Skimming the surface with Burgess Shale arthropod locomotion

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates.     

ISOPOD TRACKWAYS FROM THE CRAYSSAC LAGERSTÄTTE, UPPER JURASSIC, FRANCE

Abstract:  Well-preserved arthropod trackways are described from the laminated limestones of the Crayssac Lagerstätte (south-west France, Lower Tithonian). They occur in sediments deposited in the temporary coastal mudflats of intertidal to supratidal zones. The trackways are referred to Pterichnus isopodicus isp. nov., and are interpreted as the locomotion traces of isopods. Different trackway morphotypes are recognized and clearly resulted from variations in the original consistency of the sediment. Sinuous trackways may correspond to vagrant activity on wet mud whereas numerous straight ones indicate a more rapid crawling on a soft-to-firm substratum (e.g. tidal flat during emersion). The preferred orientation of trackways indicates that isopods were crawling in a direction perpendicular to shoreline as a result of possible taxis induced by sediment wetness and/or by a migratory behaviour controlled by tidal rhythm. Unusually long emergence of the sediments may have favoured the preservation of dense networks of trackways. An isopod identity is supported by the general morphology of the tracks and the association of trackways with isopod body fossils. Archaeoniscus , which occurs abundantly in Late Jurassic deposits of England and France, was probably the trace-maker.     

Nonmarine aspects of the Cambrian Tonto Group of the Grand Canyon, USA, and broader implications

A re-examination of the classic time-transgressive model of the Cambrian Tonto Group of the Grand Canyon challenges some of the basic assumptions regarding strandline migration during the Sauk transgression, the applicability of sequence stratigraphy to epicratonic deposits, and the criteria by which siliciclastic deposits of the North American Cambrian system are categorized as marine. Indicators for high short-term depositional rates, coupled with the complex splay and coalescence of meter-scale parasequence architectures, detract from the shoreline migration and deepening seas model that has held sway for over 60 years. Ostensibly marine signatures, represented by sparse and fragmentary trilobite fossils and abundant bedding plane trace fossils, are closely associated with nonmarine signatures, represented by nonmarine palynoflora and clay mineral assemblages, and highly oxidized paleoweathering profiles. The vast extent of three-dimensional exposure of the Cambrian section in the Grand Canyon shows a paleogeography not of a single discrete shoreline translated laterally with rising and falling sea-level, but of a single mosaic setting of emergent shoals and rocky peaks, broad fluvial and tidal channels, and shallow pools and interchannel flats. Much detail concerning sediment dynamics and paleoenvironment is lost or overlooked when summary terms like “marginal marine” or “nearshore” are applied to such settings. It is suggested here that the Tonto Group, with its mixed marine and nonmarine attributes, and considering its historic role as a teaching model, be viewed not as an exceptional Cambrian deposit, but as an exemplar of epicratonic deposits on a pre-vegetated landscape.     

Trace fossils of the arthropod Camptophyllia from the Westphalian (Carboniferous) rocks of Lancashire, UK and their palaeoenvironmental context

Two arthropod trace fossils are described and analysed from the Carboniferous Lower Westphalian (C. communis and basal A. modiolaris chronozones) coal-bearing strata of Lancashire. The biserial trackway Diplichnites triassicus consists of five overlapping en echelon sets of 7–9 tracks preserved as epichnia and hypichnia in lacustrine siltstones. The trackway suggests subaqueous in-phase walking by a multi-segmented producer with a body length of 35–40 mm, width 17–22 mm, and 7–9 appendages. Curved, clustered, or laterally repeated, hypichnial lobes with transverse striations on the base of ripple cross-laminated sandstone are identified as Rusophycus versans. This trace fossil is interpreted as shallow resting or furrowing burrows of a homopodous arthropod, 30–60 mm long, 15–30 mm wide, and probably the same kind of arthropod as produced D. triassicus.A review of contemporary arthropod body fossils from Lagerstätten in Lancashire favours the onisciform, or Arthropleura like arthropod Camptophyllia as a potential producer of both of these trace fossils in a lacustrine palaeoenvironment.This study integrates the analysis of sediments, trace fossils and body fossils for reconstructing the arthropod biota and ecology in Westphalian lacustrine and crevasse splay fluvial palaeoenvironments.     

