the brain was fixed by perfusion of fixative through the catheter and removed. Coronal vibratome sections were then processed for CR3 immunohistochemistry to examine the microglial activation. It appeared that microglial activation with both morphological transformation and an increase in CR3 immunoreactivity was induced throughout the hemisphere ipsilateral to the injury side exclusively, even in rats with elimination of circulating blood. The microglial reactions did not differ substantially from those observed in the control rats with extensive BBB disruption. The present results thus provide direct evidence that the microglial activation induced immediately after injury is independent of infiltration of circulating blood induced by concurrent BBB disruption.     

Clinal variation, episodic evolution, and possible parapatric speciation: the trilobite Flexicalymene senaria along an Ordovician depth gradient

Pygidial segmentation in the trilobite Flexicalymene senaria changes with time (as measured in relation to bentonite beds) and with depth (as measured along downslopc transects along bentomtc beds and as measured by gradient analysis of fossil assemblages) along a 'fossil' depth gradient in the Middle Ordovician Trenton Group in central New York State. The pattern of strictly temporal change resembles a 'punctuation' between 'equilibria'. This evolutionary episode, a 10⁶ year interval of continuous change, may represent parapatric speciation. The clinal gradient is strongest during this episode and is weak or not apparent outside it. Strictly temporal change takes place by way of temporal change in the cline's shape. □ Evolution, geographic variation and dines, gradient analysis. Middle Ordovician, morphometry, speciation, Trilobita.     

Diversity patterns in upper Cambrian to Lower Ordovician trilobite communities of north‐western Argentina

The hierarchical structure of biodiversity from a regional scale analysis has received much attention as an alternative approach to unravelling the principal drivers of biodiversification. To better understand the processes that control the diversification of Cambro‐Ordovician trilobite communities from the Argentine Cordillera Oriental, we explore patterns of occupancy and diversity trajectories at the local and regional scales through seven intervals (Furongian, loTr1, upTr1, loTr2, upTr2, Tr3 and Fl2–3), and across an onshore‐offshore profile. Our results indicate: (1) a decrease in regional diversity from the upper Tr2 onwards, mainly caused by a reduction in the number of rare taxa, coupled with stable beta diversity at regional scale and a constant rise in beta diversity in deep subtidal environments; (2) a higher proportion of regional diversity allocated to the within‐habitat beta component; and (3) that changes in gamma diversity are driven primarily by changes in alpha diversity during the Furongian–Tr3, whereas in the Floian, beta diversity seems to modulate regional diversity. These trends and associated patterns indicate increasing ecological differences among taxa, shifting from metacommunities where most taxa have similar ecological preferences or ‘Hubbell type’ to metacommunities with high niche differentiation or ‘Hutchinson type’. Interestingly, the timing of this shift coincides with the regional‐scale turnover between trilobite evolutionary faunas suggesting that the rise in niche differentiation among these genera may be related to the transition. Superimposed on this general trend, particular diversity structures can be understood in the light of metacommunity dynamics, such as dispersal limitation and mass effect.     

Unusual trilobite biofacies from the Lower Ordovician of the Argentine Cordillera Oriental: new insights into olenid palaeoecology

