Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

Implications de données récentes sur les Lingules actuelles dans les interprétations paléoécologiques

Some traditional concepts on lingulid paleoecology are reviewed and discussed in regard to new data on the ecology of Recent lingulid brachiopods. Fossilization in life-position occurs under several ecological conditions, e.g. dessication or emersion of the substratum, prolonged increase in salinity, perhaps very rapid sedimentation, extreme changes in temperature, and fossilization of flat-lying shells occurs under prolonged reduction in salinity, storms, natural death. Catastrophic environmental changes cause the fossilization of lingulids. Such events may affect one or more ecological features which may have a direct or indirect effect on the lingulid biology. At this time, the biotype does not correspond to the habitual one, and the survival and behavior of the lingulids depend on the severity of the environmental changes and on the tolerance of the animals to the latter.     

THE COMPOSITION AND STRUCTURE OF LOWER SILURIAN MARINE COMMUNITIES

Five benthic communities occupied the shelf regions of the British Isles, Norway, and North America in Upper Llandovery times. The communities are listed below in order of increasing distance from shore.

• 1 The Lingula Community is the least diverse; it has both infaunal elements, including a protobranch, and two lingulids, and epifaunal elements, including a rhynchonellid, a pterioid, and a cornulitid. A restricted and protected near-shore environment, such as a bay or estuary, is postulated.
• 2 The Eocoelia Community shares elements in common with the former community, but is more diverse and is dominated by epifaunal forms; the many small pedunculate brachiopods probably lived attached to the large leptostrophiid brachiopod.
• 3 The Pentamerus Community is dominated by this genus which lived free and upright on the bottom; smaller pedunculate brachiopods probably attached to this large neighbor.
• 4 The Costistricklandia Community was similar in structure to the former community with the many small pedunculate brachiopods being attached to the large Costistricklandia.
• 5 The Clorinda Community is the most diverse, with a great variety of small brachiopods which were probably able to attach to small objects in this quiet off-shore environment, or to some moderately sized brachiopods, such as Clorinda and Cyrtia, which apparently lived free on the bottom.

The brachiopod dominated communities of the Silurian clearly inhabited the ‘level bottom’, an area now occupied mainly by infaunal forms. The main attachment surfaces for the epifaunal elements of the Silurian communities were disarticulated, convex-upward shells.     

Protegulum and brephic shell of the earliest organophosphatic brachiopods

Early development stages imprinted on the shells of Cambrian brachiopods from the class Linguliformea (orders Paterinida, Lingulida, Acrotretida) were studied with scanning electron microscope based on a large collection from the Siberian Platform. Some specimens of all three orders preserved protegulum (embryonic shell); their brephic (juvenile) shells, also were studied. Many of them might lack larval development stage as it is known for the recent representatives of the family Lingulidae. But unlike recent lingulids, the surface of the juvenile dorsal valves of all studied linguliformeans bore two (rarely three) pairs of gentle elevations accommodating bundles of setae, which sometime preserve as groups of fine imprints along their margins. Recent Lingula and Glottidia lack setae in this stage. On the other hand, recent classes Craniformea and Rhynchonelliformea have setae, but they develop in the larval stage. Most of brachiopod groups have the setae in the adult stage but these setae have different origin, are short, located on the lateral and anterior ends of the shell, and grow from the marginal cells of the mantle.     

Lingulichnites, a new trace fossil genus for lingulid brachiopod burrows

Holocene lingulid brachiopods are well-known burrowing organisms. Fossil lingulid burrows have been reported from rocks in Europe and North America that range from Ordovician and Devonian through Pennsylvanian. However, the latest authoritative compendium on trace fossils, The Treatise on Invertebrate Paleontology (1975). makes no mention of lingulid burrows. The obvious reason for this omission is that the burrows, although important paleontologically and ichnologically, have never been named formally thereby making it awkward to include them in faunal lists. The monotypic genus Lingulichnites is proposed to remedy this problem. The type specimens are from Upper Devonian rocks in northeastern Ohio.     

