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.     

Morphology,faunas and genesis of ordovician hardgrounds from Southern Ontario,Canada

We have used associations of different microfacies to define facies (or microfacies associations) which form reasonably well-defined sequences, which we infer, from analogies with recent and ancient carbonate environments, to have been deposited in a shelf environment characterized by small-scale topographic differentiation into shoal, slope and basinal environments.Shoal environments are characterized by typically cross-bedded, well-sorted bioclastic sands, with intershoal areas consisting of interbedded bioclastic sands and heavily bioturbated finer-grained carbonates.Slope and “basinal” environments are typically represented by “proximal” and “distal” cycles respectively. These we compare with deposits of carbonate ramp bypass channels, and with the more thoroughly studied deep-water clastic submarine fans. Many of the strong variations in environmental energy in these proximal and distal cycles can be attributed to migration of channels on the fans and the effect of funnelling of storm surges down the channels.Although hardground morphology and faunas are mostly related to local effects such as intensity of scouring, time of exposure, topographic differentiation of the surface and other factors, differing hardground types tend to be found in different environments. Smooth and rolling hardgrounds occur in the deeper distal environments, where the beds were subject to only slight scour and often limited exposure before renewed sedimentation. Hummocky and undercut hardgrounds are characteristic of the middle parts of proximal cycles, where they developed marginally to the main bypass channel, and in intershoal areas. Both these areas are sites of intermittent sedimentation and moderate turbulence, where cemented beds may be exposed for some time in environments optimal for attached benthos. These hardgrounds usually contain the most diverse hardground biotas. Pebbly and reworked hardgrounds occur in coarse, basal units of proximal cycles, which are interpreted as the grain-flow fillings of the central parts of bypass channels, though isolated examples occur in intershoal areas and in the higher parts of proximal channels. These hardgrounds contain low-diversity faunas, reflecting the stresses imposed by intermittent or constant abrasion; though some contain more diverse faunal assemblages formed after redeposition.     

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.     

Crinoids, hardgrounds, and community succession: The Silurian Laurel—Waldron contact in southern Indiana

Halleck, Margaret S.: Crinoids, hardgrounds, and community succession: The Silurian Laurel-Waldron contact in southern Indiana.
The uppermost surface of the Silurian Laurel Limestone at its contact with the Waldron Shale in southeastern Indiana was a hardground lithified prior to the deposition of the Waldron. Evidence for this conclusion is the presence of attached palmate crinoid roots, auloporid corals, and craniid brachiopods on the Laurel surface; the irregularity of the contact with the Waldron; and a pyritic veneer at this contact. The hardground apparently had a submarine origin. In addition to the attached epifauna mentioned above, algal-sediment 'clods' formed on this surface. Some of these accumulated around the crinoid stems, causing them to produce cirral extensions. The resulting community was a crinoid 'meadow' with algal growths forming sediment traps around and between the crinoids. Later stages of Waldron Shale deposition led to the development of a soft-bottom community.     

Faunal distribution and colonization strategy in a Middle Ordovician hardground community

The attached fauna of one of the many hardgrounds from the Galena Group (Trentonian Substage) of the Upper Mississippi Valley is described. The fauna is composed of three principal elements, viz. (1) borers, including Cicatricula retiformis ichnogen. et ichnosp. nov., (2) pelmatozoans with encrusting holdfasts, and (3) bryozoans. Analysis of the distribution of members of each population on the hardground shows that most are strongly aggregated. The nature of, and reasons for, such aggregations are considered in the light of comparable Recent shallow-water marine populations. The community on this hardground, and those on other Galena Group hardgrounds, are immature. This is a consequence of frequent and damaging scour, which these organisms were poorly adapted to resist.     

Cambrian stalked echinoderms show unexpected plasticity of arm construction

Feeding arms carrying coelomic extensions of the theca are thought to be unique to crinoids among stemmed echinoderms. However, a new two-armed echinoderm from the earliest Middle Cambrian of Spain displays a highly unexpected morphology. X-ray microtomographic analysis of its arms shows they are polyplated in their proximal part with a dorsal series of uniserial elements enclosing a large coelomic lumen. Distally, the arm transforms into the more standard biserial structure of a blastozoan brachiole. Phylogenetic analysis demonstrates that this taxon lies basal to rhombiferans as sister-group to pleurocystitid and glyptocystitid blastozoans, drawing those clades deep into the Cambrian. We demonstrate that Cambrian echinoderms show surprising variability in the way their appendages are constructed, and that the appendages of at least some blastozoans arose as direct outgrowths of the body in much the same way as the arms of crinoids.     

