The Eager Formation (Cambrian Stage 4) of the Cranbrook area, British Columbia, contains abundant, sometimes beautifully preserved, trilobites and rare non-biomineralized taxa. Trilobites were first reported just over a century ago but have received little research attention, resulting in uncertainty in the number and identity of species within the assemblage and ambiguity in the age of the fauna. The trilobites of the Eager Formation in the Cranbrook area are described herein based largely upon material collected in 2015. A minimum of eight (and perhaps up to 11) trilobite species are recognized. The four species of olenelloids (Olenellus santuccii Webster n. sp., Olenellus? schofieldi, Mesonacis eagerensis, and Wanneria cranbrookense Webster n. sp.) are by far the most abundant elements of the fauna. At least two, and perhaps as many as five, species of dorypygid are present, as are two species of “ptychoparioids”. Paucity and poor preservational quality of specimens mean that the various dorypygid and “ptychoparioid” species are left in open nomenclature. Trilobite diversity in the Cranbrook Lagerstätte is comparable to that within other Lagerstätten from Cambrian Stage 4 (Series 2) of Laurentia. The diversity and abundance of trilobites, combined with biostratinomic and trace fossil data, suggest that the assemblage is autochthonous and/or parautochthonous, and that the local environment was at least periodically able to support a “typical” benthic trilobite community. The age of the Cranbrook Lagerstätte is constrained to lie within the middle Dyeran Stage of Laurentia, within what was a substantial stratigraphic gap in the distribution of Burgess Shale-type Lagerstätten.

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Radiodonta is a clade of stem euarthropods of central importance to our understanding of the evolution of this phylum. Radiodonts include some of the largest early Paleozoic animals; however, little is known about their ontogeny. We present an analysis of molting patterns and ontogeny in the radiodont Stanleycaris based on 265 exceptionally preserved specimens from the mid-Cambrian (Wuliuan) Burgess Shale. Ranging in size from 10 to 83 mm, they constitute the most extensive radiodont ontogenetic series known. Using a novel morphospace approach, we show that putative carcasses and exuviae can be quantitatively distinguished by the particular suites of structures preserved and their modes of preservation. We propose that Stanleycaris, and probably other radiodonts, molted via a suture near the anterior of the trunk. Similar anterior molting strategies, with a suture located at the head–trunk boundary, are shared with some Cambrian euarthropods and are potentially ancestral. Allometric analyses suggest that as Stanleycaris body size increased, the head sclerite and neck became relatively broader, while the trunk and flaps became slightly longer. The eyes developed precociously, indicating an important role of visual processing in juveniles. Finally, we find evidence for an initial anamorphic developmental phase, where segment number increased at least from 11 or 12 up to 17, followed by an epimorphic phase, in which growth continued without segment addition. This is consistent with the hypothesis that finite postembryonic segment addition (hemianamorphosis) is ancestral for arthropods and refines the timing of the origin of this important developmental mode.

Ordovician open marine Lagerstätten are relatively rare and widely dispersed, producing a patchy picture of the diversity and biogeography of nonmineralized marine organisms and challenging our understanding of the fate of Cambrian groups. Here, for the first time, we report soft-bodied fossils, including a well-preserved marrellomorph arthropod, fragmentary carapaces, and macroalgae, from the Late Ordovician (Katian) Upper Member of the Kirkfield Formation near Brechin, Ontario. The unmineralized elements and associated exceptionally preserved shelly biota were entombed rapidly in storm deposits that smothered the shallow, carbonate-dominated shelf. The marrellomorph, Tomlinsonus dimitrii n. gen. n. sp., is remarkable for its ornate, curving cephalic spines and pair of hypertrophied appendages, suggesting a slow-moving, benthic lifestyle. Reevaluation of marrellomorph phylogeny using new data favors an arachnomorph affinity, although internal relationships are robust to differing outgroup selection. Clades Marrellida and Acercostraca are recovered, but the monophyly of Marrellomorpha is uncertain. The new taxon is recovered as sister to the Devonian Mimetaster and, as the second-youngest known marrellid, bridges an important gap in the evolution of this clade. More generally, the Brechin biota represents a rare window into Ordovician open marine shelf environments in Laurentia, representing an important point of comparison with contemporaneous Lagerstätten from other paleocontinents, with great potential for further discoveries.