Architecture and function of the lophophore in the problematic brachiopod Heliomedusa orienta (Early Cambrian, South China)

The detailed structure of the lophophore is a key diagnostic character in the definition of higher brachiopod taxa. The problematic Heliomedusa orienta Sun and Hou, from the Lower Cambrian Chengjiang Lagerstätte of Yunnan, southwestern China, has a well-preserved lophophore, which is unlike that of any known extant or extinct brachiopods. Based on a comparative study of lophophore disposition in H. orienta and the extant discinid Pelagodiscus atlanticus, the in- and excurrent pattern and shell orientation of H. orienta are described and discussed. Reconstructions of lophophore shape and function are based on numerous specimens and comparison with P. atlanticus. The lophophore is composed of a pair of lophophoral arms that freely arch posteriorly rather than coiling anteriorly as commonly seen in fossil and recent lingulids. The lophophore is attached to the dorsal lobe of the mantle; it has neither calcareous nor chitinous supporting structures, and is disposed symmetrically on either side of the valve midline. The mouth can be inferred to be located at the base of the two brachial tubes, slightly posterior to the anterodorsal projection of the body wall. The lophophoral arms bear laterofrontal tentacles with a double row of cilia along their lateral edge, as in extant lingulid brachiopods. The main brachial axes are also ciliated, which presumably facilitated transport of mucous-bound nutrient particles to the mouth. The unique organization of the lophophore in Heliomedusa is not like any known fossil and living brachiopods. This clearly demonstrates that H. orienta is not a member of any crown group. It is here considered as a member of the brachiopod stem group, which challenges recent interpretations of a close discinid affinity.     

Provenance Of Atlantic Lingulid Brachiopods

Living lingulid brachiopods are ubiquitous in low-latitude, marine infaunas. Lingula occurs throughout the Pacific and Indian oceans with the only Atlantic species, L. parva, confined to West Africa. Glottidia is restricted to offshore America from Virginia to California and Peru, and is assumed to have descended from a Pacific Lingula during the early Tertiary. Lingulid organophosphatic shells differ structurally. That of Glottidia is characterizedby trellised rods (baculate); that of Indo-Pacific species of Lingula by spheroidal and rod-like microstructures (virgose); and that of L. parva by apatitic rods arranged as spherulites. A spherulitic fabric is unknown in fossil lingulids, but the distinction between GlottidiaLingula can be traced back to the Carboniferous, which accords with the deep molecular divergence between the two genera. The common occurrence of lingulids with baculate shells in European post-Palaeozoic sediments suggests that ancestral Glottidia entered the Atlantic by the Tethyan Current during the Late Cretaceous/early Cenozoic, and migrated into the Pacific before the formation of the Panama Isthmus. Penecontemporaneously, antecedents of L. parva possibly migrated from east Tethys along the trans-Saharan seaway.     

Turbulence,displacement, death and worms: a day in the life of a fluvial Carboniferous bivalve

Kane, I.A. 2010: Turbulence, displacement, death and worms: a day in the life of a fluvial Carboniferous bivalve. Lethaia, Vol. 43, pp. 381–395. In the Pennsylvanian Rough Rock Flags and Rough Rock of northern England trace fossils attributed to the non‐marine bivalve Carbonicola are found. Carbonicola, recorded by Lockeia and associated trace fossils, lived a semi‐infaunal lifestyle and thus were influenced by both the sediment in which they were hosted, and the currents which supplied their nutrients and oxygen. A number of palaeocurrent indictors are commonly associated with Lockeia: (1) downstream inclination of vertical burrows; (2) palaeoflow‐parallel orientation of long axes; (3) steeper scouring and higher sediment surface on the upstream side; and (4) diffuse lamination downstream of the trace or more widespread downstream erosion. Semi‐infaunal, Carbonicola bivalves were partly exposed to the prevailing flow and acted as bed defects; flow separation and acceleration enhanced flow turbulence around the bivalve leading to erosion and the development of a variably developed fan‐shaped zone of scour immediately downstream. Disturbance and destabilization of sediment in this way may have affected bivalves immediately downstream, plausibly explaining the relatively regular spacing pattern of individual or clustered Lockeia, exposed on bedding planes and revealed by nearest neighbour analyses. Bivalves that did not survive high‐energy flow events were either trapped within the sediment, or transported downstream and deposited in lower energy environments within the otherwise high‐energy deposits of the Rough Rock.     

