New brachiopod species from the order acrotretida from the Cambrian of central Kazakhstan

Four new brachiopod species from the order Acrotretida (class Lingulata): Picnotreta saryarkensis sp. nov., Stilpnotreta propria sp. nov., Anabolotreta firma sp. nov., and Batenevotreta variabilis sp. nov. are described from the Agyrek Mountains, northeastern Central Kazakhstan from two olistoliths of limestones, which contain the uppermost Middle and lowermost Upper Cambrian fauna and are located in the Upper Ordovician olistostrome. These new species supplement the described earlier brachiopod assemblage from this region (Koneva and Ushatinskaya, 2008). Strong age and intraspecific variability of some Kazakh acrotretids is shown.     

Predation of late Marjuman (Cambrian) linguliformean brachiopods from the Deadwood Formation of South Dakota, USA

One hundred and twenty-five linguliformean brachiopods of late Marjuman (Cambrian) age with shell perforations, presumably caused by predation, were recovered from shallow-water limestones at two localities of the Deadwood Formation in the Black Hills of South Dakota, USA. Three-quarters of the perforated valves belonged to taxa in the order Acrotretida, while one-quarter of the specimens belonged to those of the order Lingulida. This is the first report of predation of fossil lingulids. In both orders there was a marked selection for valve type. Ninety-five per cent of all perforations of acrotretids were in the ventral valve, while 87% of all perforations of lingulids were in the dorsal valve. The highest rate of predation of collected acrotretids, at any stratigraphic horizon, was 22%, while the rate of predation of collected lingulids, at any given stratigraphic horizon, was as high as 9%. Half of the perforated valves had round holes with a sharp, non-beveled exterior edge, and half had irregularly shaped holes with chipped interior edges. The former type is attributed to either boring or a swift, piercing percussive strike, while the latter is attributed to a smashing percussive strike with a blunt appendage. A third type of perforation consisting of minute, roughly circular holes is thought to be too small to be the work of predators, and is assumed to be the result of an encrusting organism or parasite. The different types of perforation seen in the brachiopod valves indicate that there was more than one type of predator attacking them, including possibly one of the earliest durophages. Various hypothetical predators are suggested as potential candidates for causing the shell perforations. The criteria for their selection as possible linguliformean predators include possessing the ability to produce one of the two types of shell perforation, and being small enough to warrant preying on small (< 2 mm) brachiopods.     

Cambrian-Ordovician lingulate brachiopods from Scandinavia, Kazakhstan, and South Ural Mountains

Lingulate brachiopods are described from the Upper Cambrian - Lower Ordovician (Tremadoc-Arenig) of Scandinavia (Sweden, Denmark, and Norway), South Ural Mountains, northeastern Central Kazakhstan, and the southern Kendyktas Range in southern Kazakhstan. The faunas comprise a total of 56 species of which 20 are new these are assigned to 40 genera, of which the lingulids Agalatassia and Keskentassia , the siphonotretid Siphonotrerella. and the acrotretids Galinella, Longipegma, Ottenbyella, Akmolina, Mamatia, Sasyksoria , and Otariella are new. The new Subfamily Elliptoglossinae is proposed. The poorly known Cambrianardovician stratigraphy of the South Urals. northeastern Central Kazakhstan, and the southern Kendyktas Range is reviewed. Many sequences in these areas that were previously referred to the Upper Cambrian and Tremadoc can now be correlated with the lower Arenig Hunneberg Stage in Baltoscandia. Three main types of faunal assemblages can be distinguished: (1) the Broeggeria assemblage; (2) several microbrachiopod assemblages; and (3) the Leptembolon-Thysanotos assemblage. The Broeggeria assemblage is distributed world-wide in the Tremadoc of the southern Kendyktas Range, Scandinavia, Belgium, Great Britain, Canada, and Argentina, while the Leptembolon-Thysanotos assemblage is confined to the Arenig of an area surrounding the East European platform, including northern Estonia, Poland, Germany, Bohemia, Serbia, and the South Urals. The microbrachiopod assemblages are known mainly from the Upper Cambrian - Arenig of Scandinavia, South Ural Mountains, northeastern Central Kazakhstan, and the southern Kendyktas Range.     