Obituary: Ron K. Pickerill (1947–2018)

Recently discovered tracks in alluvial sediments of the Late Paleocene Bullion Creek Formation of western North Dakota have been identified as belonging to the large eusuchian crocodile Borealosuchus formidabilis and an associated small arthropod. They are described as two new ichnotaxa. The first of these, Borealosuchipus hanksi igen. et isp. nov., is based on a partly complete trackway having fore- and hind-footprints as well as body marks preserved in hyporelief. These tracks are directly associated with numerous body fossils including articulated bones which provide strong implications for their probable origin. The second specimen, Koupichnium pentapodus isp. nov., is founded on a trackway of a small invertebrate in which a series of five pairs of footprints are arranged along a midline drag mark which is continuous throughout the length of its trackway. This trackway shares features with other fossils as well as living arthropod trackways in which four small feet and a larger posterior hind “pusher” foot are present. The two described new ichnotaxa represent the first recognized tracks from the Wannagan Creek local fauna.     

Reassessment of sediments and trace fossils from old red sandstone (Lower devonian) of Dunure,Scotland, described by John Smith (1909)

In 1909 John Smith, a Scottish naturalist and geologist,described 23 «genera and 51 «species of trace fossils from small patches of sediment associated with andesite lava flows at Dunure, Ayrshire. He interpreted the traces as evidence of a diverse invertebrate fauna which inhabited small pools and fissures in the lava surface between eruptions. Smith's collection (c. 300 specimens) was presented to the British Geological Survey, Edinburgh but has remained largely unstudied.Re-examination of the Smith collection shows thatit came from 3 separate localities which differ in composition of the ichnofaunas, associated sediments and sedimentary structures. Arthropod trackways dominate the ichnofauna occurring in laminated siltstone frequently with ripple marks and foam marks suggesting formation in shallow lacustrine conditions. Preservational and behavioural analysis of trace fossils reveals about 10 valid ichnogenera of locomotion and swimming trackways, resting traces, feeding trails but few burrows. Presence of early terrestrial arthropod traces is uncertain. Palaeocological interpretation is of ephemeral lakes in distal braid-plain situation subject to subsurface invasion of andesite lava producing fluidization and deformation of wet sediment. The Dunure ichnocoenosis shows unique diversity of Devonian arthropod trace fossils.     

Trace fossils in the Ediacaran-Cambrian transition: Behavioral diversification, ecological turnover and environmental shift

After taxonomic revision, trace fossils show a similarly explosive diversification in the Ediacaran-Cambrian transition as metazoan body fossils. In shallow-marine deposits of Ediacaran age, trace fossils are horizontal, simple and rare, and display feeding strategies related to exploitation of microbial matgrounds. Equally notable is the absence of arthropod tracks and sinusoidal nematode trails. This situation changed in the Early Cambrian, when a dramatic increase in the diversity of distinct ichnotaxa is associated was followed by the onset of vertical bioturbation and the disappearance of a matground-based ecology (‘‘agronomic revolution’’). On deep sea bottoms, animals have been present already in the Ediacaran, but ichnofaunas were poorly diverse and dominated by the horizontal burrows of undermat miners. As shown by the ichnogenus Oldhamia, this life style continued to be predominant into the Early, and to a lesser extent, Middle Cambrian. Nevertheless, there was an explosive radiation of behavioral programs during the Early Cambrian. When exactly the bioturbational revolution arrived in the deep sea is uncertain. In any case, the Nereites ichnofacies was firmly established in the Early Ordovician. The rich ichnofauna in the Early Cambrian Guachos Formation of northwest Argentina probably marks a first step in this ecological onshore-offshore shift.     

Vestiges of a beginning? Paleontological and geochemical constraints on early animal evolution

Inspired by molecular clock estimates, some biologists have proposed that animals in general, and bilaterian metazoans in particular, began to diverge substantially earlier than fossils indicate. Balavoine and Adoutte [Science 280 (1998) 397] specifically hypothesized that the Cambrian explosion documents parallel radiations within three major bilaterian clades that diverged from one another relatively early in the Neoproterozoic Era. The geological record is permissive with respect to such hypotheses, but not encouraging. The earliest evolution of animals certainly took place before the initial appearance of phosphatic animal microfossils or Ediacaran macrofossils, but bilaterian clades need not have been part of this metazoan “pre-history”. Alternatively, the evolution of large size, made possible by late Neoproterozoic oxygen increase, may have provided the selective environment in which stem bilaterians differentiated. Cambrian events per se appear to have begun in the wake of environmental perturbation and accompanying extinction near the Proterozoic–Cambrian boundary.     

