Showing papers in "Lethaia in 2004"

Geologic Time Scale 2004 – why, how, and where next!

Abstract: A Geologic Time Scale (GTS2004) is presented that integrates currently available stratigraphic and geochronologic information. The construction of Geologic Time Scale 2004 (GTS2004) incorporated different techniques depending on the data available within each interval. Construction involved a large number of specialists, including contributions by past and present subcommissions officers of the International Commission on Stratigraphy (ICS), geochemists working with radiogenic and stable isotopes, stratigraphers using diverse tools from traditional fossils to astronomical cycles to database programming, and geomathematicians. Anticipated advances during the next four years include formalization of all Phanerozoic stage boundaries, orbital tuning extended into the Cretaceous, standardization of radiometric dating methods and resolving poorly dated intervals, detailed integrated stratigraphy for all periods, and on-line stratigraphic databases and tools. The geochronological science community and the International Commission on Stratigraphy are focusing on these issues. The next version of the Geologic Time Scale is planned for 2008, concurrent with the planned completion of boundary-stratotype (GSSP) definitions for all international stages.

A new 'great-appendage' arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages

Abstract: The uniramous ‘great appendages’ of several arthropods from the Early to Middle Cambrian are a characteristic pair of pre-oral limbs, which served for prey capture. It has been assumed that the morphological differences between the ‘great-appendage’ arthropods indicate that raptorial antero-ventral and anteriorly pointing appendages evolved more than once in arthropod phylogeny. One set of Cambrian ‘great-appendage’ arthropods has, however, very similar short antero-ventral appendages with a peduncle of two segments angled against each other (elbowed) and with stout distally or medio-distally directed spines or long flexible flagellate spines on each of the four distal segments. Moreover, the head appendages of all these forms comprise the ‘great appendages’ and three pairs of biramous limbs. To this set of taxa we can add a new form from the Lower Cambrian Maotianshan Shale of southern China, Haikoucaris ercaiensis n. gen. and n. sp. It is known from three specimens, possibly being little abundant in the faunal community. It can be distinguished from all other taxa by the prominence of the proximal claw segment of its ‘great appendages’ and by only three distal spines (one on each of the distal segments). The similarity of the short, spiky ‘great appendages’ of Haikoucaris with the chelicera of the Chelicerata leads us to hypothesize that this particular type of ‘great appendages’ was the actual precursor of the chelicera. Homeobox gene and developmental data recently demonstrated the homology between the antenna of ateloceratans and the antennula of crustaceans on one side and the chelicera of chelicerates on the other. To this we add palaeontological evidence for the homology between the chelicerae of chelicerates and the ‘short great appendages’ of certain Cambrian arthropods, which leads us to hypothesize that the evolutionary path went from the ‘short great appendages’, by progressive compaction, toward the chelicera with only a two-spined chela. The new form from China is regarded as the possible latest offshoot, whereas the other ‘great appendages’ arthropods with similar short grasping limbs were derivatives of the stem lineage of the crown-group Chelicerata. Consequently, the chelicera with a chela with one fixed and one mobile finger is an autapomorphy of the crown group of Chelicerata, whereas a raptorial, but more limb-like antenna, with more distal spine-bearing segments, characterized the ground pattern of Chelicerata. Further taxa having ‘great appendages’, including the large Anomalocarididae, are also discussed in the light of their possible affinities to the Chelicerata and possible monophyly of all of these arthropods with raptorial anterior appendages.

Global Standard Stratotype‐section and Point (GSSP) of the Furongian Series and Paibian Stage (Cambrian)

Abstract: The Global Standard Stratotype-section and Point (GSSP) of the Furongian Series (uppermost series of the Cambrian System) and the Paibian Stage (lowermost stage of the Furongian Series), has been recently defined and ratified by the International Union of Geological Sciences (IUGS). The boundary stratotype is 369 metres above the base of the Huaqiao Formation in the Paibi section, northwestern Hunan Province, China. This point coincides with the first appearance of the cosmopolitan agnostoid trilobite Glyptagnostus reticulatus, and occurs near the base of a large positive carbon isotopic excursion (SPICE excursion).

