Palaeoscolecidan worms and a possible nematode from the Early Ordovician of South China

Several species of palaeoscolecidan worm have been found in the Tonggao Formation (earliest Floian) of Guizhou Province, China. Two new palaeoscolecidan taxa are described (Sanduscolex regularis n. gen. n. sp. and Ashetscolex nuppus n. gen. n. sp.); another palaeoscolecidan (Wronascolex sp.) is left in open nomenclature. This is the youngest known occurrence of Wronascolex. A possible large free-living nematode is described and illustrated, representing the earliest known nematode in the fossil record, and a rare example from a marine setting.     

Cyclopygid Trilobites from the Ordovician Builth-Llandrindod Inlier, Central Wales

Cyclopygids are a minor element in the abundant trilobite fauna of the Ordovician Llanfawr Mudstones Formation ( teretiusculus – gracilis biozones) of the Builth-Llandrindod inlier. The rarity of these trilobites may be due to the accumulation of these sediments close to the shoreward limit of their depth range, and if this is the case, their presence, together with sedimentological evidence, can be used to infer the bathymetric limits within which the Llanfawr Mudstones Formation might have been deposited. Species of Degamella , Microparia ( Microparia ), Microparia ( Heterocyclopyge ), Sagavia and Emmrichops are described; some are common to the Dobrotivá Formation in Bohemia, whilst others have affinities with species from approximately coeval strata in Kazakhstan and north-west China.     

Spicule structure and affinities of the Late Ordovician hexactinellid‐like sponge Cyathophycus loydelli from the Llanfawr Mudstones Lagerstätte,Wales

Exceptionally well‐preserved specimens of the reticulosan sponge Cyathophycus loydelli from the Sandbian (Late Ordovician) Llanfawr Mudstones Formation of Llandrindod, Waes, UK, have been examined using scanning electron microscopy (SEM). The specimens include exquisitely detailed pyritized spicules, and granular pyritization of surrounding soft tissues. Spicules frequently show axial canals of diameter similar to those of modern siliceous sponges, with hexagonal symmetry typical of modern demosponges rather than hexactinellids. In one case, the axial filament is also preserved. The largest spicules (ray diameter >20 μm) show a complex structure, with a laminar external region similar to that of the extant hexactinellid Monorhaphis. Some spicules preserve sub‐micron detail of the spicule surface, resembling the reticulate collagenous sheath of Monorhaphis. The hexagonal symmetry of the canal confirms that at least some Reticulosa are not crown‐group hexactinellids, but stem‐group Hexactinellida or Demospongea, or stem‐group Silicea. This suggests that a square canal is a sufficient diagnostic feature of total‐group Hexactinellida, but that hexagonal canals were more widely distributed among Early Silicea and were probably not restricted to demosponges. Alternatively, comparison with the structure of modern verongiid fibres suggests that these may be homologous with the outer layers of Cyathophycus spicules, and Cyathophycus may instead be a stem‐group demosponge. The preserved detail of the surface layer shows that pyritization can preserve certain material with extraordinarily fine resolution.     

Unravelling causal components of the Ordovician Radiation: the Builth Inlier (central Wales) as a case study

Hypotheses about the causes of biodiversification during the Ordovician have been focused in three main areas: tectonic activity and nutrient supply, palaeogeography, and ecological escalation. There is as yet no consensus on mechanisms, and it is unclear whether it is better to study the patterns at local or regional scales. By applying ecological knowledge to the available palaeontological information, it can be shown that neither tectonic nor palaeogeographic effects could account for the permanence of the diversity rise, in the absence of elements of ecological escalation. However, it may be possible to identify trigger mechanisms resulting in enhanced speciation or reduced extinction. Areas of local diversity increase should be distinguished from speciation centres. An ongoing study of the Middle Ordovician Builth-Llandrindod Inlier of central Wales, conducted over 10 years, has identified elements of all three of the above categories of causal mechanisms affecting local diversity. This implies that the patterns of causal relationship and diversification are complex even at very local scales, and at this stage we should not anticipate a clear correlation of global diversity with any single factor. More data are needed from small-scale but intensive studies before we can generalize about the causal mechanisms of the Ordovician Radiation.     