Tracking the last elephants in Europe during the Würm Pleniglacial: the importance of the Late Pleistocene aeolianite record in SW Iberia

Abstract

In his short joint ventures across the world of vertebrate tracks, Richard Bromley recognized the aeolian sands as unsuitable soft substrates for their preservation. Only after his work in the Balearic Islands, a more systematic study of coastal aeolianites worldwide revealed that these depositional systems could preserve a highly important record of behavioural trace fossils for the evolution of vertebrates, especially in the Pleistocene, including extinct megafauna and the escalation of the Homo. Here we describe coastal aeolianites from the upper Pleistocene of SW Iberia, namely SW Portugal and Gibraltar, with trackways, tracks and trampled surfaces of the last elephants in mainland Europe. Photogrammetric 3?D modelling and analysis of the relevant proboscidean track levels allowed revision of and support for previous ichnotaxonomic identification to Proboscipeda panfamilia and behavioural interpretations of the producer. Smaller and very large trackways and footprints attributed to Palaeoloxodon antiquus are described and discussed according to new and recent, but always rare findings. The seeming progressive and definitive extinction of this species towards southern Iberia, following the same pattern for the replacement of the Neanderthals during the last initial Pleniglacial (until ca. 28 ka), suggests evidence for co-evolution.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

Trace fossils from lower Cambrian Hongjingshao Formation,Yunnan, China: Taxonomy,palaeoecology, palaeoenvironment

The Hongjingshao Formation (Cambrian Series 2, Stage 3) yields abundant, complex trace fossils that have not been systematically investigated up till now. Our detailed ichnological study of the traces from this formation in Malong area, Yunnan, China, recognizes four groups: (1) simple horizontal or sub-horizontal burrows; (2) complex branched burrow systems; (3) arthropod trails and; (4) simple vertical burrows belonging to the Cruziana ichnofacies. A trophic web is reconstructed for the community in Hongjingshao Formation based on both trace and body fossils. The ichnological and sedimentological features of the formation indicate an intertidal setting. The trace fossil assemblage demonstrates that Early Cambrian organisms were able to colonize very shallow marine environments, which further supports the landward expansion of the Early Cambrian ecosystems.     

Evolution of eye development in arthropods: phylogenetic aspects

The architecture of the adult arthropod visual system for many decades has contributed important character sets that are useful for reconstructing the phylogenetic relationships within this group. In the current paper we explore whether aspects of eye development can also contribute new arguments to the discussion of arthropod phylogeny. We review the current knowledge on eye formation in Trilobita, Xiphosura, Myriapoda, Hexapoda, and Crustacea. All euarthropod taxa share the motif of a proliferation zone at the side of the developing eye field that contributes new eye elements. Two major variations of this common motif can be distinguished: 1. The “row by row type” of Trilobita, Xiphosura, and Diplopoda. In this type, the proliferation zone at the side of the eye field generates new single, large elements with a high and variable cell number, which are added to the side of the eye and extend rows of existing eye elements. Cell proliferation, differentiation and ommatidial assembly seem to be separated in time but spatially confined within the precursors of the optic units which grow continuously once they are formed (intercalary growth). 2. The “morphogenetic front type” of eye formation in Crustacea + Hexapoda (Tetraconata). In this type, there is a clear temporal and spatial separation of the formation and differentiation processes. Proliferation and the initial steps of pattern formation take place in linear and parallel mitotic and morphogenetic fronts (the mitotic waves and the morphogenetic furrow/transition zone) and numerous but small new elements with a strictly fixed set of cells are added to the eye field. In Tetraconata, once formed, the individual ommatidia do not grow any more. Scutigeromorph chilopods take an intermediate position between these two major types. We suggest that the “row by row type” as seen in Trilobita, Xiphosura and Diplopoda represents the plesiomorphic developmental mode of eye formation from the euarthropod ground pattern whereas the “morphogenetic front type” is apomorphic for the Tetraconata. Our data are discussed with regard to two competing hypotheses on arthropod phylogeny, the “Tracheata” versus “Tetraconata” concept. The modes of eye development in Myriapoda is more parsimonious to explain in the Tetraconata hypothesis so that our data raise the possibility that myriapod eyes may not be secondarily reconstructed insect eyes as the prevailing hypothesis suggests.     