Status of Divisions of the International Geologic Time Scale

Abstract: Each chronostratigraphic unit of the International Geologic Time Scale will be defined at its base by a Global Stratotype Section and Point (GSSP) or Global Standard Stratigraphic Age (GSSA). Nearly 50 GSSPs and 10 GSSAs have now been ratified. Ideally, the GSSP coincides with events having a global correlation potential. The international stage divisions of some systems, such as the Jurassic or Neogene, are similar to traditional usage in European geology. However, in order to utilize global correlation horizons, the international stage divisions of other systems, such as the Ordovician or Permian, have required assembling new stage nomenclatures or hybrids of different regional stages. A reference table by the International Commission on Stratigraphy itemizes the current or potential GSSP and GSSA definitions of all international geologic time units.

Calcite precipitation and dissolution of biogenic aragonite in shallow Ordovician calcite seas

Abstract: The Ordovician was a time of extensive and pervasive low-magnesium calcite (LMC) precipitation on shallow marine sea floors. The evidence comes from field study (extensive hardgrounds and other early cementation fabrics in shallow-water carbonate sequences) and petrography (large volumes of marine calcite cement in grainstones). Contemporaneous sea-floor events, particularly relationships with boring and encrusting organisms and reworking in sequences of intraformational conglomerates, confirm the early timing of such LMC cementation, and also of widespread associated aragonite dissolution. Local evidence points to the dissolved aragonite as a significant source of the calcite cement. This scenario, and the fabrics that provide the evidence for it, are likely to be pointers to other times in the stratigraphic record when LMC was the predominant shallow marine precipitate (Calcite Sea times). The combination of rapid calcite precipitation and aragonite dissolution at a time early in the Phanerozoic when many major invertebrate groups were becoming established may have acted as an influence on the evolution of both their skeletal mineralogy and their ecology.

The two Early Toarcian (Early Jurassic) extinction events in ammonoids

Abstract: The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass-extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass-extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass-extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long-ranging ammonoids) and Polyplectus (relatively long-ranging compared to other Ammonitina).

A new high‐resolution planktic foraminiferal zonation and subzonation for the lower Danian

Abstract: A new planktic foraminiferal zonation has been established for the lower Danian, based on some of most expanded and continuous pelagic sections known to date (from Spain, Tunisia and Mexico). This biozonation is considered valid for low and middle latitudes. The maximum stratigraphical distribution of the index-species approximately coincides in all the studied sections. The index-species are abundant and easily recognizable. We propose the following biozones and subzones: Guembelitria cretacea Zone and the Hedbergella holmdelensis and Parvularugoglobigerina longiapertura subzones; the Parvularugoglobigerina eugubina Zone, which is subdivided into the Parvularugoglobigerina sabina and Eoglobigerina simplicissima subzones and the Parasubbotina pseudobulloides Zone with the Eoglobigerina trivialis and Subbotina triloculinoides subzones. A biomagnetostratigraphic correlation and calibration of the stratigraphical ranges of these species suggest that the biohorizons used to define the new biozonation are very isochronous, at least in the geographical areas analysed.

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Abstract: Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstatten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Lou & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.

Vertical sections through dinosaur tracks (Late Triassic lake deposits, East Greenland) – undertracks and other subsurface deformation structures revealed

Abstract: Tracks and trackways of theropod dinosaurs (Grallator footprints) are abundant in the Late Triassic lake sediments of East Greenland. For this study we selected a rather diffuse theropod track preserved on the upper surface of a red heterolithic mudrock, and a better preserved track seen on the upper surface of a greyish mudrock. In order to examine undertracks and other subsurface deformation structures, both slabs were sectioned vertically at closely-spaced intervals, perpendicular to the length of the axis of the impression of digit III. Each section was subsequently polished and internal structures revealed. The digit impressions of both tracks were associated with well-defined undertracks which were cut by deep and narrow claw imprints at the distal end of the digit impressions. Marginal ridges at the tracking surfaces were typically associated with subsurface marginal folds. The marginal ridges were asymmetrically developed suggesting an outward movement of the proximal part of the foot, probably during the kick-off; this is in contrast to what is observed in tracks from Lower Jurassic theropods. The study shows that cross-sections through dinosaur tracks display large structural variation and it is suggested that some disturbed layers in continental deposits could be the result of trampling by vertebrates.