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Exceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

Radiodonts (stem Euarthropoda) were ecologically diverse, but species generally displayed limited functional specialization of appendages along the body axis compared with crown group euarthropods. This is puzzling, because such functional specialization is considered to have been an important driver of euarthropod ecological diversification. One way to circumvent this constraint could have been the functional specialization of different parts of the frontal appendages, known to have been ecologically important in radiodonts. This hypothesis has yet to be tested explicitly. Here we redescribe the poorly known mid-Cambrian hurdiid radiodont Stanleycaris hirpex from the Burgess Shale (Stephen Formation) and quantitatively assess functional specialization of the frontal appendages of stem euarthropods. The appendages of Stanleycaris are composed of 14 podomeres, variously differentiated by their possession of pectinate endites, mono- to trifurcate medial gnathites, and outer spines. The oral cone is tetraradially organized and can be uniquely distinguished from those of other hurdiids by the presence of 28 rather than 32 smooth tridentate plates. Our phylogenetic analysis finds Stanleycaris in a grade of hurdiids retaining plesiomorphic raptorial appendicular functionality alongside derived adaptations for sweep feeding and large, bilaterally opposed gnathites. We conclude that the latter performed a masticatory function, convergent with gnathal structures like mandibles in various panarthropods. Taken together, Stanleycaris and similar hurdiids provide an extreme example of the evolution of division of labor within the appendage of a stem euarthropod and suggest that this innovation may have facilitated the functional transition, from raptorial to sweep feeding, at the origin of the hurdiid clade.

The middle (Wuliuan Stage) Cambrian Burgess Shale is famous for its exceptional preservation of diverse and abundant soft-bodied animals through the “thick” Stephen Formation. However, with the exception of the Walcott Quarry (Fossil Ridge) and the stratigraphically older Tulip Beds (Mount Stephen), which are both in Yoho National Park (British Columbia), quantitative assessments of the Burgess Shale have remained limited. Here we first provide a detailed quantitative overview of the diversity and structure of the Marble Canyon Burgess Shale locality based on 16,438 specimens. Located 40 km southeast of the Walcott Quarry in Kootenay National Park (British Columbia), Marble Canyon represents the youngest site of the “thick” Stephen Formation. We then combine paleoecological data sets from Marble Canyon, Walcott Quarry, Tulip Beds, and Raymond Quarry, which lies approximately 20 m directly above the Walcott Quarry, to yield a combined species abundance data set of 77,179 specimens encompassing 234 species-level taxa. Marble Canyon shows significant temporal changes in both taxonomic and ecological groups, suggesting periods of stasis followed by rapid turnover patterns at local and short temporal scales. At wider geographic and temporal scales, the different Burgess Shale sites occupy distinct areas in multivariate space. Overall, this suggests that the Burgess Shale paleocommunity is far patchier than previously thought and varies at both local and regional scales through the “thick” Stephen Formation. This underscores that our understanding of Cambrian diversity and ecological networks, particularly in early animal ecosystems, remains limited and highly dependent on new discoveries.

The discovery of a new naraoiid nektaspid in the Upper Silurian (Pridolian) of southeastern Ontario significantly extends the range of this unusual group. Nektaspids are nonmineralized arthropods typical of Early and Middle Cambrian soft-bottom communities, but were thought to have become extinct in the Late Ordovician. The unique holotype specimen of Naraoia bertiensis n. sp. comes from a Konservat–Lagerstätte deposit renowned for its eurypterid fauna (the Williamsville Member of the Bertie Formation). Naraoia bertiensis lacks thoracic segments and is morphologically similar to Naraoia compacta from the Middle Cambrian Burgess Shale, save for the presence of a long ventral cephalic doublure and a subtly pointed posterior shield. To examine the phylogenetic relationships of the new naraoiid, we coded characters of the holotype specimen and of nine previously described nektaspids. The results confirm a sister taxon relationship between Naraoia compacta and Naraoia bertiensis and the monophyly of nektaspid forms lacking thoracic segments (family Naraoiidae). This latter group may have arisen from an ancestral segment-bearing form through heterochronic loss of thoracic segments early in the Cambrian. The disjunct occurrence of a naraoiid nektaspid in the Late Silurian resembles the reappearance of other “Lazarus taxa” that were thought to have been eliminated during mass extinction events. The naraoiid lineage survived the Late Ordovician biotic crisis, but in this case the “Lazarus effect” seems likely to be taphonomic in origin.