Megarhizoliths in Pleistocene aeolian deposits from Gran Canaria (Spain): Ichnological and palaeoenvironmental significance

The Pleistocene dune field of Tufia, located on the east of Gran Canaria (Spain), contains different stratigraphic levels of indurated pillar-like structures that are interpreted as megarhizoliths. The megarhizoliths occur at the top of different aeolian sets and reach 31.5 cm in diameter and over 1 m in height. These scattered, free-standing, vertical, cylindrical-to-slightly conical columns usually appear as hollow cylinders, displaying elliptical cross-sections aligned with the prevailing wind. On the leeward side of some specimens the external wall shows a tail of rock matrix resembling a sort of “wind shadow”. These tails and other remains of the associated rock matrix show a texture composed of long, horizontal, parallel cylinders orientated with the wind.Internally the most complete structures show five concentrically arranged zones: Zone (a), is a central pore corresponding to the cavity originally occupied by the root; Zones (b) and (c), which include alveolar and laminated carbonate textures indicating that carbonate precipitation was mostly induced by the roots and their associated microorganisms; and Zones (d) and (e), consisting mostly of aeolian sands. In (d) the sand grains show thin micritic coatings whereas in (e) vadose aragonite cements can be seen on the grain surface suggesting a less biogenic influence in their formation. The degree of cementation and the time of the precipitation of carbonate around the roots controlled the preservation of these zones. Thus, in some cases, Zones b, c and/or d are not preserved. Cylinders are up to 30× the diameter of the root that nucleated them.The presence of the megarhizoliths at the top of the aeolianite beds indicates that aeolian sedimentation halted several times, allowing soil formation and plant colonisation during slightly more humid periods. The occurrence of megarhizoliths is further proof of the alternation of arid and slightly more humid climates in the north Atlantic during the last glacial period. It is also noted that they may be misinterpreted as animal trace fossils or tree trunk casts, resulting in incorrect ichnological or palaeoenvironmental interpretations.     

The first record of xiphosurid (arthropod) trackways from the Saltwick Formation, Middle Jurassic of the Cleveland Basin, Yorkshire

Trackways attributed to a xiphosurid (arthropod) maker are described for the first time from the Middle Jurassic Saltwick Formation, Ravenscar Group, of the Cleveland Basin, Yorkshire. The trackways are assigned to Kouphichnium aff. variabilis (Linck, 1949) and clearly demonstrate the heteropody and varied behaviour of the maker. Their occasional asymmetry indicates a slight rotation of the body relative to the direction of locomotion. The Saltwick Formation has generally been regarded as a coastal plain deposit, and the presence of xiphosurid tracks reinforces the suggestions of periodic marine influences.     

The ‘Orsten’—More than a Cambrian Konservat-Lagerstätte yielding exceptional preservation

In several areas of southern Sweden, limestone nodules, locally called Orsten occur within bituminous alum shales. These shales and nodules were deposited under dysoxic conditions at the bottom of what was most likely a shallow sea during the late Middle to Upper Cambrian (ca. 500 million years ago). Subsequently, the name ‘Orsten’ has been referred to particular, mainly arthropod, fossils from such nodules, and, in a wider sense, to the specific type of preservation of minute fossil through secondarily phosphatization. This preservation is exceptional in yielding uncompacted and diagenetically undeformed three-dimensional fossils. ‘Orsten’-type preservation resulted from incrustation of a thin external layer and also by impregnation by calcium phosphate and, therefore, mineralization of the surface of the former animals during early diagenesis. Primarily, this type of preservation seems to have affected only cuticle-bearing metazoans such as cycloneuralian nemathelminths and arthropods. ‘Orsten’ preservation in this sense seems to be limited by size, in having yielded no partial or complete animals larger than 2 mm. On the other end of the scale, even larvae 100 μm long are preserved, often more complete than larger specimens, and details such as setules and pores smaller than 1 μm can be observed. Fossils preserved in such a manner are almost exclusively hollow carcasses, but can be filled secondarily; less common are completely phosphatized compact specimens. The high quality of preservation makes the Swedish ‘Orsten’ a typical Konservat-Lagerstätte. Yet, its special type of preservation is more widespread in time and geographical distribution than assumed initially, and the origin of the phosphate is not necessarily restricted just to one source. Subsequent to the first discoveries of limb fragments of Cambrian arthropods in 1975, animals in this special preservational type have been discovered in several continents and across a broad stratigraphic range including even Proterozoic strata. The latter have yielded early cleavage and metazoan embryonic stages, expanding knowledge on the preservational capacities of the ‘Orsten’. Here, we report the recent status of our research on the ‘Orsten’ and give perspectives for future exploration on a worldwide scale, particularly in light of a recently formed international research group named Center of Orsten Research and Exploration (C.O.R.E.).     