A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale

Abstract: Wiwaxia corrugata and the indisputable polychaetes of the Middle Cambrian Burgess Shale, particularly Canadia spinosa, have figured prominently in recent hypotheses about the early evolution of polychaete annelids. Based on similarities between the sclerites of Wiwaxia and the notochaetae of Canadia with the broad notochaetae (paleae) of Recent chrysopetalid polychaetes, these two fossil taxa have been variously treated as closely related to the most highly derived stem forms of the polychaete (and annelid) crown group or as members of a specific, Recent subgroup within Polychaeta, the order Phyllodocida. Chrysopetalidae is a member of Phyllodocida, which is part of the major polychaete clade Aciculata; the latter two taxa are distinguished by four and six well defined autapomorphic characters, respectively. The best preserved or otherwise appropriate fossils of Wiwaxia corrugata, Canadia spinosa and the other polychaetes of the Burgess Shale have been studied in detail in order to determine whether they possess any characters that could support the homology of wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae. Most of these fossil taxa have significant autapomorphies but the specific characters of the Aciculata and Phyllodocida are entirely absent. It is demonstrated that constraining cladograms in such a manner that wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae are homologous leads to results that are markedly unparsimonious. Furthermore, Canadia and the other polychaetes of the Burgess Shale cannot be referred to any extant subgroup within the Polychaeta and cannot be used to polarize character evolution within the annelid crown group. Apart from its dubious sclerites, Wiwaxia has no characters that could indicate any close relationship with Polychaeta or Annelida.

Early Mesozoic evolution of alivincular bivalve ligaments and its implications for the timing of the ‘Mesozoic marine revolution’

Abstract: The early Mesozoic radiation of the Pteriomorphia was accompanied and furthered by the development of several new types of alivincular ligaments. These new types evolved as modifications of the primitive alivincular-areate (new term) ligament, which is characterized by an ontogenetic shift of both the central resilium and the straight lateral ligament in the direction of main shell growth. Arching of the attachment surface of the ligament led to the alivincular-arcuate (new term) ligament type, which has been realized by the Ostreidae only. By contrast, a replacement of the lateral ligament by hinge teeth, limiting the (primary) ligament to a central groove (alivincular-fossate, new term), has evolved independently in three families (Dimyidae, Plicatulidae and Spondylidae). Functionally, both kinds of modification effectively impede shearing of the valves and are interpreted as an antipredatory adaptation advantageous in the cemented habit of these families. The alivincular-alate (new term) ligament of the Entoliidae and Pectinidae differs from the other types of alivincular ligaments by different growth directions of resilium and lateral ligament, which result in an internal position of the resilium suitable for fast and powerful opening of the valves. This arrangement is an important prerequisite for effective swimming, which, in its turn, is a behaviour chiefly used to escape from predator attacks. The simultaneous early Mesozoic appearance of different antipredatory adaptations within independent clades hints at increased predator pressure as a stimulant and may therefore point to a contemporaneous proliferation of durophagous predators. Hence, an important aspect of the ‘Mesozoic marine revolution’ might have started earlier than previously thought.