河北完县中、晚寒武世无铰纲腕足类

研究了河北完县清醒剖面中寒武统张夏组至上寒武统凤山组酸解法所获得的无铰纲腕足类化石5500多件,详细描述了乳孔贝科2新属、16新种、3未定种,圆货贝科1新属、2新种.自张夏组至凤山组建立了8个无铰纲腕足类化石组合.对乳孔贝亚科的幼壳形态进行了分类,并研究幼壳的演化趋向.     

The Cambrian radiation of bilaterians: Evolutionary origins and palaeontological emergence; earth history change and biotic factors

Evidence from a variety of research areas, including phylogenetic palaeobiogeographic studies of trilobites, indicates that there may be a fuse to the Cambrian radiation, with a duration on the order of 20–70 myr. Evolution in trilobites appears to have been powerfully influenced by the tectonic changes occurring at the end of the Neoproterozoic: especially the breakup of Pannotia. This continental fragmentation may have also elevated opportunities for vicariance and speciation in trilobites, and other metazoans, given that speciation rates at this time period were high, though not phenomenally so. This provides clear evidence that abiotic factors played an important role in motivating evolution during this key episode in the history of life; biotic factors probably also played a role. The evidence for the role of biotic factors is considered in light of information from some problematic Cambrian taxa. These may show affinities with modern problematic pseudocoelomate phyla, although Cambrian and modern exponents differ dramatically in body size.     

黔东早—中寒武世凯里组疑源类组合及其界线意义

通过对台江八郎和丹寨两条剖面凯里组中疑源类化石的分析,研究,台江剖面产疑源类化石有17属35各（12未定种）,其中以Leiosphaeridia,Synsphareidium,Cymatiosphaera,Pterospermella最为丰富,可划分为3个组合,即Cymatiosphaera cf.cristata-Fimbriaglomerella memebrancea组合,Cristallinium-Micrhystridium-Pterospermella组合和Dictyotidium-Granomarginata组合,丹寨平寨剖面凯里组所产疑源类化石有：13属21种（5未定种）,同样划分为3个组合,即Leiosphaeridia-Tasmanites组合,Retisphaeridium-Micrhystridium tentatium组合和Baltisphaeridium-Bubomorpha hunjiangensis组合,本文对两条剖面中的凝源类化石的组成,相对含量做了详细的统计,发现在八郎剖面9－2层和平寨剖面3层疑源类的丰度,分异度开始发生明显的变化,表现由早寒武世向中寒武世疑源类组合面貌的转变,这一疑源类转变层位正好是与三叶虫划分的中,下寒武统界线的层位位置相一致,这充分表明疑源类化石可作为划分中,下寒武统的极有价值的微体生物化石证据。     

Microreticulation in Cambrian Ostracoda and its relation to similar patterns in tertiary and Recent Ostracoda

The Cambrian bradoriid ? Parahoulongdongella sp. is shown to have a microreticulate (2nd order) surface sculpture. A possible organic sheet‐like origin is proposed for this reticulation. The relation between this second order pattern and those found in tertiary to Recent Ostracoda is discussed.     

A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia

Two Cambrian lepidote metazoans known from different respective types of preservation have been compared in order to elucidate their biology and affinities. The widely distributed Lower Cambrian Halkieria is represented by isolated hollow sclerites, probably of originally calcareous composition. The Middle Cambrian Wiwaxia is known from the Burgess Shale as isolated sclerites (scales and spines) and as more or less complete individuals. Although Halkieria sclerites were mineralized and those of Wiwaxia were probably not, there are fundamental structural and morphological similarities between the two. Both bad an imbricating scaly and spiny armour consisting of hollow sclerites with a longitudinally fibrous structure. The sclerites did not grow, but were probably moulted during the course of ontogenetic growth. Halkieria and Wiwaxia are regarded as closely related. Both are referred to the Order Sachitida He 1980. The sclerite armour of Halkieria is reconstructed on the template provided by Wiwaxia. The interpretation of sachitid sclerites as protective armour is an alternative to the interpretation by Jell (1981, Alcheringa 5 )that sachitid sclerites were respiratory organs in an animal of probable annelid affinities. Sachitids are interpreted as sluggish, benthic deposit feeders that do not belong to any recognized phylum.     