Tommotiids from the early Cambrian (Series 2, Stage 3) of Morocco and the evolution of the tannuolinid scleritome and setigerous shell structures in stem group brachiopods

An assemblage of tannuolinid sclerites is described from the Amouslek Formation (Souss Basin) of the Anti‐Atlas Mountains in Morocco. The assemblage contains two species, Tannuolina maroccana n. sp., which is represented by a small number of mitral and sellate sclerites, and Micrina sp., represented by a single mitral sclerite. Tannuolina maroccana differs from other species of the genus in the presence of both bilaterally symmetrical and strongly asymmetrical sellate sclerites. This observation suggests that the scleritome of Tannuolina was more complex than previously thought and that this tommotiid may have held a more basal position in the brachiopod stem group than previously assumed. The shell structure of both T. maroccana and Micrina sp. is well preserved and exhibits two fundamentally different sets of tubular structures, only one of which was likely to contain shell‐penetrating setae. Based on these observations, the structure of the tannuolinid shell is discussed and its implications for the evolution of tubular microstructures in stem and crown group brachiopods are analysed.     

RHYNCHONELLIFORMEAN BRACHIOPODS WITH SOFT-TISSUE PRESERVATION FROM THE EARLY CAMBRIAN CHENGJIANG LAGERSTÄTTE OF SOUTH CHINA

Abstract:  Cosmopolitan kutorginates, the most abundant Early Cambrian rhynchonelliformean brachiopods, became extinct at the end of the Middle Cambrian. Consequently, any information concerning the anatomy of this peculiar lineage of brachiopods has great phylogenetic significance with regard to their extant relatives for analogies with the stem-group clade. Such data have been supplied from fossils of which the soft parts have been preserved in exceptional detail. A new brachiopod, Kutorgina chengjiangensis sp. nov., from the Early Cambrian Chengjiang Lagerstätte of southern China, is described here. It is the first articulated brachiopod species collected from this deposit. The specimens preserve a set of soft-body parts, i.e. lophophore, digestive tract and pedicle, all previously poorly known in almost all Palaeozoic calcareous brachiopod taxa. The lophophore attains an early spirolophe stage, clearly homologous to that in the coeval lingulids. The digestive tract consists of a mouth, oesophagus, swollen stomach, intestine and a terminal anus. The pedicle protruding between the valves is stout and elongate, with annulated lamellae on the surface, and contains a putative cœlomic cavity. K. chengjiangensis displays the characteristics of the stem group of calcareous brachiopods, and illustrates anatomical similarities between Cambrian phosphatic- and calcareous-shelled brachiopods, thus corroborating that the Brachiopoda are a monophyletic group.     