History, philosophy, and application of the Global Stratotype Section and Point (GSSP)

Abstract: The history, philosophy, and application of the concept of the Global Stratotype Section and Point (GSSP) are reviewed. Geochronologic units defined by GSSPs serve as practical classificatory pigeonholes for the subdivision of geologic time. Accordingly, the main factor involved in the definition of GSSPs must be global correlatability. Early opposition to the GSSP concept centered around the desire for a traditional biochronologic time scale defined conceptually in terms of palaeobiological events, but such time scales are inherently unstable and thus unsuitable for the use of all geoscientists. The GSSP concept is also generally incompatible with the desire for ‘natural’ geochronologic boundaries. GSSPs have been defined mainly on the basis of biostratigraphic guiding criteria, but magnetic polarity reversals and chemostratigraphic and cyclostratigraphic horizons are now playing an important role. Most primary guiding criteria used to place a ‘golden spike’ will eventually become problematical in some way, so GSSPs should be defined so as to be correlatable by as many different lines of age-significant information as possible. The ‘Global Standard Stratigraphic Age’ (better renamed ‘Standard Global Numerical Age’) is a numerical analogue of the golden spike. Numerical definitions are currently appropriate for the formal subdivision of the Precambrian, and perhaps also for the Pleistocene/Holocene boundary. Recent suggestions to abandon chronostratigraphic terms (system, series, stage) in favor of geochronologic terms (period, epoch, age) are logically defensible, but could perpetuate the continuing confusion between various stratigraphic categories.

An ethological framework for animal bioerosion trace fossils upon mineral substrates with proposal of a new class, fixichnia

Abstract: Animal bioerosion trace fossils upon mineral substrates are analyzed from the point of view of the Seilacherian ethological classification. Several of the currently accepted ethological classes: cubichnia, fugichnia, repichnia, fodinichnia, agrichnia, calichnia and aedificichnia are not represented in these substrates. This fact points out a lower behavioral diversity of hard substrate trace fossils when compared with soft sediment trace fossils. Bioerosion traces can be classified in just five classes: domichnia, pascichnia, equilibrichnia, praedichnia and fixichnia. Fixichnia is here erected to gather superficial etching scars resulting from the anchoring of fixation of sessile epiliths by means of a soft or skeletal body part. Praedichnia and fixichnia are exclusive of the bioerosion realm.

The oldest Mesozoic nearshore Zoophycos: evidence from the German Triassic

Abstract: The trace fossil Zoophycos has been described from the Middle Triassic carbonates of the German Basin for the first time. It occurs in a calcilutite bed at the top of a shallowing-upward cycle (parasequence) in the transgressive systems tract of the Middle to Upper Muschelkalk sequence of Thuringia (Germany). Based on sedimentological and palaeontological features, the studied interval is interpreted as deposited in a marine nearshore environment with proximal storm deposits (tempestites). Zoophycos occurs in a very simple planar form with lobate spreiten, which were most likely produced by a worm-like animal by strip mining. The upper tier of the ichnofabric consists of Zoophycos, whereas the lower tier is occupied by cylindrical trace fossils of unknown taxonomic affiliation and with decreasing size towards the bottom. Associated trace fossils such as Rhizocorallium, Balanoglossites and Trypanites indicate a partly firm to hard substrate. No mixed layer is developed at the top of the trace fossil bearing succession. The ichnofabric together with the sedimentological features (disseminated pyrite, blue-grey colour) and palaeontological circumstances (poor benthic fauna, meiofauna with a small body size) support an interpretation of a dysaerobic environment. In the view of evolutionary change, Palaeozoic Zoophycos occurs in both deep and shallow marine deposits, whereas Mesozoic and Cenozoic Zoophycos is only common in shelfal and deeper-marine deposits. The new finding from the shallow-marine Middle Triassic represents the first reliable occurrence of Zoophycos after the end-Permian mass extinction and shows close similarities to its Palaeozoic precursors. It demonstrates that the producer survived the end-Permian mass extinction, became re-established in the nearshore realm and progressively colonized deeper-marine environments during the Mesozoic and Cenozoic.