THE LOWER CAMBRIAN FOSSIL TOMMOTIA

The genera Tommotia Missarevskij 1970 (1966) and Camenella Missarevskij 1966 from the Lower Cambrian Tommotian Stage of the Siberian Platform represent different skeletal parts of the same organism and are consequently synonymous. Tommotia is selected as the valid name. Tommotia baltica n.sp. is described from the Lower Cambrian zone of Mobergella holsti of the Kalmarsund area in the southern part of the Baltic Sea. The Lower Cambrian families Tommotiidae Missarzevskij 1970 (1969) and Tannuolinidae Fonin & Smirnova 1967 show important similarities in their basic structure and are brought together in the new order Mitrosagophora, characterized by a composite outer skeleton with two types of skeletal elements, termed 'mitral' and 'sellate', each occurring in symmetrical pairs. The group is of uncertain systematic position but may have affinities with the Machaeridia.
The Scandinavian zone of Mobergella holsti is correlated with the Siberian zone of Dokidocyathus lenaicus and Majatheca tumefacta , the uppermost zone of the Tommotian Stage.     

Early Cambrian radiolarians and sponge spicules from the Niujiaohe Formation in South China

Most of the reported Cambrian radiolarians are from middle Cambrian (Miaolingian Series, Wuliuan Stage) and onward, the radiolarians from lower Cambrian (Terreneuvian and Cambrian Series 2), on the other hand, are poorly documented, thus their morphological characteristics have not been well understood. In this study, we extracted spherical radiolarians and sponge spicules from the chert of the Cambrian Series 2 Niujiaohe Formation in southern Jiangxi Province, China. The well-preserved radiolarian fossils, identified as Paraantygopora porosa, consist of perforate plate shells pierced by dense pores with elevated rims, and display similarities to those from Lower Ordovician. Seven forms of sponge spicules are recognized, including monaxons, diaxons, tetractines, hexactines, triaxon tetractines, triaxon pentactines and irregular tetraxon tetractines, which are similar to those from the Qiongzhusian Stage in South China. Our results, combined with previously reported early Cambrian radiolarians from South China, indicate that the early Cambrian radiolarians developed advanced spherical skeletons.     

Middle and Upper Cambrian Protoconodonts and Paraconodonts from Hunan, South China

More than 4200 kg of limestone, representing 980 productive samples, from the upper Middle Cambrian, Upper Cambrian, and lowermost Ordovician in western and north-western Hunan were processed for paraconodonts, protoconodonts, and euconodonts. The focus of the present paper is on the upper Middle and lower Upper Cambrian interval as it is developed at two, apparently stratigraphically continuous, key sections. The collections studied, which include more than 20,000 specimens and are quite diverse taxonomically, prove that some protoconodonts and paraconodonts are useful biostratigraphically. Previously proposed protoconodont-paraconodont biozones in Hunan are revised and correlated with recently revised trilobite biozones, as well as with protoconodont-paraconodont and trilobite biozones in North China. Twenty-six species and seven conditionally identified species belonging to 13 genera are described. Among these, two genera ( Huayuanodontus and Yongshunella ) and two species ( Westergaardodina elegans and Yongshunella polymorpha ) are new. This study has special interest for Cambrian biostratigraphy because the two studied sections are global stratotype candidates for the Middle-Upper Cambrian series boundary. The level of the Upper-Middle Cambrian Series boundary, as currently recognized in China (at the base of the Linguagnostus reconditus Trilobite Biozone) is well marked in the paraconodont succession and can be traced into the Swedish standard succession using these fossils. By contrast, another level recently proposed as a potential global Middle-Upper Cambrian Series boundary level, the base of the Glyptagnostus reticulatus Trilobite Biozone, does not coincide with any marked change in the Hunan conodont species succession useful for local and regional correlation.     