Late Cambrian hard substrate communities from Montana/Wyoming: the oldest known hardground encrusters

Hardground surfaces from the Late Cambrian Snowy Range Formation in Montana/Wyoming are the oldest known non-reefal hard substrates exhibiting encrusting fossils. These surfaces range in age from Early Franconian to early Trempealeauan. Hardgrounds were developed on slightly hummocky to planar, truncated surfaces of glauconite-rich, carbonate, flat pebble conglomerates, which were deposited during episodes of storm scouring in shallow subtidal environments of the Montana/Wyoming shelf. Snowy Range hardgrounds are encrusted by a low diversity assemblage of fossils dominated by simple discoidal holdfasts of pelmatozoans, probably crinoids, and including small conical spongiomorph algae? and probable stromatolites. Macroborings (e.g. Trypanites) are notably absent from all hardground surfaces, although sharp-walled, vertical, cylindrical holes (borings?) occur in micrite clasts imbedded in certain flat pebble conglomerates. No evidence of faunal succession or microecologic partitioning of irregular surfaces was observed on these Cambrian hardgrounds.     

Evolution of pelagic swells from hardground analysis (Bathonian–Oxfordian, Eastern External Subbetic, southern Spain)

The Middle Bathonian to Middle Oxfordian interval in the Eastern External Subbetic (Betic Cordillera, SE Spain) is characterized by Ammonitico Rosso facies including various stratigraphic breaks. Five hardground-bounded units are recognized in relation to hiatuses in the ammonite record at the following stratigraphic boundaries: Hg1 (Lower–Middle Bathonian), Hg2 (Middle–Upper Bathonian), Hg3 (Lower–Middle Callovian), Hg4 (Middle–Upper Callovian), and Hg5 (Callovian–Oxfordian). Interesting features of these hardgrounds include their microfacies, ferruginous crusts and macro-oncoids, taphonomy of macroinvertebrates, trace fossils, neptunian dykes, and the hiatuses associated with each of them. The main hardgrounds (Hg1, Hg2, and Hg5) contain trace fossils of the Cruziana and Trypanites ichnofacies as well as abundant fossil macroinvertebrates with taphonomic features evidencing corrasion, early diagenesis, and reworking, indicating substrate evolution from softground to hardground. Neptunian dykes affected the trace fossils and ammonoid moulds, and their walls and the hardground surfaces were colonized by ferruginous microbial crusts. These features are characteristic of the External Subbetic pelagic swells, where the absence of sedimentation, sediment bypassing and erosion, and early diagenesis during relative sea-level falls produced hardgrounds. The neptunian dykes are indicative of tectonic activity in the areas of pelagic swells. Ferruginous crusts and macro-oncoids developed only on hardground surfaces and neptunian dykes walls prior to deposition of condensed bioclastic beds, which are interpreted as the first deposits after hardground development and are related to the onset of transgression. The varying ranges of the gaps as well as lateral facies changes are related to different local paleobathymetry controlled by the activity of listric faults.     

Patterns of magnesium content in Arctic bryozoan skeletons along a depth gradient

A growing body of evidence suggests that ocean acidification acting synergistically with ocean warming alters carbonate biomineralization in a variety of marine biota. Magnesium often substitutes for Ca in the calcite skeletons of marine invertebrates, increasing their solubility. The spatio-environmental distribution of Mg in marine invertebrates has seldom been studied, despite its importance for assessing vulnerabilities to ocean acidification. Because pH decreases with water depth, it is predicted that levels of Mg in calcite skeletons should also decrease to counteract dissolution. Such a pattern has been suggested by evidence from echinoderms. Data on magnesium content and depth in Arctic bryozoans (52 species, 103 individuals, 150 samples) are here used to test this prediction, aided by comparison with six conceptual models explaining all possible scenarios. Analyses were based on a uniform dataset spanning more than 200 m of coastal water depth. No significant relationship was found between depth and Mg content; indeed, the highest Mg content among the analyzed taxa (8.7 % mol MgCO₃) was recorded from the deepest settings (>200 m). Our findings contrast with previously published results from echinoderms in which Mg was found to decrease with depth. The bryozoan results suggest that ocean acidification may have less impact on the studied bryozoans than is generally assumed. In the broad context, our study exemplifies quantitative testing of spatial patterns of skeletal geochemistry for predicting the biological effects of environmental change in the oceans.     

Hettangian (Early Jurassic) Dinosaur Tracksites from the Mecsek Mountains,Hungary

A rare example of a North American Jurassic hardground is found in the Carmel Formation of southwestern Utah. The Carmel hard‐ground was formed across a carbonate lagoon from an oolitic shoal seaward to a subtidal shelly facies landward. It has an abundant bivalve fauna consisting of thick layers of encrusters (the oyster Liostrea and the plicatulid Plicatula), borers (the ichnofossil Gastro‐chaenolites with the mytilid Lithophaga often preserved inside), and nestlers (the mytilid Modiolus). A rare soft‐bodied bryozoan (Arach‐nidium) is preserved by bioimmuration in the attachment scars of Liostrea; this is the first bioimmuration recorded from the Jurassic of North America, and the first bioimmuration recorded from a hard‐ground. The phoronid boring Talpina is present in some Liostrea shells; it was apparently excavated after the death of these oysters. The Carmel hardground community does not contain other fossils, such as serpulids, brachiopods, foraminiferans, and skeletal bryo‐zoans, typical of Jurassic hardgrounds elsewhere. It represents a low diversity molluscan community developed in a restricted marine environment.     

