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.     

华北寒武系第二统辛集组软舌螺动物化石群*

软舌螺动物是华北板块寒武系小壳化石基本组成之一,也是寒武纪生命大爆发在华北地区的重要见证者。本文系统描述了华北板块南缘-西南缘(河南-陕西地区)寒武系第二统辛集组软舌螺化石9属14种及1未定种。包括直管螺类6属9种:Conotheca australiensis、Cupitheca holocyclata、Cupitheca costellata、Cupitheca? decollata、Tegminites hymenodes、Triplicatella disdoma、Triplicatella xinjia、Protomicrocornus triplicensis和Longxiantheca mira;软舌螺类3属5种及1未定种:Microcornus eximius、Microcornus petilus、Parkula cf. esmeraldina、Parkula bounites、Parakorilithes mammillatus和Parakorilithes sp.。基于大量保存的精美聚合标本,本文准确重建了锥壳和口盖在软舌螺不同属种中的配置。此外,新材料显示华北辛...     

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale

Pettersson Stolk, S., Holmer, L. E. and Caron, J ‐B. 2010. First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale. —Acta Zoologica (Stockholm) 91 : 150–162 The organophosphatic shells of linguloid brachiopods are a common component of normal Cambrian–Ordovician shelly assemblages. Preservation of linguloid soft‐part anatomy, however, is extremely rare, and restricted to a few species in Lower Cambrian Konservat Lagerstätten. Such remarkable occurrences provide unique insights into the biology and ecology of early linguloids that are not available from the study of shells alone. Based on its shells, Lingulella waptaensis Walcott, was originally described in 1924 from the Middle Cambrian Burgess Shale but despite the widespread occurrence of soft‐part preservation associated with fossils from the same levels, no preserved soft parts have been reported. Lingulella waptaensis is restudied herein based on 396 specimens collected by Royal Ontario Museum field parties from the Greater Phyllopod Bed (Walcott Quarry Shale Member, British Columbia). The new specimens, including three with exceptional preservation of the pedicle, were collected in situ in discrete obrution beds. Census counts show that L. waptaensis is rare but recurrent in the Greater Phyllopod Bed, suggesting that this species might have been generalist. The wrinkled pedicle protruded posteriorly between the valves, was composed of a central coelomic space, and was slender and flexible enough to be tightly folded, suggesting a thin chitinous cuticle and underlying muscular layers. The nearly circular shell and the long, slender and highly flexible pedicle suggest that L. waptaensis lived epifaunally, probably attached to the substrate. Vertical cross‐sections of the shells show that L. waptaensis possessed a virgose secondary layer, which has previously only been known from Devonian to Recent members of the Family Lingulidae.     

Small shelly fossils and trace fossils near the Precambrian-Cambrian boundary in the Yukon Territory, Canada

In the past an ‘explosion’ in diversity and abundance of small shelly fossils and of trace fossils has served to mark the base of the Cambrian. However, no evidence has been presented to prove that the ‘explosions’ of the two groups were synchronous. We describe small shelly fossils and trace fossils from the same phosphatic limestone beds that indicate that the two events were separate in time. The small shelly fossils are Anabarites trisulcatus, Hyolithellus cf. H. isiticus, Microcornus? sp., Protohertzina anabarica, P. unguliformis, P. sp. A, Pseudorthotheca sp. A, Rushtonia? sp. A, four types of tuberculate plates and one type of reticulate plate. These fossils represent a restricted, ‘pre-explosion’ fauna and are assigned to the Anabarites-Circotheca-Protohertzina Assemblage Zone, an uppermost Precambrian zone in the Meishucun Stage, Yunnan Province, China. A point at the top of this zone has received strong international endorsement for future designation as the base of the Cambrian. Associated with the small shelly fossils are the trace fossils Cruziana sp. A, Cruziana? sp. B, Rusophycus sp. A, Palaeophycus rubdark and arthropod scratch marks. If found in isolation, this trace fossil assemblage would be considered as post-Precambrian because it includes large, highly organized arthropod traces that are traditionally accepted as occurring above the trace fossil ‘explosion’. We therefore conclude that the trace fossil ‘explosion’ predates the small shelly fossil ‘explosion’. If the proposed location of the base of the Cambrian in Yunnan is accepted, the small shelly fossil ‘explosion’ concept and its relationship to the boundary would not be greatly modified. The trace fossil ‘explosion’, however, would no longer indicate the base of the Cambrian and the ranges of some trace fossils would be extended into the Precambrian.     

Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs

Runnegar, Bruce 1981 12 15: Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs. Lethaia, Vol. 14, pp. 311–322. Oslo. ISSN 0024–1164. Well preserved muscle scars have been discovered on one or more specimens of the early Palaeozoic univalved molluscs Pelagiella, Matherella, Sinuites and ?Bucania. From this information it is argued that the bellerophonts Sinuites and ?Bucania were untorted, that Pelagiella had undergone about 10° of torsion, and that Matherella may have been a fully torted, hyperstrophic descendant of Pelagiella. Other early gastropods such as Aldanella could have been independently derived from Pelagiella and therefore the Class Gastropoda may be at least diphyletic. In general, small dextrally coiled archaeogastropods such as Aldanella and the pelagiellids are the most common asymmetrical univalves of the Early Cambrian, but rare ultradextral (hyperstrophic) forms are also known. Only pelagiellids are known from the Middle Cambrian, and ultradextral forms dominate in the Late Cambrian. As a result, slit-bearing archaeogastropods are not common before the Ordovician. Yuwenia bentleyi gen. et sp. nov. is a new taxon proposed in this work. *Mollusca, Gastropoda, Monoplacophora, anatomy, evolution, Cambrian, Ordovician, Yuwenia n.g.     

A new mobergellan (small shelly fossils) from the early middle cambrian of morocco and its significance

A new mobergellan genus and species,Tateltella ranoculata, is described from the early Middle Cambrian (Agdzian Stage) of Morocco. The new taxon is characterized by only four pairs of muscle scars and is furthermore distinguished from other mobergellans by its strongly concave shape and its distally rising muscle scars. The individual specimens ofTateltella ranoculata distinctly vary in size and display different ontogenetic stages. Juvenile, intermediate, and adult stages can be distinguished by means of the development of the muscle scars that differ in Position relative to the apex, size, and distinctness between individual stages. The shell ofT. ranoculata is composed of a succession of thin phosphatic lamellae separated by interlamellar gaps, presumably originally filled by organic material. The interlamellar gaps may be divided by septum-like structures producing discrete cavities. The specimens are the youngest mobergellans known so far and correlation of their stratigraphic position suggests a correspondence with the lower part of the Amgan stage of the Siberian Platform. In addition, they are the first reported mobergellans from the present day continent Africa. Other mobergellan taxa and mobergellan-like species are briefly reviewed and the genusHippoklosma Missarzhevsky, previously assigned to the Mobergellidae, is excluded from the family due to its different shell structure. An evolutionary trend of reduction in the number of muscle scars from 14 in the early Early Cambrian to only eight in the early Middle Cambrian is apparent among mobergellans.      

陕南寒武系底部宽川铺组微体化石人工智能识别

我国陕南寒武纪初期的宽川铺组(距今大约535百万年)以盛产磷酸盐化的微体小壳化石和球状动物胚胎化石而闻名于世,是探索早期动物门类起源和寒武纪生命大爆发研究中的一个非常重要的窗口。在宽川铺组中寻找微体化石新类型主要依赖显微镜下人工挑选的传统方法日趋低效,迫切需要新的技术手段来解决这些问题。而微体化石数量庞大,大小接近,形态简单,非常适合于采用机器学习的手段进行人工智能分类。我们在宽川铺组微体化石中尝试使用机器学习的手段来进行化石图像识别和机器分拣。采用方向梯度直方图来提取化石图像的主要向量特征,并设计了二叉树型多分类识别器进行化石数字分类。目前处理了5 000多张微体化石照片的人工智能识别,已经取得了较高的准确率。     