Lapworthellids and other skeletonised microfossils from the Cambrian Stage 3 of the northern Montagne Noire,southern France

The Montagne Noire (southern France) possesses one of the most complete Cambrian successions in the western peri-Gondwana margin and might provide a good stratigraphic reference for both regional charts and international correlations. However, to date, the lower Cambrian succession of the northern Montagne Noire has been supposed to be devoid of biostratigraphically significant fossils. The complex tectonostratigraphic framework of the area (a range divided into Axial Zone, northern and southern Montagne Noire) exacerbated problems related to regional correlations and palaeogeographic reconstructions. As a result, the chronostratigraphic context of the lower Cambrian of northern Montagne Noire is still uncertain and stratigraphic reports have broadly relied on putative lithostratigraphic correlations with the southern Montagne Noire. The purpose of this study is to characterise, for the first time, the fossil record of carbonate beds and lenses of the northern Montagne Noire occurring at the top of the siliciclastic-dominated Marcory Formation, in order to provide regional bio- and chronostratigraphic constraints on lower Cambrian strata. Moreover, this study is aimed at improving international chronostratigraphic correlation. Carbonate beds and lenses cropping out along the Orque Cliff, in the Mélagues thrust slice, were investigated. They yielded a faunal assemblage constituted of molluscs (Igorella cf. ungulata and Igorella moncereti n. sp.), hyoliths (Conotheca brevica), chancelloriids (Archiasterella cf. pentactina and Allonnia sp.) and tommotiids (Lapworthella rete). L. rete is recorded in upper Meishucunian (Cambrian Stage 3) strata of the Yangtze Platform (South China) where it is used as index fossil of the Cambrian Stage 3 Sinosachites flabelliformis–Tannuolina zhangwentangi Assemblage Zone. Therefore, the presence of this tommotiid provides evidence that the studied carbonate beds and lenses are Cambrian Age 3 (Atdabanian according to the Siberian chart). The upper part of the Marcory Formation in the Mélagues slice, dated as Cambrian Stage 3, might represent a lateral equivalent of the mixed (carbonate-siliciclastic) Pardailhan Formation defined in the southern Montagne Noire.     

Tardigrades as ‘Stem-Group Arthropods’: The Evidence from the Cambrian Fauna

The Cambrian fauna can now reasonably be seen as containing many taxa that lie in the stem-groups of the extant phyla. As such, these fossils suggest how both the ‘body plans’ of extant phyla were assembled, and also how various ‘minor’ phyla relate to the larger groupings of today such as the arthropods and annelids.

The various arthropod and lobopod taxa of the Cambrian faunas have been controversial and have generally been considered either as lying in the crown or (occasionally) stem groups of the euarthropods, onychophorans and tardigrades. However, phylogenetic analysis strongly suggests that many of even the most euarthropod-like taxa do not lie within the euarthropod crown-group but are more basal. Further, the commonly expressed view that Cambrian lobopods are in effect stem- or crown-group onychophorans also seems not to be well supported. Lobopods in the Cambrian appear to be diverse and not particularly closely related to one another, and certainly cannot be combined in a monophyletic clade.

Both these advances offer hope that the tardigrades (placed as the sister group to the euarthropods in many analyses of extant taxa, here collectively named the Tactopoda) may be more closely related to some of these Cambrian taxa than others. The challenge for both neontologists and palaeontologists is to refine the systematic analysis of both living and fossil taxa in order to maximise the usefulness of the (admittedly few) characters that unite tardigrades to their Cambrian forbears.     

A New Algal Fossil from Early Cambrian in Qingzhen County,Guizhou Province,China

A new algal fossil was collected from the lower part of the Lower Cambrian, a Sinian-Cambrian boundary section in Abazhai of Qingzhen County, Guizhou Province. It was discovered in petrographic thin sections of a black laminated chert. The new fossil, Symphysosphaera radialis gen. et sp. nov. is a fossilized muhicellular colony characterized by a surface layer of cells enclosed in an envelope, inner cavity permeated with dark dense mucilage embedded with numerous spherical units. Its difference from all morphs discovered in Precambrian, indicates that the genus may represent member of a new order at the beginning of Cambrian. hs stratigraphic and biological significance merits our attantion.     