Tentaculate Fossils from the Cambrian of Canada (British Columbia) and China (Yunnan) Interpreted as Primitive Deuterostomes

Molecular and morphological evidence unite the hemichordates and echinoderms as the Ambulacraria, but their earliest history remains almost entirely conjectural. This is on account of the morphological disparity of the ambulacrarians and a paucity of obvious stem-groups. We describe here a new taxon Herpetogaster collinsi gen. et sp. nov. from the Burgess Shale (Middle Cambrian) Lagerstätte. This soft-bodied vermiform animal has a pair of elongate dendritic oral tentacles, a flexible stolon with an attachment disc, and a re-curved trunk with at least 13 segments that is directed dextrally. A differentiated but un-looped gut is enclosed in a sac suspended by mesenteries. It consists of a short pharynx, a conspicuous lenticular stomach, followed by a narrow intestine sub-equal in length. This new taxon, together with the Lower Cambrian Phlogites and more intriguingly the hitherto enigmatic discoidal eldoniids (Cambrian-Devonian), form a distinctive clade (herein the cambroernids). Although one hypothesis of their relationships would look to the lophotrochozoans (specifically the entoprocts), we suggest that the evidence is more consistent with their being primitive deuterostomes, with specific comparisons being made to the pterobranch hemichordates and pre-radial echinoderms. On this basis some of the earliest ambulacrarians are interpreted as soft-bodied animals with a muscular stalk, and possessing prominent tentacles.     

ORDOVICIAN (ARENIG–CARADOC) SYNTROPHIIDINE BRACHIOPODS FROM THE EAST BALTIC REGION

Abstract:  Syntrophiidine brachiopods are a rare and poorly known component of Ordovician Baltoscandian faunas. They appear in the East Baltic in the Billingenian (lower Arenig) as part of the earliest known benthic assemblages dominated by elements of the Palaeozoic Evolutionary Fauna. These faunal assemblages usually include bryozoans, ostracodes, and the earliest known porambonitoids, strophomenides and endopunctate orthides, such as Idiostrophia and Orthidium , which later became characteristic of the Whiterockian brachiopod assemblages in Laurentia, but by that time had disappeared from Baltica. The superfamily Syntrophioidea reappears in Baltoscandia in the mid Caradoc. In contrast, Porambonitoidea remained the integral part of the Baltoscandian brachiopod associations through the Ordovician. Porambonites , herein redefined on the basis of restudy of the type species P. intermedius , includes only smooth porambonitoids; taxa with the distinctive ornament of radiating rows of pits first appeared in the group in the mid Arenig. The taxa Eoporambonites gen. nov., Tetralobula peregrina sp. nov., Idiostrophia prima sp. nov. and Idiostrophia tenuicostata sp. nov. are erected.     

Revision of Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia

Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia was originally described as the earliest crinoid(?) known from the fossil record. Recently, Conway Morris and Ausich & Babcock have questioned whether Echmatocrinus is in fact an echinoderm, comparing it instead to cnidarians with a polyp-like body and pinnate tentacles, and other authors are beginning to use this reinterpretation. We studied the well-preserved holotype of Echmatocrinus brachiatus, two paratypes, and 18 new specimens recovered from different levels in the Burgess Shale sequence at three localities. All are preserved as pyrite films in dark shale with relatively little relief, suggesting a lightly skeletized body. Complete specimens have a long, slightly tapering, large-plated attachment stalk, a conical cup or calyx with numerous small to medium-sized irregular plates, and 7–10 short arms with heavier plating and (in the holotype) soft appendages alternating from opposite sides of several arms. Several morphologic features indicate that Echmatocrinus is an echinoderm and has crinoid affinities: (1) Sutured plates, shown by darker depressed sutures, slightly raised plate centers, and oriented plate ornament, cover all major parts of the body; (2) reticulate surface ornament in the pyrite film on the plates of all specimens matches the ornament in the Burgess Shale edrioasteroid Walcottidiscus, an undoubted echinoderm, but not the pyritized surfaces of other metazoans in the fauna; (3) this distinctive ornament may represent the surface expression of microporous stereom; (4) possible ligament or muscle pads are present between the arm ossicles to fold and unfurl the more heavily plated arms. Within the echinoderms, only crinoids commonly have a calyx attached by a stalk or stem to the substrate and bear erect, moveable, uniserial arms for feeding. Although Echmatocrinus shows some resemblance to octocorals in overall body shape as an attached suspension feeder, almost all the details are different, indicating that Echmatocrinus is most likely unrelated to this group. All complete specimens of Echmatocrinus are attached to hard substrates, either another fossil or skeletal debris. The new specimens indicate that Echmatocrinus was twice as common (about 0.02%) in the Burgess Shale fauna as previously recorded and represents one of the earliest attached, medium-level, skeletized, suspension feeders or microcarnivores in the fossil record.     