Balseiro, D., Waisfeld, B.G. & Buatois, L.A. 2010: Unusual trilobite biofacies from the Lower Ordovician of the Argentine Cordillera Oriental: new insights into olenid palaeoecology. Lethaia, Vol. 44, pp. 58–75. The study of biofacies has proven to be relevant in the understanding of trilobite palaeoecology, palaeobiogeography and macroevolution. The widespread Olenid biofacies is one of the best known, and is usually interpreted as occuring in dysoxic environments. Tremadocian successions of the Argentinian Cordillera Oriental bear a diverse and long‐studied olenid‐dominated fauna. Based on cluster analysis, five distinct biofacies are defined for the middle Tremadocian (Tr2 stage slice), distributed from shelf (below storm wave base) to lower‐shoreface settings (above fair‐weather wave base). Ordination shows biofacies along two gradients, a bathymetrical one and another related to oxygen content. All of them are dominated both taxonomically and ecologically by olenids. This detailed quantitative palaeoecological study challenges current views suggesting instead that the Olenidae dominated a broad range of environments, from oxygenated shallow‐marine to dysoxic deep‐marine. Comparisons with largely coeval trilobite records from geodynamically and palaeobiogeographically disparate sites suggest that siliciclastic sedimentation appears as the most influential controlling environmental factor upon olenid distribution and dominance. Further comparisons across different climatic belts show that siliciclastic input controlled trilobite diversity gradients, even more than latitude. From an autoecological viewpoint distribution of traditional olenid morphotypes shows no relation to depth or to oxygen content, and at least some members of the group appear to have had the possibility of coping with low oxygen content, rather than being restricted to oxygen‐deficient environments. The analysis performed herein, together with recent research on the group, demonstrate that factors controlling olenid distribution are more complex than previously envisaged. □Biofacies, diversity, Olenidae, palaeoecology, Tremadocian, trilobite.     

Techniques of trilobite exuviation

Examples of the trilobites Toxochasmops extensus (Boeck, 1838), Asaphiscus wheeleri Meek, 1874, Encrinurus mitchelli Foerste, 1888, Ogygopsis klotzi (Rominger, 1887), Paradoxides davidis Salter, 1863 and Oryctocephalus spp. which are interpreted as exuvial configurations, are described. Four specimens of Toxochasmops extensus arc known in which the pygidium rests either directly behind the eephalon, or with only three intervening thoracic segments. It is considered that during exuviation the old pygidium became wedged behind the cephalon. This facilitated its removal. An ecdysial configuration of Asaphiscus wheeleri is described which possesses inverted and partially rotated free cheeks. In addition, part of the thorax of the specimen is wedged obliquely behind the cephalon. This is considered to have aided withdrawal of the trilobite from its old exoskeleton. Two specimens of Encrinurus mitchelli are described which possess free cheeks inverted beneath the cranidium by lateral rotation, in a manner similar to that of A. wheeleri. Three examples of Ogygopsis klotzi are described, one a failed exuvia and two in which the free cheeks were inverted and rotated through 180° with respect to their original position and came to rest beneath the thorax. An identical exuvia of Paradoxides davidis trapezopyge is also described. Two specimens of Oryctocephalus exhibiting two different arrangements of inverted free cheeks are recorded. Possible mechanisms for each of these free cheek inversions are proposed.     

A reconstruction of Telephina bicuspis, a pelagic trilobite from the Middle Ordovician of the Oslo Region, Norway

Well preserved parts of Telephina bicuspis described from dark limestone concretions of Llanvirn age, in the Oslo Region, Norway, exhibit many features associated with a pelagic life style for this trilobite. The free cheek has a huge holochroal eye with perfectly square lenses and a long genal spine which is unique in being directed vertically upwards rather than ventrally. Additional spines include an occipital spine, the macro spine on the rachis of the 6th thoracic segment and paired spines from each of the two rachial rings of the pygidium. These spines may have had a hydrodynamic and stabilizing function. The genal spine could have had a sensory function or been a dimorphic feature. The hypostoma is figured for the first time and a reconstruction is given for the complete exoskeleton.     

Severe traumatic brain injury after the assault with an axe handle

Traumatic brain injuries represent a major cause of death and disability. We present a case of a 47-year-old patient who sustained a severe brain injury after being assaulted with a handle of an axe. The patient underwent numerous surgeries by various specialists during several months. Following a few failed attempts to cover the skull defects, the vacuum-assisted closure system had been utilized with great success in healing of her complex head wound. Traumatic brain injury requires great effort and collaboration in order to rehabilitate people to the most independent level of functioning possible.     