Extinction process and patterns of Middle Permian Fusulinaceans in southwest China

Abstract: Study of the extinction process of 179 fusulinacean species belonging to 34 genera in southwest China demonstrates that diversity changes in the small (shell length ≤6 mm) and large (shell length >6 mm) species groups were very similar throughout the ‘Maokouan’ (∼Guadalupian). However, significant differences in the timing of extinction pulses occur between different fusulinacean clades with different wall structures, i.e. the nankinellids, schwagerinids, verbeekinids and neoschwagerinds, and between the large and small species groups within the schwagerinids and verbeekinids. Fusulinacean diversity reveals that the Guadalupian mass extinction began in the Middle ‘Maokouan’ and greatly intensified in the late ‘Maokouan’. With only seven species of five genera surviving into the Late Permian, the extinction of fusulinaceans in the Guadalupian mass extinction is 96% at species level and 85% at generic level in southwest China. The preferential extinction of large, morphologically complicated species and the survival of simpler, ecologically more tolerant species of the nankinellids suggest that the extinction of Guadalupian fusulinaceans was caused by falling sea level and the consequent effects related to salinity, temperature and substrate changes. It is also demonstrated that biological characters of fusulinaceans such as shell size and test structures could have certain effects on survivorship of species in the early stages of mass extinction when extinction pressure was less intensive but were ineffectual during the extinction climax.

Experimental fragmentation patterns of modern Nautilus shells and the implications for fossil cephalopod taphonomy

Abstract: Shells of modern Nautilus pompilius from the Philippines were experimentally fragmented designed to mimic: (1) transport with sediment; (2) sediment loading; and (3) collision during floating. The breaking patterns by other mechanisms (predation and implosion by hydrostatic pressure) documented in the literature were also considered. The breaking patterns produced by various mechanisms are distinct and can therefore be differentiated. The results allow identification of the distinct mechanisms responsible for specific fragmentation not only in modern Nautilus but also in fossil cephalopods. This is a new approach for the more complete recognition of post-mortem transport of fossil cephalopods and their early taphonomic history, and contributes to our understanding of their palaeobiogeography and palaeoecology.

Shell structure, ontogeny and affinities of the Lower Cambrian bivalved problematic fossil Mickwitzia muralensis Walcott 1913

Abstract: Exceptionally preserved carbonate- and shale-hosted Mickwitzia muralensis from the Lower Cambrian Mural Formation, southern Canadian Rocky Mountains, complement one another to yield an unusually complete account of its ontogeny, ecology and phylogenetic relationships. The shell of M. muralensis is composed of dense phosphatic layers interspersed with porous organic-rich layers. At the insertion of shell-penetrating tubes, shell layers deflect inwards to produce inwardly pointing cones. The tubes are interpreted as having hosted setae that were secreted by outer-epithelial follicles. Follicular setae also occurred at the mantle margin, where they were oriented within the plane of the shell as in modern brachiopods. During ontogeny, the initial setae oriented in the plane of the shell occurred before the first shell-penetrative setae. In the juvenile and early-mature stages of shell secretion, a posterior opening was present between both valves and was used for the protrusion of an attachment structure. In the late-mature shell, this opening became fixed in the ventral valve. Based on the posterior margin and the shell microstructure, a close relationship between Mickwitzia and the paterinids is proposed with differences interpreted as heterochronic. The shell-penetrative setal apparatus of M. muralensis is distinct from that previously described of Micrina, though both types are conceivably homologous to adult and juvenile setae of modern brachiopods.

Towards a ‘natural’ time scale for the Precambrian – A proposal

Abstract: A proposal is put forward to redefine the geological time scale for the Precambrian. Flaws of the present, chronometrically defined, time scale are discussed and illustrated. It is concluded that we need to go back to the rock record to define a “natural” time scale, in which major divisions (eons, eras, etc.) are defined in terms of first-order events and transitions in the observable stratigraphic record. For the earliest part of Earth history, we need a time scale, including a formalized Hadean eon, that is fully consistent with rapidly evolving insights from planetary science.

The youngest African clymeniids (Ammonoidea, Late Devonian) – failed survivors of the Hangenberg Event

Abstract: A co-occurrence of the clymeniid Postclymenia evoluta Schmidt, 1924 and the goniatite Acutimitoceras hilarum Korn, 2002 is reported from the Anti-Atlas of Morocco. Both species occur in the same limestone horizon within the Acutimitoceras prorsum Zone, that has yielded an exclusive conodont fauna of the Upper praesulcata Zone (latest Devonian, above the Hangenberg Black Shale). This record is firm evidence that some clymeniids survived the global Hangenberg Event, but soon later became extinct without descendants.