Graptolite (Hemichordata,Pterobranchia) preservation and identification in the Cambrian Series 3

Due to inadequate preservation, pterobranchs are often difficult to identify in the fossil record, and a better understanding of preservational modes and diagenetic and metamorphic effects is needed for their recognition. Pterobranch hemichordates are common in Cambrian Stage 5 and younger sedimentary rocks, but are frequently overlooked. Often, pterobranch hemichordate colonies have been considered to be algal remains or hydroids. Re‐examination of Cambrian Burgess Shale algae reveals that the genera Yuknessia and Dalyia can be recognized as putative early representatives of pterobranch hemichordates. Distinct fusellar construction of the individual zooidal tubes and branching of the creeping proximal part of the colonies are found in the morphologically similar rhabdopleurid pterobranch genus Sphenoecium. The erect tubes of Sphenoecium do not branch and can reach a length of several centimetres. The development of the fusellar construction in this taxon shows a highly irregular development of the suture patterns, but a fairly consistent height of the individual fuselli. The taxon is widely distributed in the Cambrian Series 3, but has regularly been identified as a hydroid or an alga. Sphenoecium wheelerensis from the Cambrian Wheeler Shale of Utah is described as new.     

A cryptic record of Burgess Shale‐type diversity from the early Cambrian of Baltica

Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).     

New data on molluscs and their shell microstructures from the Middle Cambrian Gowers Formation, Australia

Abstract: Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation (Ptychagnostus atavus/Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi‐nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite‐shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre‐like microstructures in many of these fossils reveals how widespread these strong (fracture‐resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov.     

Cambrian to Cretaceous changes in hardground communities

The changing nature of the communities of boring and encrusting taxa found on upward-facing hard-grounds has been studied from the standpoints of (a) diversity, (b) faunal composition, and (c) nature of the niches occupied. After a rapid initial increase in the early Palaeozoic, diversity remained at much the same level from the Middle Ordovician until the late Cretaceous. However, there is a considerable turnover in the identity of the individual taxa between successive sample intervals. The incoming and outgoing of the major groups parallel their fortunes in the marine realm as a whole. Niche analysis suggests that the same feeding levels are occupied for most of the history of hardground communities, but Mesozoic faunas contain a much higher proportion of species with true exoskcletons, or which lived infaunally. The evolution of these forms was probably influenced by the Mesozoic radiation of marine predators and duriphages, but it also resulted in Mesozoic hardground faunas being more resistant than their Palaeozoic counterparts to episodic corrasion. Resulting higher population densities in the Mesozoic were probably one reason why cavity faunas beneath some of these hardground surfaces are more diverse than those beneath Palaeozoic examples. □ Hardground, community, evolution.     

Phylogenetic Significance of the Burgess Shale Crustacean Canadaspis

The crustaceans, like the other major living groups of arthropods, have a long evolutionary history. The earliest examples occur in the Cambrian, and fossils of this age are a critical source of evidence of relationships both within the Crustacea, and between the Crustacea and other major arthropod groups. Canadaspis perfecta, from the Middle Cambrian Burgess Shale, is important as one of the oldest well-documented crustaceans. The evidence for reconstructing its remarkable combination of primitive and derived characters is reviewed, and its possible phylogenetic significance re-assessed.     

Dumbbell-shaped gogiid clusters: the oldest evidence of secondary tiering for stalked echinoderms

Among 381 specimens of Cambrian stalked echinoderms from eastern Guizhou, China examined, several slabs ( n  = 19) contain either dumbbell-shaped or v-shaped echinoderm clusters. Four slabs of Globoeocrinus globules Zhao, Parsley & Peng, 2008 from the middle-upper part (Cambrian Series 3 portion) of the Kaili Formation are prepared to reveal the attachment sites. Articulated gogiid echinoderms are reported to be attached to both sides of inarticulate (organophosphatic) brachiopods; thus, allowing me to interpret that the larvae of these gogiids were capable of attaching to live benthic brachiopods. This study documents the one of the earliest examples of echinoderms employing secondary tiering, which elevates an organism higher into the benthic boundary layer. Many of the gogiid echinoderm pairs attached to a live brachiopod are similar in size, indicating they were from a single larval spatfall.     

Deciphering the early evolution of echinoderms with Cambrian fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.