Cope's rule in the Ordovician trilobite family Asaphidae (order Asaphida): patterns across multiple most parsimonious trees

Cope's rule defines lineages that trend towards an increase in body size through geological time. The trilobite family Asaphidae is one of the most diverse of the class Trilobita and ranges from the Upper Cambrian through to the Upper Ordovician. The group is one trilobite clades that displays a large size range and contains several of the largest trilobite species. Reduced major axis correlations between the lengths of cephala and pygidia and the total sagittal length of complete individuals have high support and were used to standardise all incomplete specimens to total axial length. Phylogenetic studies into Cope's rule tend to use supertrees, composite trees or a single tree selected through a fit criterion. Here, for the first time, all trees recovered from a maximum parsimony analysis were analysed equally. Maximum likelihood was used to fit four evolutionary models: random walk, directional, Ornstein–Uhlenbeck (evolution towards an adaptive optimum) and stasis. These were compared equally using Akaike weights. Fitting of evolutionary models by maximum likelihood supports stasis as consistently the most likely model across all trees with low support for directionality.     

Neuroprotective properties of levosimendan in an in vitro model of traumatic brain injury

Background  

We investigated the neuroprotective properties of levosimendan, a novel inodilator, in an in vitro model of traumatic brain injury.     

New mechanics of traumatic brain injury

The prediction and prevention of traumatic brain injury is a very important aspect of preventive medical science. This paper proposes a new coupled loading-rate hypothesis for the traumatic brain injury (TBI), which states that the main cause of the TBI is an external Euclidean jolt, or SE(3)-jolt, an impulsive loading that strikes the head in several coupled degrees-of-freedom simultaneously. To show this, based on the previously defined covariant force law, we formulate the coupled Newton–Euler dynamics of brain’s micro-motions within the cerebrospinal fluid and derive from it the coupled SE(3)-jolt dynamics. The SE(3)-jolt is a cause of the TBI in two forms of brain’s rapid discontinuous deformations: translational dislocations and rotational disclinations. Brain’s dislocations and disclinations, caused by the SE(3)-jolt, are described using the Cosserat multipolar viscoelastic continuum brain model.

Internal structures of trilobite trace fossils indicative of an open surface furrow origin

X-radiography of sections cut from Cruziana and Rusophycus of Cambro-Ordovician age, show that they contain cross-laminated, parallel-laminated, pelleted and homogenous fills. The absence of bioturbation or mottling textures and the undisturbed nature of the primary sedimentary structures does not support a burrow origin for these trilobite-produced trace fossils.An open surface furrow origin is supported both by the presence of internal primary sedimentary structures and by the precise current orientation of many of these types of trace fossils.     

Latest Ordovician brachiopod and trilobite assemblage from Yuhang,northern Zhejiang,East China: a window on Hirnantian deep-water benthos

A moderately diverse brachiopod and trilobite assemblage, the Leangella–Dalmanitina (Songxites) Assemblage, occurs in the upper Yankou Formation (Hirnantian, probably equivalent to the Normalograptus persculptus Biozone) at Shizi Hill, Yuhang, west of Hangzhou, northern Zhejiang, E China. The brachiopods are rare, characterised by minute, thin shells with very small body cavities, preserved in mudstones as moulds. They may have inhabited quiet, deep-water and dysaerobic slope environments with low levels of nutrients, equivalent to Benthic Assemblage 5. Most genera were adapted for life in deep water and either remained there or alternatively migrated into relatively shallower habitats to evade perturbations during the first phase of the end Ordovician extinctions. The slope environments were recolonised from outer shelf and upper slope communities during the early Hirnantian, but isolated biotas may also have survived in deeper-water habitats by reducing their population size and diversity during the crisis. The Leangella–Dalmanitina (Songxites) Assemblage provides an unique Hirnantian window through which we can monitor the changes in the deep-water biofacies following the first phase of the extinctions. Significantly, parts of the deep water marine environment may have survived intact, the end Ordovician extinctions.