A probable hyalonematid sponge (Hexactinellida: Amphidiscophora) from the Middle Ordovician of the Builth Inlier,Wales

Modern hexactinellid sponges are diverse, but almost exclusively deep-marine organisms with a very intermittent fossil record. Aside from the fused skeletons of hexactinosan lineages (which are also exceptionally rare in Palaeozoic sediments), identifying other families is challenging due to the microscopic nature of many diagnostic characters, and the need for exceptional preservation in a deep-water palaeoenvironment. Among the more distinctive living families is the Hyalonematidae, which have several preservable diagnostic features. A new sponge (Nectocollare zakdouli n. gen. n. sp.) from the Middle Ordovician Castle Bank fauna of Wales, UK, shows several of these characters, including pinular pentactine dermalia, unbundled choanosomal diactins, and a reticulate marginal rim at the apex. No root tuft is preserved, but these are often detached from fossil sponges. Although there remains some uncertainty over the assignment to Hyalonematidae, this sponge represents the only probable example from the Palaeozoic. Even in the absence of diagnostic microscleres, it likely represents a derived, crown-group hexactinellid, and further confirms the diversification of the class during the earliest Palaeozoic.     

Trace fossil evidence of predation upon bone‐eating worms on a baleen whale skeleton from the Oligocene of New Zealand

The osteophagous worm Osedax (Annelida: Siboglinidae) colonizes vertebrate bones in deep‐sea environments globally. Osedax bioerosion of modern bones suggests a potentially destructive agent in the marine vertebrate fossil record, but the dearth of published reports of abundant Osedax traces suggests an uncertain taphonomic influence of this organism. This study reports Osedax traces (Osspecus boreholes, pockmarks and collapsed galleries) in an Oligocene baleen whale (Cetacea: Eomysticetidae) from New Zealand, which constitute the first record of fossil Osedax traces from the southern hemisphere. Some Osedax traces are cross‐cut by linear biogenic scrape marks, implying that sharks or bony fish fed upon Osedax worms, a process which compounds or potentially accelerates worm‐inflicted damage to vertebrate bones in marine environments.     

Diverse bryophyte mesofossils from the Triassic of Antarctica

Compared with the fossil record of vascular plants, bryophyte fossils are rare; this circumstance is probably related to a lower preservation potential compared with that of vascular plants. We searched for bryophyte remains in extensive collections of plant‐fossil assemblages from the Triassic of Antarctica and identified three assemblages with surprisingly well‐preserved bryophyte fossils. Although most bryophyte remains are too fragmented to conclusively place them in a detailed systematic context, they exhibit features sufficient to suggest the presence of at least four types of leafy bryophytes and two orders of thallose liverworts (Pallaviciniales and Metzgeriales) in the high‐latitude Triassic ecosystems of Antarctica. The leafy bryophytes exhibit combinations of morphological features (e.g. keeled and entire‐margined, ecostate leaves with elongated cells) that today occur in only a few small, systematically isolated groups, but were common among Palaeozoic and especially Mesozoic bryophytes. The diverse morphologies of the bryophyte fossils add further support to previous hypotheses that during warmer periods in the Earth's history, bryophyte vegetation may have been particularly rich and diverse in high‐latitude regions. Through analysis of the sedimentology and taphonomy of these assemblages, we identify a combination of key factors that may explain the preservation of bryophyte fossils in these deposits: (1) punctuated, high‐energetic sedimentary events causing traumatic removal and incorporation of bryophytes into sediment‐laden flood waters; (2) limited transport distance, and short period of suspension, followed by rapid settling and burial as a result of a rapidly decelerating flow discharging into a floodplain environment; and (3) early‐diagenetic cementation with iron hydroxides in locally anoxic zones of the organic‐rich, muddy substrate.     