Phylogenetic affinities and taphonomy of Brooksella from the Cambrian of Georgia and Alabama, USA

Siliceous “star cobbles”, referred to the enigmatic genus Brooksella, are abundant in the Conasauga Formation of the Coosa River Valley of Alabama and Georgia, USA. Explaining the phylogenetic affinities and taphonomic history of Brooksella has been difficult and contentious. Brooksella has, at times, been referred to: 1, the cnidarian order Scyphomedusae; 2, the cnidarian class Protomedusae (order Brooksellida); 3, as algae; 4, as a trace fossil; and 5, as a feature of inorganic origin.Macroscopic, microscopic, and computer-assisted tomographic analysis of Brooksella from the Conasauga Formation suggests that the “star cobbles” represent exceptionally preserved body fossils of simple construction. Morphology of star cobbles is most consistent with a siliceous (hexactinellid) sponge interpretation. Specimens show wide morphologic variation, including gradational patterns, suggesting that a single species name (Brooksella alternata) should be used to embrace all forms described from the Coosa Valley. B. alternata includes specimens having a variable number of radially disposed lobes divided by radial grooves, and often a central opening inferred to be an osculum on one side. Lobes in many specimens terminate in small openings. Small craterlike structures, inferred to be ostia, are present on the external surface. Radial internal cavities occupy the lobes. Specimens from the Conasauga Formation have siliceous spicules preserved surficially and internally.The three-dimensional nature of most “star cobbles” suggests rapid fossil diagenesis, perhaps mediated by the activities of microbial consortia that quickly formed biofilms around the dead hosts.     

Perinatal adaptation in mammals: the impact of metabolic rate

Mammalian birth is accompanied by profound changes in metabolic rate that can be described in terms of body size relationship (Kleiber's rule). Whereas the fetus, probably as an adaptation to the low intrauterine pO₂, exhibits an “inappropriately” low, adult-like specific metabolic rate, the term neonate undergoes a rapid metabolic increase up to the level to be expected from body size. A similar, albeit slowed, “switching-on” of metabolic size allometry is found in human preterm neonates whereas animals that are normally born in a very immature state are able to retard or even suppress the postnatal metabolic increase in favor of weight gain and O₂ supply. Moreover, small immature mammalian neonates exhibit a temporary oxyconforming behavior which enhances their hypoxia tolerance, yet is lost to the extent by which the size-adjusted metabolic rate is “locked” by increasing mitochondrial density. Beyond the perinatal period, there are no other deviations from metabolic size allometry among mammals except in hibernation where the temporary “switching-off” of Kleiber's rule is accompanied by a deep reduction in tissue pO₂. This gives support to the hypothesis that the postnatal metabolic increase represents an “escape from oxygen” similar to the evolutionary roots of mitochondrial respiration, and that the overall increase in specific metabolic rate with decreasing size might contribute to prevent tissues from O₂ toxicity.     

Devonian tetrapod trackways and trackmakers; a review of the fossils and footprints

The earliest tetrapods are known from the Upper Devonian. Their remains are becoming better known from increasing numbers of specimens, localities, environments and ichnofossils. Each of the eight (or possibly nine) genera now represented by skeletal fossils is reviewed in its sedimentological, faunal and stratigraphic context, with an assessment of what might be inferred about the habitus and locomotory capabilities of each. Fossil trackways and their interpretations are then re-examined in the context of the known body forms, and consideration given to the degree of fit between the skeletal fossils, the trackways and their interpretations. The currently known Devonian tetrapods are unlikely to have made any of the known tracks, unless they were produced under water. Neither the skeletal fossils nor the trackways show good evidence of terrestrial locomotion among Devonian tetrapods. When the fossil material and recent phylogenetic analyses are taken in combination, it appears that neither tetrapods nor limbs with digits are likely to have arisen before the Frasnian. This should be borne in mind in palaeoecological studies of these animals.