Yuknessia Walcott, 1919 recently was transferred from the green algae to the Phylum Hemichordata on the basis of new details observed for the type species, Y. simplex, from the Burgess Shale Formation (Cambrian Stage 5) of British Columbia. This has prompted reexamination of material attributed to Yuknessia from various Cambrian localities in South China. Findings preclude both a Yuknessia and a hemichordate affinity for all of the Chinese study material, and most of this material is formally transferred to Fuxianospira Chen and Zhou, 1997, a taxon common in the Chengjiang biota. Comparable material from the Cambrian Marjum, Wheeler, and Burgess Shale formations of North America is also assigned to Fuxianospira, and this reassignment expands both the paleogeographic and stratigraphic range of this taxon. All aspects of the study specimens, including details obtained from scanning electron microscopy, are consistent with an algal affinity, as proposed in the original descriptions of the Chinese material.

Lagerstätten from the Precambrian–Cambrian transition have traditionally been a relatively untapped resource for understanding the paleoecology of the “Cambrian explosion.” This quantitative paleoecological study is based on 10,238 fossil specimens belonging to 100 animal species, 11 phyla, and 15 ecological categories from the lower Cambrian (Series 2, Stage 3) Chengjiang biota (Mafang locality near Haikou, Yunnan Province, China). Fossils were systematically collected within a 2.5-meter-thick sequence divided into ten stratigraphic intervals. Each interval represents an induced time-averaged assemblage of various event (obrution) beds of unknown duration. Overall, the different fossil assemblages are taxonomically and ecologically similar, suggesting the presence of a single community type recurring throughout the Mafang section. The Mafang community is dominated by epibenthic vagile hunters or scavengers, sessile suspension feeders, and infaunal vagile hunters or scavengers represented primarily by arthropods, brachiopods, and priapulids, respectively. Most species have low abundance and low occurrence frequencies, whereas a few species are numerically abundant and occur frequently. Overall, in structure and ecology the Mafang community is comparable to the Middle Cambrian (Series 3, Stage 5) Burgess Shale biota (Walcott Quarry, Yoho National Park, British Columbia, Canada). This suggests that, despite variations in species identity within taxonomic and ecological groups, the structure and ecology of Cambrian Burgess Shale-type communities remained relatively stable until at least the Middle Cambrian (Series 3, Stage 5) in subtidal to relatively deep-water offshore settings in siliciclastic soft-substrate environments.

The Tulip Beds locality on Mount Stephen (Yoho National Park, British Columbia) yields one of the most abundant and diverse (~10,000 specimens in 110 taxa) Burgess Shale fossil assemblages in the Canadian Rockies. Detailed semi quantitative and quantitative analyses of this assemblage suggest strong similarities with the Walcott Quarry on Fossil Ridge. Both assemblages are dominated by epibenthic, sessile, and suspension feeding taxa, mostly represented by arthropods and sponges and have comparable diversity patterns, despite sharing only about half the genera. However, the Tulip Beds has a higher relative abundance of suspension feeders and taxa of unknown affinity compared to the Walcott Quarry. These biotic variations are probably largely attributable to ecological and evolutionary differences between the two temporally distinct communities that adapted to similar, but not identical, environmental settings. For instance, the Tulip Beds is farther away from the Cathedral Escarpment than the Walcott Quarry. The Tulip Beds and Walcott Quarry assemblages are more similar to each other than either one is to the assemblages of the Chengjiang biota, although the relative diversity of major taxonomic groups and ecological patterns are similar in all assemblages. The conserved diversity patterns and ecological structures among sites suggest that the ecological composition of Cambrian Burgess Shale-type communities was relatively stable across wide geographic and temporal scales.

The morphology of Tuzoia is reinterpreted in the light of abundant new specimens from the Middle Cambrian Burgess Shale (British Columbia, Canada) and Kaili (Guizhou, China) Lagerstätten. Tuzoia was a very large (up to 180 mm long) bivalved arthropod with a nonmineralized domelike carapace strengthened by prominent pointed features and often flanked by a lateral ridge bearing a spiny frill. The reticulate pattern of Tuzoia is comparable with that of present-day crustaceans (e.g., myodocope ostracods) and is interpreted as a structural compromise between exoskeletal lightness and high resistance to mechanical stress. Tuzoia had a pair of large, stalked, spherical, possibly compound eyes facing forward. Flagella-like antennae protruded through the anterior notch. No other appendages are known except possible filamentous setae underlying the carapace. Tuzoia typically occurs as laterally (lc) or dorsoventrally (dvc) compacted carapaces or single valves. Each type (lc or dvc) emphasizes particular aspects of the morphology (e.g., spiny lateral ridge, ventral margin) that were often interpreted as specific differences by previous authors. A revision of Tuzoia validates only 7 of the 23 named species. Tuzoia is placed tentatively within a group of large bivalved arthropods along with Isoxys and the possible ancestors of Thylacocephala (Lower Cambrian–Upper Cretaceous). In the Middle Cambrian, Tuzoia occurs across Laurentia, South and North China, and the Perigondwanan area (Bohemia) within a relatively narrow subtropical belt, indicating a high dispersal capability and possible latitudinal control on its distribution. Functional morphology, taphonomy, and the distributional pattern indicate that Tuzoia was a free-swimming arthropod.