Buckman's Paradox: variability and constraints on ammonoid ornament and shell shape

Abstract: Buckman's Law of Covariation states that ammonoid shell shape and ornamentation are typically correlated, such that compressed, involute forms have light ornament while more inflated, evolute forms have heavier ornament. Such covariation has been observed in many ammonoid groups, and implies a link between the morphogenesis of shell shape and ornamentation. However, other evidence suggests that while ornament growth is controlled by the genetic-developmental program of the ammonoid, shell shape is strongly influenced by environmental factors. These differing viewpoints lead to Buckman's Paradox – are ornamentation and shell shape tightly linked, as implied by Buckman's covariation, or is the morphogenesis of ornament controlled genetically, while shell shape is controlled environmentally? To address this issue, the variability of shell shape and rib morphology has been compared for a group of endemic acanthoceratid ammonites from the Cretaceous Western Interior Seaway of North America. If Buckman's Law holds due to a morphogenetic connection between shell shape and ornamentation, we would expect taxa with more variable shell shapes to also show more variable rib features and growth. Morphometric analysis of seven shell shape and two rib characters for the Western Interior acanthoceratids finds no such correlation, suggesting that shell shape and rib growth are controlled by different processes. Indeed, rib growth appears to be more constrained than shell shape, consistent with the view that ornamentation is more tightly controlled by the developmental-genetic growth program of the ammonoid. These results emphasize the complexity of ammonoid morphogenesis and highlight our limited understanding of the causes underlying Buckman's Law.

The lobopodian Paucipodia inermis from the Lower Cambrian Chengjiang fauna, Yunnan, China

Abstract: New specimens of Paucipodia inermis Chen, Zhou & Ramskold, 1995, are described from the Lower Cambrian Chengjiang Lagerstatte in Haikou, Kunming. Details not previously seen in the Chengjiang material appear to be caused by early diagenetic processes. Some features not previously observed in Palaeozoic lobopodians include details of the dermomuscular sac, body cavities, contents of the gut, possible paired ventral nerve ganglia, and a rasping or biting apparatus with teeth. The latter implies a fundamental difference from onychophorans and rules out an ancestral position for Palaeozoic lobopodians. The supposed tail is shown to be the head, and it is shown that this animal possessed nine pairs of lobopods rather than six, as originally stated. The family Paucipodiidae n. fam. is introduced.

Palaeopathology and injury in the extinct mosasaurs (Lepidosauromorpha, Squamata) and implications for modern reptiles

Abstract: Three fossilized dentaries provide an insight into the healing of fractures in a major group of extinct marine predators, mosasaurs. The data has implications for modern day reptiles in which such information is scanty. All three dentaries have callus formation. Both dentaries of Mosasaurus hoffmanni show fracture non-union, possibly resulting from intervening tissue. They also show evidence of osteomyelitis. Bone remodeling is complete in an earlier fracture in one of the M. hoffmanni dentaries. Given that the most recent fracture (non-union) occurred at a new and much deeper part of the dentary and not in the region of the earlier fracture, it can be assumed that remodeled bone in mosasaurs (probably in reptiles generally) could withstand powerful stresses such as those encountered during predation and fighting. The splenial bone, attached in life to the dentary only by connective tissue, may have acted as a natural splint to immobilize the fracture to the dentary and maintain its alignment during healing. Possible explanations for the injuries are feeding on hard-shelled prey, such as turtles, and fighting.