Reappraising the early evidence of durophagy and drilling predation in the fossil record: implications for escalation and the Cambrian Explosion

The Cambrian Explosion is arguably the most extreme example of a biological radiation preserved in the fossil record, and studies of Cambrian Lagerstätten have facilitated the exploration of many facets of this key evolutionary event. As predation was a major ecological driver behind the Explosion – particularly the radiation of biomineralising metazoans – the evidence for shell crushing (durophagy), drilling and puncturing predation in the Cambrian (and possibly the Ediacaran) is considered. Examples of durophagous predation on biomineralised taxa other than trilobites are apparently rare, reflecting predator preference, taphonomic and sampling biases, or simply lack of documentation. The oldest known example of durophagy is shell damage on the problematic taxon Mobergella holsti from the early Cambrian (possibly Terreneuvian) of Sweden. Using functional morphology to identify (or perhaps misidentify) durophagous predators is discussed, with emphasis on the toolkit used by Cambrian arthropods, specifically the radiodontan oral cone and the frontal and gnathobasic appendages of various taxa. Records of drill holes and possible puncture holes in Cambrian shells are mostly on brachiopods, but the lack of prey diversity may represent either a true biological signal or a result of various biases. The oldest drilled Cambrian shells occur in a variety of Terreneuvian‐aged taxa, but specimens of the ubiquitous Ediacaran shelly fossil Cloudina also show putative drilling traces. Knowledge on Cambrian shell drillers is sorely lacking and there is little evidence or consensus concerning the taxonomic groups that made the holes, which often leads to the suggestion of an unknown ‘soft bodied driller’. Useful methodologies for deciphering the identities and capabilities of shell drillers are outlined. Evidence for puncture holes in Cambrian shelly taxa is rare. Such holes are more jagged than drill holes and possibly made by a Cambrian ‘puncher’. The Cambrian arthropod Yohoia may have used its frontal appendages in a jack‐knifing manner, similar to Recent stomatopod crustaceans, to strike and puncture shells rapidly. Finally, Cambrian durophagous and shell‐drilling predation is considered in the context of escalation – an evolutionary process that, amongst other scenarios, involves predators (and other ‘enemies’) as the predominant agents of natural selection. The rapid increase in diversity and abundance of biomineralised shells during the early Cambrian is often attributed to escalation: enemies placed selective pressure on prey, forcing phenotypic responses in prey and, by extension, in predator groups over time. Unfortunately, few case studies illustrate long‐term patterns in shelly fossil morphologies that may reflect the influence of predation throughout the Cambrian. More studies on phenotypic change in hard‐shelled lineages are needed to convincingly illustrate escalation and the responses of prey during the Cambrian.     

Traces of predation in the Cambrian

Series two marks a revolution in Cambrian predation when new predators and new predation methods appeared, which led to general increase in predation intensities and in the diversity of prey groups. The number of bored taxa and taxa with the predation scars is similar in the Cambrian. Most of the borings are associated with brachiopods and most of the scars with trilobites. Brachiopods, arthropods, molluscs, cnidarians and echinoderms were the most common prey in the Cambrian. The Cambrian record of predation is dominated by damage inflicted on brachiopods and trilobites. The fossils with predation signs are known from a majority of paleocontinents and all the Cambrian series.     

A revision of Holm's Early and early Mid Cambrian hyoliths Of Sweden

ABSTRACT. Re-examination of type specimens of Early Cambrian and early Mid Cambrian hyoliths from Sweden confirms placement of Hyolithes teretiusculus Linnarsson in Hexitheca Syssoiev, and reassignment to the order Hyolithida rather than Orthothecida. Inclusion of Hyolithes affinis Holm within Decoritheca Syssoiev is re-confirmed, and Hyolithes socialis Linnarsson is now referred to Nevadotheca Malinky and Slapylites Marek. An operculum included under Hyolithes oelandicus Holm is transferred provisionally to Maxilites Marek, and conchs of that species are placed under Tulenicornus Val'kov with question. No other hyolith species of Holm can be definitely assigned to genus. The rarity of well preserved opercula is a major impediment to recognizing hyolith taxa confidently, and we follow Holm in attributing the comparative rarity of opercula to the fact that in some hyolith taxa the operculum may have been composed of a different, and perhaps less stable, material than the rest of the conch.     