Spanning the gap: experimental determination of paratenic host specificity of horsehair worms (Nematomorpha: Gordiida)

Abstract. Details of the life cycle of freshwater nematomorphs (gordiids) remain unclear. Free-living aquatic larval gordiids must make a critical transition from an epibenthic aquatic environment to terrestrial hosts. In order to identify potential hosts capable of bridging this ecological gap, the specificity of paratenic hosts of three common species of North American gordiids was investigated. All three species were characterized by an identical infection pattern: low host specificity. Gordiids were able to encyst within annelids, mollusks, crustaceans, insects and a vertebrate. Three species of putative host (a turbellarian, a water mite, and a mosquito larva) were not infected with any of the gordiid species. Internal defense reactions (IDR) and feeding behaviors are implicated as preventing infection in these species. Several of the other host species produced either an IDR or an immune reaction to the cysts, although reactions to the cysts were highly variable between species. In most species, IDR did not cause noticeable harm to the encysted larvae. It is proposed that although many species are easily infected with gordiid cysts, most do not act as natural paratenic hosts. For some of these host groups, especially snails, a role as reservoir host is suggested. Of all hosts included in this study, aquatic insects were identified as the hosts likely responsible for spanning the ecological gap and acting as true hosts for gordiids.     

The disparity of priapulid, archaeopriapulid and palaeoscolecid worms in the light of new data

Priapulids and their extinct relatives, the archaeopriapulids and palaeoscolecids, are vermiform, carnivorous ecdysozoans with an armoured, extensible proboscis. These worms were an important component of marine communities during the Palaeozoic, but were especially abundant and diverse in the Cambrian. Today, they comprise just seven genera in four families. Priapulids were among the first groups used to test hypotheses concerning the morphological disparity of Cambrian fossils relative to the extant fauna. A previous study sampled at the generic level, concluding that Cambrian genera embodied marginally less morphological diversity than their extant counterparts. Here, we sample predominantly at the species level and include numerous fossils and some extant forms described in the last fifteen years. Empirical morphospaces for priapulids, archaeopriapulids and palaeoscolecids are relatively insensitive to changes in the taxon or character sample: their overall form has altered little, despite the markedly improved sampling. Cambrian and post-Cambrian genera occupy adjacent rather than broadly overlapping regions of these spaces, and Cambrian species still show lower morphological disparity than their post-Cambrian counterparts. Crucially, the significance of this difference has increased with improved taxon sampling over research time. In contrast with empirical morphospaces, the phylogeny of priapulids, archaeopriapulids and palaeoscolecids derived from morphological characters is extremely sensitive to details of taxon sampling and the manner in which characters are weighted. However, the extant Priapulidae and Halicryptidae invariably resolve as sister families, with this entire clade subsequently being sister group to the Maccabeidae. In our most inclusive trees, the extant Tubiluchidae are separated from these other living taxa by a number of small, intervening fossil clades.     

Larval horsehair worms (Nematomorpha) from the tissues of native freshwater fish in New Zealand

Abstract

Encysted larval horsehair worms (Phylum Nematomorpha) were recovered from the tissues of upland bullies (Gobiomorphus breviceps) and a galaxiid (Galaxias vulgaris) from the Seaward River, Canterbury. The cysts were mostly in the alimentary submucosa. A few occurred in the liver. An adult digenean fluke (Coitocaecum sp.), parasitic within the gut of a bully, also contained nematomorph larvae in its tissues. It is suggested that the fish became infected by eating nematomorph larvae or egg strings.     

A new aglaspidid arthropod from the Lower Ordovician of Wales

A new genus and species of aglaspidid arthropod, Tremaglaspis unite , is described from the well-known upper Tremadoc locality of Y Garth in North Wales, and compared with other Cambrian and Ordovician taxa. It is remarkable in having a large hypostome, a character shared with other primitive arachnomorphs.     