A trace fossil assemblage from fluvial Old Red deposits (Wood Bay Formation; Lower to Middle Devonian) of NW‐Spitsbergen, Svalbard

Abstract: From the fluvial Old Red Sandstone (ORS) of the Lower to Middle Devonian Wood Bay Formation (NW-Spitsbergen), a diverse trace fossil assemblage, including two new ichnotaxa, is described: Svalbardichnus trilobus igen n, isp n is interpreted as the three-lobed resting trace of an early phyllocarid crustacean (Rhinocarididae) Cruziana polaris isp n yields morphological details that point towards a trilobite origin This occurence of presumably marine trace makers in a fluvial red bed sequence raises the question of whether we are dealing with marine ingressions that are not sedimentologically expressed, with homeomorphy, or with an adaptation of marine groups to non-marine environments

An unusual crinoid-barnacle association in the type area of the Danian (Paleocene), Denmark

Abstract: The specialization of holopodid crinoids to a sessile habit, by directly cementing the cup to a hard substrate and having short arms that interlock on closure to give a watertight seal, led F.A. Bather to speculate in 1928 that they might once have led an intertidal, barnacle-like existence in shallow water. Although this remains unproven, holopodid-barnacle associations from the Danian now provide indirect supporting evidence of their similar environmental preferences at that time. It has long been recognized that the holopodid Cyathidium holopus Steenstrup was encrusted by the verrucomorph Verruca prisca ? Bosquet in Danian submarine caves. This crinoid is now known to occur with abundant plates of the brachylepadomorph barnacle Pycnolepas bruennichi Withers in open sea bed settings, suggesting that they lived together on a substrate now lost due to diagenesis. These were most likely to have been azooxanthellate scleractinian corals (benthic; 100+ m estimated water depth) and/or logs (either on sunken benthic islands (100+ m) or floating as pseudoplankton). If the tight, barnacle-like closure of the holopodid arms was primarily an anti-predatory adaptation, suggested as an alternative hypothesis, then it was of limited effectiveness and insufficient to prevent the group having to migrate from shallow into deeper water with the stalked crinoids since the Late Cretaceous.

Early Palaeozoic diversification of chitons (Polyplacophora, Mollusca) based on new data from the Silurian of Gotland, Sweden

Abstract: Isolated, well-preserved silicified sclerites of an unusually diverse Silurian paleoloricate assemblage from Gotland preserve morphological features that are important in interpreting palaeobiology. The typically granular dorsal ornament is comparable with Recent chitons, and is hence possibly linked functionally with sensory aesthetes. Ventral structures particularly in thickened shells indicate major muscle attachment sites sub-apically or, equivalently, beneath the rim of the apical area, and also marginally. In early chitons such as Cambrian Matthevia, deep ventral cavities represent comparable sub-apical sites. Three Gotland genera with an unusual, holoperipheral shell growth style apparently represent plated aplacophorans (cf. Acaenoplax), which coexisted with paleoloricate chitons in shallow inshore carbonate shelf environments. Sclerite features of all the Gotland genera are discussed together since they share most characteristics. The new family Heloplacidae includes those genera and Acaenoplax, which in a preliminary cladistic analysis form a sister group to other Lower Palaeozoic paleoloricates. Multiplated skeletons in paleoloricates and this group of aplacophorans represent parallel evolution of dorsal armour, which in chitons resulted in overlapping, articulating sclerites. The diversity of the mid-Silurian Gotland assemblage is examined against early evolutionary diversification of polyplacophorans, aplacophorans, and in relation to the overall record of Palaeozoic paleoloricate and neoloricate chitons. Peaks in diversity in early Ordovician, mid-Silurian and early Carboniferous times correspond to periods with widespread development of low latitude carbonate shelves. Neoloricates, apparently with an additional shell layer that contributed articulatory plates, appeared in the Devonian from where the fossil record remains poorly known.

Billingen (Lower Arenig/Lower Ordovician) acritarchs from the East European Platform and their palaeobiogeographic significance