A review of the Cambrian biostratigraphy of South Australia

Cambrian rocks in South Australia occur in the Stansbury, Arrowie, eastern Officer and Warburton Basins. The succession in the Stansbury and Arrowie Basins can be divided into three sequence sets (supersequences), 1, 2 and 3. Sequence set 1 can be divided into five third-order sequences: 1.0, 1.1A, 1.1B, 1.2 and 1.3. Trilobites from the Stansbury and Arrowie Basins are restricted largely to the lower part of the succession. Four trilobite zones are recognized: Abadiella huoi (latest Atdabanian–earliest Botoman), Pararaia tatei, Pararaia bunyerooensis and Pararaia janeae Zones (all Botoman). Trilobites higher in the succession are known from only a few horizons and in part correlate with the upper Lower Cambrian Lungwangmiaoan Stage of China, equivalent to the top Toyonian. Pagetia sp. has been reported in the Coobowie Formation of the Stansbury Basin, thus suggesting an early Middle Cambrian age.The Cambrian faunas of the Warburton Basin range in age from early Middle Cambrian (Late Templetonian) to very Late Cambrian, although the richest faunal assemblages are late Middle Cambrian (Ptychagnostus punctuosus to Goniagnostus nathorsti Zones). Conodonts, including Cordylodus proavus, occur in a Datsonian fauna.The Arrowie Basin contains the most complete and best studied archaeocyath succession in the Australia–Antarctica region. The Warriootacyathus wilkawillensis, Spirillicyathus tenuis and Jugalicyathus tardus Zones from the lower Wilkawillina Limestone (Arrowie Basin) and equivalents are correlated with the Atdabanian. Botoman archaeocyathids occur higher in the Wilkawillina Limestone. The youngest (Toyonian) archaeocyath fauna in Australia occurs in the Wirrealpa Limestone (Arrowie Basin).Brachiopods and molluscs of the Arrowie and Stansbury Basins can be divided into four biostratigraphic assemblages. Several informal Early Cambrian SSF biostratigraphic assemblages are recognized. Probable tabulate-like corals occur in the Botoman Moorowie Formation. Seven informal acritarch assemblages occur in the Early Cambrian of the Stansbury and Arrowie Basins. Trace fossils may mark the Precambrian–Cambrian boundary. Only two of several tuffaceous horizons from the Stansbury and Arrowie Basins have been dated (i) a date of 522.0 ± 2.1 Ma from the Heatherdale Shale of the Stansbury Basin, about 400 m above latest Atdabanian archaeocyathids and (ii) a date of 522.0 ± 1.8 Ma from the lower part of the Billy Creek Formation in the Arrowie Basin. Neither date is regarded as reliable.     

Post-embryonic development of the Furongian (late Cambrian) trilobite Tsinania canens: implications for life mode and phylogeny

SUMMARY The current concept of the order Asaphida was proposed to accommodate some Cambrian and Ordovician trilobite clades that are characterized by the possession of a ventral median suture. The family Tsinaniidae was recently suggested to be a member of the order Asaphida on the basis of its close morphological similarity to Asaphidae. Postembryonic development of the tsinaniid trilobite, Tsinania canens , from the Furongian (late Cambrian) Hwajeol Formation of Korea, reveals that this trilobite had an adult-like protaspis. Notable morphological changes with growth comprise the effacement of dorsal furrows, sudden degeneration of pygidial spines, regression of genal spines, and loss of a triangular rostral plate to form a ventral median suture. Programmed cell death may be responsible for degenerating the pygidial and genal spines during ontogeny. Morphological changes with growth, such as the loss of pygidial spines, modification of pleural tips, and effacement of dorsal furrows, suggest that T. canens changed its life mode during ontogeny from benthic crawling to infaunal. The protaspid morphology and the immature morphology of T. canens retaining genal and pygidial spines suggest that tsinaniids bear a close affinity to leiostegioids of the order Corynexochida. Accordingly, development of a ventral median suture in T. canens demonstrates that the ventral median suture could have evolved polyphyletically, and thus the current concept of the order Asaphida needs to be revised.     

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.