Bryozoans as taphonomic engineers,with examples from the Upper Ordovician (Katian) of Midwestern North America

A combination of encrusting calcitic bryozoans and early seafloor dissolution of aragonitic shells recorded in the Cincinnatian Series of the upper Midwest of North America allowed the preservation of abundant moulds of mollusc fossils bioimmured beneath the attachment surfaces of the bryozoans. We here call this preservational process ‘bryoimmuration’, defined as a bryozoan‐mediated subset of bioimmuration. The bryozoans moulded very fine details of the mollusc shells, usually with more accuracy than inorganic sediment moulds. Most of the bryozoans are heterotrypid trepostomes with robust low‐Mg calcite skeletons. The molluscs are primarily bivalves, gastropods, nautiloids and monoplacophorans with their originally aragonitic shells now dissolved. Many of the encrusting bryozoans are so thin and broad that they give the illusion of calcitic mollusc shells clinging to the moulds. Some molluscs in the Cincinnatian, especially monoplacophorans and epifaunal bivalves, would be poorly known if they had not been bryoimmured. Unlike internal and external moulds in sediment, bryoimmured fossils could be transported and thus record aragonitic faunas in taphonomic assemblages (e.g. storm beds) in which they would otherwise be rare or absent. In addition, bryoimmurations of aragonitic shells often reveal the ecological succession of encrustation on the shells by exposing the earliest encrusters and borings that were later overgrown. Bryoimmuration was common during the Late Ordovician because the calcite sea at the time quickly dissolved aragonitic shells on the seafloor before final burial, and large calcitic bryozoans very commonly used molluscs as substrates. Bryoimmuration is an important taphonomic process for preserving aragonitic faunas, and it reveals critical information about sclerobiont palaeoecology. Several Cincinnatian mollusc holotypes are bryoimmured specimens. Bryozoans involved in bryoimmuration enhance the preservation of aragonitic fauna and thus act as taphonomic engineers.     

Scontrone (central Italy), signs of a 9‐million‐year‐old tragedy

The fossilierous bonebeds of Scontrone (Abruzzo region, central Italy) are preserved in tidal‐flat aeolian calcarenites at the base of the Lithothamnion Limestone, a Miocene carbonate ramp widespread in the central‐southern Apennines. The site bears evidence of a catastrophic event at 9 Ma. Reported are the results of the palaeobiological and taphonomic analysis conducted on the rich vertebrate assemblage, particularly on the remains of Hoplitomeryx (Mammalia, Artiodactyla, Ruminantia), recovered from the so‐called Scontrone calcarenites between 1992 and 2012. This is the first taphonomic study of a Late Miocene continental bone assemblage preserved in coastal deposits. The bones are not in primary context. They were likely exhumed during the initial phase of a marine transgression after a period of primary ‘storage’ within a possibly flood‐generated deposit in an estuarine environment. The mortality patterns indicate that the carcasses accumulated in a short time (within a year). The bones of the disarticulated skeletons were then removed, broken in a dry and brittle state, scattered over wide carbonate ramps along an arid to semi‐arid, wind‐exposed coastline and eventually buried again in aeolian calcarenites that drape transgressive tidal‐flat creek deposits. The analysis also reveals that hoplitomerycids were possibly seasonal reproducers and that the land they inhabited, the so‐called Apulia Platform, was probably swept by sudden, disastrous, storm‐supplied flash floods.     

Disarticulation patterns in Ordovician crinoids: Implications for the evolutionary history of connective tissue in the Crinoidea

Taphonomic information is examined to evaluate the early history of connective tissues in the Crinoidea. The pattern of stalk segmentation of Middle and Late Ordovician crinoids is consistent with the two-ligament (intercolumnal and through-going ligaments) pattern present in living isocrinid crinoids and interpreted for fossil isocrinids, holocrinids, and Lower Mississippian crinoids. A single rhombiferan was also examined; its taphonomic pattern is also indicative of this style of tissue organization. Furthermore, the taphonomy of all Middle and Late Ordovician crinoids may reflect that they lacked discretely organized muscles between arm brachials, which is consistent with the hypothesis that muscles evolved as a connective tissue between plates only once within the Crinoidea, during the Early Devonian. These data indicate that the two-ligament organization of the stalk is a primitive feature among the Crinoidea and perhaps even among stalked echinoderms. Therefore, the autotomy function of this column-tissue organization among living crinoids is an exaptation. On the other hand, discretely organized muscles as connective tissue in crinoid arms is a derived trait that first appeared during the middle Paleozoic; this adaptation proved very successful for the advanced cladid crinoids.     