Decay experiments are becoming a more widespread tool in evaluating the fidelity of the fossil record. Character interpretations of fossil specimens stand to benefit from an understanding of how decay can result in changes in morphology and, potentially, total character loss. We performed a decay experiment for the Class Enteropneusta to test the validity of anatomical interpretations of the Burgess Shale enteropneust Spartobranchus tenuis and to determine how the preservation of morphological features compares with the sequence of character decay in extant analogues. We used three species of enteropneust (Saccoglossus pusillus, Harrimania planktophilus, and Balanoglossus occidentalis) representing the two major families of Enteropneusta. Comparisons between decay sequences suggest that morphological characters decay in a consistent and predictable manner within Enteropneusta, and do not support the hypothesis of stemward slippage. The gill bars and nuchal skeleton were the most decay resistant, whereas the gill pores and pre-oral ciliary organ were unequivocally the most decay prone. Decay patterns support the identification of the nuchal skeleton, gill bars, esophageal organ, trunk, and proboscis in Spartobranchus tenuis and corroborate a harrimaniid affinity. Bias due to the taphonomic loss of taxonomically informative characters is unlikely. The morphologically simple harrimaniid body plan can be seen, therefore, to be plesiomorphic within the enteropneusts. Discrepancies between the sequence of decay in a laboratory setting and fossil preservation also exist. These discrepancies are highlighted not to discredit the use of modern decay studies but rather to underline their non-actualistic nature. Paleoenvironmental variables besides decay, such as the timeframe between death and early diagenesis as well as postmortem transport, are discussed relative to decay data. These experiments reinforce the strength of a comprehensive understanding of decay sequences as a benchmark against which to describe fossil taxa and understand the conditions leading to fossilization.

To investigate the phylogenetic affinity of Yuknessia simplex Walcott, 1919, scanning electron microscopy was applied to the Burgess Shale (Cambrian Series 3, Stage 5) type material and to new material from the Trilobite Beds (Yoho National Park) and specimens from the Cambrian of Utah. On the basis of fine-scale details observed using this approach, including banding structure interpreted as fusellae, Yuknessia Walcott, 1919 is transferred from the algae, where it resided for nearly a century, to the extant taxon Pterobranchia (Phylum Hemichordata). Considered as such, Yuknessia specimens from the Trilobite Beds and Spence Formation (Utah) are amongst the oldest known colonial pterobranchs. Two morphs regarded herein as two different species are recognized from the Trilobite Beds based on tubarium morphology. Yuknessia simplex has slender erect tubes whereas Yuknessia stephenensis n. sp., which is also known in Utah, has more robust erect tubes. The two paratypes of Y. simplex designated by Walcott (1919) are formally removed from Yuknessia and are reinterpreted respectively as an indeterminate alga and Dalyia racemata Walcott, 1919, a putative red alga.

Although Banffia constricta was first described in 1911, the affinities of this soft-bodied fossil animal have remained unclear. Banffia is redescribed here, on the basis of over 300 specimens. Banffia has a bipartite body measuring up to 10 cm in length. The entire body is axially twisted along a spiral, clockwise as seen from the front. The anterior section consists of two fused carapace-like units with a prominent crown-like structure at the anterior end surrounding the mouth. This crown-like structure is composed of three concentric circlets. A single antenniform outgrowth lies posterior to the mouth. The posterior section of Banffia is composed of 40 to 50 lightly sclerotised segments. The anus is terminal, and situated in a caudal notch. The gut is straight. Simple serially repeated structures along the gut may represent metameric mid-gut diverticulae. Banffia is considered to have been an epibenthic gregarious animal and possibly a deposit-feeder.

Banffia is putatively classified as an end-member of the phylum Vetulicolia, in a new class, the Banffozoa. If the vetulicolids represent a stem-group deuterostome, the absence of gills and a possible endostyle in Banffia is problematic. The presence of mid-gut diverticulae in Banffia may represent a link with the protostomes.