Abstract: Billingen (Lower Arenig/Lower Ordovician) sediments of the St. Petersburg region, northwest Russia and the Leba area, northern Poland of the East European Craton yield acritarch assemblages, which are largely homogenous though displaying minor compositional differences that probably reflect a gradient from inner to outer shelf environments. Comparison with coeval acritarch microflora from the Yangtze Platform, South China, shows an overall similarity between Baltoscandian and South Chinese phytoplankton. The widespread uniformity in the fossil microphytoplankton may be related to the extensive global ‘evae’ sea-level transgression, which characterized the Billingen time. This suggests that during the Tremadoc through early Arenig times, acritarch assemblages displayed essentially an undifferentiated cold-water and oceanic character along the whole margin of Perigondwana in the South, as well as on the South Chinese and Baltic platforms, at middle latitudes (Mediterranean oceanic Realm). Despite this overall similarity, however, some typical taxa of the high-latitude Mediterranean Province (Arbusculidium, Coryphidium and Striatotheca) occur in South China, but are absent in Baltica. This discrepancy is explained as caused by differences in climatic and physiographic conditions that prevailed at the two palaeocontinents at this time. The inferred pattern of oceanic circulation during the Lower Ordovician is consistent with the palynological evidence of a prevailing warmer climate in Baltica than in South China, although the two palaeocontinents occupied the same palaeolatitudinal position.

A mineralogical method to determine cyclicity in the taphonomic and diagenetic history of fossilized bones

Abstract: The study of authigenic minerals within the voids of a fossilized bone can reveal its diagenetic history. When the order of precipitation of minerals is plotted on an Eh/pH diagram any cyclicity in the diagenetic history is revealed. If one cycle is displayed then it can be assumed that the bone has been found in its original bed of deposition; if two cycles or more are revealed then reworking or environmental change may have taken place. This is demonstrated in a case study of two bones from the Wealden Group (Lower Cretaceous) from the Isle of Wight, UK.

Sexual dimorphism in Champsosaurus (Diapsida, Choristodera)

Abstract: Two nearly complete specimens of Champsosaurus (Diapsida, Choristodera) with distinctive morphologies, from the Tullock Formation (Early Paleocene) of northeastern Montana, USA, were described as different species. The limb bones of C. ambulator are more robust than those of C. laramiensis, indicating that C. ambulator was more adapted for walking than C. laramiensis. The phylogenetic significance of these limb bone morphologies, however, appears questionable because similar dimorphic variations occur in a closely related genus and champsosaurs from other geologic ages and locations. Female champsosaurs may have been better adapted to a terrestrial life than males due to nesting behavior on land, resulting in variable limb bone morphologies between sexes. The observed morphologic variations are, hence, hypothesized to reflect sexual dimorphism rather than sympatry of species. The C. ambulator -shaped humeri and femora, demonstrating a terrestrial adaptation, are suggested to belong to females and C. laramiensis -shaped limb bones to males. No significant variations of humeral and femoral morphologies occur in small champsosaur specimens, suggesting an aquatic niche for juveniles like adult males.

A paradox survival – report of a repaired syn vivo perforation in a nautiloid phragmocone

Abstract: The nautiloid Trocholites depressus (Eichwald, 1840) from the Lasnamagi regional stage (Darrivillian, Middle Ordovician) of Vaike Pakri Island (North-West Estonia) is the only known ectocochleate cephalopod that survived and healed a perforation of the phragmocone. Two chambers of the specimen were broken during its lifetime. The injury is located on the venter of the conch directly above the peristomal opening of the body chamber. It is reconstructed that the peristomal mantle tissue carried out an initial sealing of the injured chambers. The complete calcified sealing and compensation of the irregular shell surface started late with the overgrowth of the septa of the preceding whorl. The position and diameter of the siphuncle were not disturbed by the regeneration showing that these characters allowed a low phenotypic variability. Despite the trauma in the midlife growth history of this specimen, it appears to have reached maturity.

Morphological adaptations of the late Mississippian problematic alga Calcifolium to fluctuating palaeoecologic environments

Abstract: Well-preserved specimens of Calcifolium ? punctatum Maslov from the Guadiato area (Spain) show a great adaptability to environmental modifications Six morphotypes are recognized: flabellum-shaped cups, massive erect cups, stockade cups, crustose growth, petals and inter-cup segments These morphologies are an adaptative response to fluctuation of the following factors: substratum, light/turbidity/depth, competition for space, influx of micrite/rate of sedimentation, hydrodynamic energy and bioturbation