Life mode of in situ Conularia in a Middle Devonian epibole

Abstract: Exceptionally abundant specimens of Conularia aff. desiderata Hall occur in multiple marine obrution deposits, in a single sixth‐order parasequence composed of argillaceous and silty very fine sandstone, in the Otsego Member of the Mount Marion Formation (Middle Devonian, Givetian) in eastern New York State, USA. Associated fossils consist mostly of rhynchonelliform brachiopods but also include bivalve molluscs, orthoconic nautiloids, linguliform brachiopods and gastropods. Many of the brachiopods, bivalve molluscs and conulariids have been buried in situ. Conulariids buried in situ are oriented with their aperture facing obliquely upward and with their long axis inclined at up to 87 degree to bedding. Most specimens are solitary, but some occur in V‐like pairs or in radial clusters consisting of three specimens, with the component specimens being about equally long or (less frequently) substantially different in length. The compacted apical end of Conularia buried in situ generally rests upon argillaceous sandstone. With one possible exception, none of the examined specimens terminates in a schott (apical wall), and internal schotts appear to be absent. The apical ends of specimens in V‐like pairs and radial clusters show no direct evidence of interconnection of their periderms. The apical, middle or apertural region of some inclined specimens abuts or is in close lateral proximity to a recumbent conulariid or to one or more spiriferid brachiopods, some of which have been buried in their original life orientation. The azimuthal bearings of Conularia and nautiloid long axes and the directions in which conulariids open are nonrandom, with conulariids being preferentially aligned between 350 and 50 degree and with their apertural end facing north‐east, and nautiloids being preferentially aligned between 30 and 70 degree. Otsego Member Conularia were erect or semi‐erect, epifaunal or partially infaunal animals, the apical end of which rested upon very fine bottom sediment. The origin of V‐like pairs and radial clusters remains enigmatic, but it is probable that production of schotts was not a regular feature of this animal’s life history. Finally, conulariids and associated fauna were occasionally smothered by distal storm deposits, under the influence of relatively weak bottom currents.     

Unusual sensory setae of the raptorial Branchinecta gigas (Branchiopoda: Anostraca)

In many vernal pools, visibility is very poor because of the turbidity from suspended clay particles. For predatory species like Branchinecta gigas, these conditions can be detrimental to successful prey capture. In vernal lakes in central California, B. gigashave developed specialized hunting modes to capture anostracan prey in pools of low visibility. The position of their body, the kinematics of their locomotion, and their reduced eye size suggested the possibility of novel sensory structures on their antennae and/or their cercopods designed to enhance their prey capture abilities. Using Scanning Electron Microscopy, we investigated the presence and design of sensory setae on the antennae and cercopods of B. gigas. On both males and females, there are dense patches of sensilla along the length of the antennae. They are oriented ventrally and slightly anteriorly. These antennal setae appear to be chemosensory in structure and position; they resemble antennal setae of other branchiopods. However, the setae of the cercopods are unusual in their morphology and location on the appendage. The cercopods, which are bent over the head in the hunting position, have a linear arrangement of specialized setae on their ventral side. They are jointed setae with an anterior crown of protective spines. The setal joint only permits limited abduction either toward the head in the hunting position or ventrally when swimming. These setae appear to be mechanosensory in function and may be adaptations to a raptorial lifestyle. They correlate well with the behavioral components of hunting in B. gigasand their complex prey capture mechanism.     

Symbiotic interactions in the Silurian of North America

There are at least 47 different symbiotic pairs of taxa and 16 symbiotic associations in the Silurian of North America. Crinoids are most common host species and they hosted variety of epibiotic and endobiotic symbionts, including Tremichnus, platyceratid gastropods, brachiopods, microconchids, cornulitids, cyclostome bryozoans and favositid tabulates. Eighteen symbiotic pairs contain at least one colonial partner. Stromatoporoids hosted the most diverse fauna of endobiotic symbionts, including cornulitids, lingulids, Chaetosalpinx, Heliocosalpinx and rugosans. Among 16 symbiotic associations of Silurian of North America, 8 are common between North America and Baltica. North American symbiotic associations involving stromatoporoid hosts are the most similar to their Baltic equivalents.     

EXTINCT TRANSITIONAL FAGACEAE FROM THE OLIGOCENE AND THEIR PHYLOGENETIC IMPLICATIONS

Fruits, catkins, and associated leaves of at least two extinct trigonobalanoid taxa have been discovered at an Oligocene fossil plant locality rich in fagaceous remains. These fossils exhibit a mosaic of fruit and pollen characters found in the two extant subfamilies Castaneoideae and Fagoideae of Fagaceae. Comparison with cladograms based on modern taxa suggests that these extinct taxa were similar to the ancestors of subfamily Fagoideae and may have been intermediate between Fagus and the modern trigonobalanoid genera. Pollen types isolated from the fossil staminate catkins provide unique character states that are transitional between modern pollen types in Fagaceae and are important in understanding the evolution of exine micromorphology within the family. This analysis provides a striking example of the use of character data from fossils to determine character-state adjacency prior to polarization of characters using outgroup comparison. Because of the mosaic nature of their character complexes, these fossils support monophyly in both the family Fagaceae and the subfamily Fagoideae. In addition, the occurrence of trigonobalanoid fossils in the Oligocene of North America has interesting biogeographic implications and provides insights into the nature of North American Fagaceae during the Tertiary.     