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

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & 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.     

Chordodes mizoramensis (Nematomorpha, Gordiida), a new species of horsehair worm from Mizoram, North-East India

Chordodes mizoramensis, a new species of freshwater gordiid horsehair worm, is described from Mizoram, NE India on the basis of scanning electron microscopic and morphometric studies. The new species can be distinguished from its congeners in that the apical filaments of the crowned areoles are branched several times, a pattern that has not been observed in other species. An additional distinguishing character is that it has more bulging areoles, which are distributed among simple areoles alone or in groups, do not form clear patterns.     

A cephalopod fauna from the middle Ordovician of Saudi Arabia

A small collection of cephalopods from the middle Ordovician of the Tabuk region of Saudi Arabia consists entirely of orthoconic longicones tentatively referred to the actinocerid genus Mesaktoceras? and pseudorthocerid Sactoceras?. Both taxa have Australasian affinities. Circumpolar Gondwana cephalopod faunas in the mid Ordovician were of low diversity and dominated by orthocerids, whilst actinocerids were largely restricted to low palaeolatitudes. The presence of Mesaktoceras? and Sactoceras? in a benthonic fauna belonging to the Neseuretus community is regarded as anomalous. Comparison is made with the occurrence of Trocholites in middle Ordovician Iberian successions. It has been suggested that Trocholites may have arrived in the Iberian area from Baltica through the transfer of bodies of warm water in storm masses. Mesaktoceras? and Sactoceras? may have arrived (albeit temporarily) in Saudi Arabia from Australasia through a similar process.     

Chondrichthyan‐like scales from the Middle Ordovician of Australia

Abstract: Microvertebrate sampling of the Stairway Sandstone (Darriwilian, Middle Ordovician, central Australia) has yielded scales that are chondrichthyan‐like in their overall construction, and Tantalepis gatehousei gen. et sp. nov. is erected here to describe these specimens. Tantalepis gatehousei gen. et sp. nov. is the stratigraphically oldest microsquamous taxon described thus far, and the ‘shark‐like’ appearance of the scales may extend the chondrichthyan lineage back into the Middle Ordovician. The presence of ‘shark‐like’ scales in the fossil record some 50 myr prior to the first articulated chondrichthyan body fossils and 44 myr before the first clearly identifiable chondrichthyan teeth suggests there is a considerable scope for the recovery of articulated specimens with which to document the early history of crown gnathostomes. Traditional hypotheses of phylogenetic relationships among early jawed vertebrates were recently challenged by the proposal of a radically different tree topology. However, the development of a new data set specifically addressing scale‐based characters is required before taxa such as Tantalepis, that are based upon disarticulated remains alone, can be firmly placed within the emerging, revised, evolutionary narrative.     

A well‐preserved isopod from the Middle Jurassic of southern Germany and implications for the isopod fossil record

A new isopod species, Eonatatolana geisingensis gen. et sp. nov., is described from Middle Jurassic shallow‐water sediments of southern Germany. It shows not only the almost completely preserved dorsal morphology but, in addition, details of the cephalic appendages, the pereiopods, pleopods and uropods. The presence of ambulatory pereiopods I–VII of a wide tridentate mandibular incisor with prominently developed posteriormost tooth and a narrow frontal lamina indicates that the new species belongs to the subfamily Conilerinae of family Cirolanidae within the suborder Cymothoida. It is closer to the species of the modern genus NatatolanaBruce than to any fossil isopod hitherto described. The isopod fossil record as well as current practices of isopod taxonomy in palaeontology are discussed, and the facies distribution and fossilization of isopods is reviewed with examples from the Jurassic.