Brachiopod morphology,life mode,and crushing predation in the Ordovician Arnheim Formation,Kentucky, USA

Predation on ancient shelled prey is an often-studied topic in paleoecology, but the early Paleozoic and the brachiopods that dominated the seafloor at that time are relatively underrepresented in the predation literature. We assessed predatory repair scar frequencies among the brachiopod genera from the Early Richmondian (Late Ordovician) Oregonia Member (Arnheim Formation) near Flemingsburg, Kentucky. We found higher repair frequencies on the concavo-convex Rafinesquina and Leptaena relative to the bi-convex genera. There were no trends in repair frequency through the stratigraphic section and no relationships between repair frequency and community diversity metrics. It is possible that concavo-convex brachiopods’ flat shape, thin shell profile, and free-lying (no pedicle attachment) lifestyle made them more likely or appealing targets of Ordovician crushing predators. It is also possible that concavo-convex brachiopods were better suited to survive crushing attacks than biconvex taxa. We also found differences in shell ornament that may influence the visibility of repair scars.     

Dynamic palaeoredox and exceptional preservation in the Cambrian Spence Shale of Utah

Garson, D.E., Gaines, R.R., Droser, M.L., Liddell, W.D. & Sappenfield, A. 2011: Dynamic palaeoredox and exceptional preservation in the Cambrian Spence Shale of Utah. Lethaia, Vol. 45, pp. 164–177. Burgess Shale‐type faunas provide a unique glimpse into the diversification of metazoan life during the Cambrian. Although anoxia has long been thought to be a pre‐requisite for this particular type of soft‐bodied preservation, the palaeoenvironmental conditions that regulated extraordinary preservation have not been fully constrained. In particular, the necessity of bottom water anoxia, long considered a pre‐requisite, has been the subject of recent debate. In this study, we apply a micro‐stratigraphical, ichnological approach to determine bottom water oxygen conditions under, which Burgess Shale‐type biotas were preserved in the Middle Cambrian Spence Shale of Utah. Mudstones of the Spence Shale are characterized by fine scale (mm‐cm) alternation between laminated and bioturbated intervals, suggesting high‐frequency fluctuations in bottom water oxygenation. Whilst background oxygen levels were not high enough to support continuous infaunal activity, brief intervals of improved bottom water oxygen conditions punctuate the succession. A diverse skeletonized benthic fauna, including various polymerid trilobites, hyolithids, brachiopods and ctenocystoids suggests that complex dysoxic benthic community was established during times when bottom water oxygen conditions were permissive. Burgess Shale‐type preservation within the Spence Shale is largely confined to non‐bioturbated horizons, suggesting that benthic anoxia prevailed in intervals, where these fossils were preserved. However, some soft‐bodied fossils are found within weakly to moderately bioturbated intervals (Ichnofabric Index 2 and 3). This suggests that Burgess Shale‐type preservation is strongly favoured by bottom water anoxia, but may not require it in all cases. □Anoxia, Burgess Shale, Burgess Shale type‐preservation, Langston Formation, Spence Shale Member, Utah.     

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

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.     

Downsized Dinosaurs: The Evolutionary Transition to Modern Birds

Living birds are the most diverse land vertebrates and the heirs of a rich chapter in the evolution of life. The origin of modern birds from animals similar to Tyrannosaurus rex is among the most remarkable examples of an evolutionary transition. A wealth of recently discovered fossils has finally settled the century-old controversy about the origin of birds and it has made the evolutionary saga toward modern birds one of the best documented transitions in the history of life. This paper reviews the evidence in support of the origin of birds from meat-eating dinosaurs, and it highlights the array of fossils that connect these fearsome animals with those that fly all around us.