Early Cambrian organophosphatic brachiopods from the Xinji Formation,at Shuiyu section,Shanxi Province,North China

Abundant and diverse small shelly fossils have been reported from Cambrian Series 2 in North China, but the co-occurring brachiopods are still poorly known. Herein, we describe seven genera, five species and two undetermined species of organophosphatic brachiopods including one new genus and new species from the lower Cambrian Xinji Formation at Shuiyu section, located on the southern margin of North China Platform. The brachiopod assemblage comprises one mickwitziid (stem group brachiopoda), Paramickwitzia boreussinaensis n. gen. n. sp., a paterinide, Askepasma toddense Laurie, 1986, an acrotretoid, Eohadrotreta cf. zhenbaensis Li and Holmer, 2004, a botsfordiid, Schizopholis yorkensis (Holmer and Ushatinskaya in Gravestock et al., 2001) and three linguloids, Spinobolus sp., Eodicellomus cf. elkaniiformis Holmer and Ushatinskaya in Gravestock et al., 2001 and Eoobolus sp. This brachiopod assemblage suggests a late Age 3 to early Age 4 for the Xinji Formation and reveals a remarkably strong connection with coeval faunas from East Gondwana, particularly the Hawker Group in South Australia. The high degree of similarity (even at species level) further supports a close palaeogeographic position between the North China Platform and Australian East Gondwana during the early Cambrian as indicated by small shelly fossil data.     

寒武纪早期岩家河生物群:研究进展和展望*

埃迪卡拉纪-寒武纪转换时期动物的起源、演化和"寒武纪大爆发"一直是国际古生物学界研究的热点问题,其中寒武纪早期小壳化石群与埃迪卡拉纪化石群和寒武纪早期澄江化石库之间的内在关系是古生物学界研究的难题,其主要原因是寒武纪早期与小壳化石群伴生的宏体动、植物化石的缺乏。发现于峡东地区的寒武纪早期岩家河生物群填补了这一缺失环节,该生物群包含宏体动物、宏观藻类、小壳化石、球形化石(可能的胚胎化石)、微古植物和蓝菌类等化石,部分宏体化石显示了从埃迪卡拉纪向寒武纪过渡色彩。化石保存方式有碳质膜、黄铁矿化、磷酸盐化、硅化。因此对岩家河生物群生物多样性和埋藏学进行综合研究,将可提供纽芬兰世(梅树村期)碳酸盐台地—碳酸盐台地内部的局部凹陷盆地相的一个较完整的生物景观图,对探索"寒武纪大爆发主幕"前夕生物的辐射、演化模式及保存机制具有重要的科学意义。     

New data on the enigmatic Ocruranus–Eohalobia group of Early Cambrian small skeletal fossils

Abstract: The Ocruranus–Eohalobia group, whose members were variously considered to be brachiopods, bivalves, chitons, tommotiids and coeloscleritophorans, are difficult to classify because of lack of morphological detail and evidence for skeletal reconstruction. New specimens from South China reveal more information about Ocruranus–Eohalobia and allow progress towards deciphering the skeletal reconstruction and phylogenetic affinity of this enigmatic group. Many specimens have a phosphatic inner and outer coat (mould) with empty space in between that resulted from dissolution of the original shell. Moreover, many of the internal moulds show a previously unknown type of shell microstructure that consisted of stacked layers of highly organized, acicular crystallites that radiated from the apex of the shell towards the aperture. The dissolved shell and needle‐like crystals suggest an original calcareous, probably aragonitic, shell mineralogy. A few specimens also show a polygonal texture in regions that suggests the shell had a thin, prismatic inner shell microstructure. Ocruranus and Eohalobia belong to the same skeleton, and we herein synonymize Eohalobia with the older Ocruranus. Moreover, new specimens from Meishucun reveal a third type of shell plate, similar in form and inferred placement to intermediate valves of chitons. Ocruranus is likely a mollusc, and possibly a member of the chiton stem lineage. If so, then the beginning of the known record of chitons would be extended back from late Cambrian (Saukia Zone; Furongian) to early Cambrian (Meishucunian; Series 1).     

Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China

The Chengjiang Lagerstätte in the Lower Cambrian of South China yields a small, larva‐like arthropod, which was considered to be a protaspis of naraoiids by many authors. The discovery of a large number of well‐preserved specimens from many new localities has allowed the original study to be revised. The relatively large size, stable morphology and unusual structure of the appendages indicate that these specimens represent adults of a new arthropod, Primicaris larvaformis. The larva‐like outline is considered to have arisen by the heterochronic process of progenesis. In addition, this animal displays primitive aspects of bodyplan and limb morphology that suggest a basal position within arachnomorphs, or perhaps even arthropods, and the similarities to the Vendian arthropod‐like animal Parvancorina probably provide an evolutionary link between Vendian forms and Cambrian arthropods.     

贵州寒武系第二统杷榔组有壳变形虫的新发现*

有壳变形虫(testate amoebae)的演化历史最早可追溯至新元古代早期, 以该时期北美、华北、挪威以及澳洲等多个地区浅海相碳酸盐岩、页岩中发现的瓶状微体化石(vase-shaped microfossils)为标志。此前认为, 显生宙的有壳变形虫最早出现在早泥盆世。长期以来, 早古生代的地层中未发现这类原生生物的明确化石证据。本研究通过对岩石样品进行常规孢粉酸泡分析处理和切磨岩石薄片, 获取原位保存的化石标本的技术方法, 从贵州东部剑河县交榜剖面出露的寒武系杷榔组(第2统第4阶)中获得数枚有壳变形虫(testate amoebae)化石标本。基于标本的显微形态特征, 并结合激光拉曼光谱等研究, 对原先记述为疑源类的Plagasphaera balangensis和P. sp. A两形态种进行重新认识和描述。由于它们在结构和形态上与一些现生的鳞壳虫目(Euglyphida)有壳变形虫极为相似, 因此将先前定为疑源类的Plagasphaera balangensis和P. sp. A两形态属、种名, 分别修订为Palaeoassulina balangensis gen. et sp. nov.和?Palaeoassulina sp. A。该发现不仅将显生宙有壳变形虫的原有化石记录从晚古生代泥盆纪向前延伸至寒武纪早期, 还为调查研究有壳变形虫的系统演化提供关键的生物化石证据。     

New data on the morphology and systematics of hyolithelminthes (Cambrian problematic organisms)

Phosphatic structures are discovered in the tube interior of the hyolithelminth species Hyolithellus vitricus from the Lower Cambrian of the Siberian Platform. Anatomic interpretation of these structures suggests that these small-sized shelly fossils represent the earliest worm-shaped organisms probably closely related to modern Nemathelminthes.     

Ecology and phylogenetic affinity of the early Cambrian tubular microfossil Megathrix longus

Abundant, exquisitely preserved specimens of the enigmatic tubular microfossil Megathrix longus are reported from the early Cambrian Yanjiahe Formation in the Yangtze Gorges area. Studies of their morphology and taphonomy reveal that the inner cross‐wall of Megathrix longus is topographically flat, the trichome terminus is blunt and closed, and incomplete cross‐walls are regularly intercalated between complete ones. The deformation of trichomes, as well as the corrugation of their cross‐walls, are postmortem features caused by compaction, rather than biological features as previously interpreted. Statistical analysis indicates that chamber division is accomplished by the closure of incomplete cross‐walls. This process, which contributed to trichome growth, is extremely similar to that of modern Oscillatoriaceae. Trichome fragmentation was also observed in a series of specimens, which closely resembles that of cyanobacteria, particularly the Oscillatoriaceae. The termini of Megathrix longus suggest that it was planktonic. In general, Megathrix longus bears a close affinity to the extant Oscillatoriaceae (Cyanophyta).     

贵州织金寒武系戈仲伍组钉形类骨片化石研究*

钉形类(nail-shaped)骨片化石是华南寒武纪早期地层中常见的一类具矿化骨骼的小壳化石。本文系统研究了贵州织金戈仲伍剖面寒武系戈仲伍组的钉形类骨片化石, 发现织金地区钉形类化石主要以织金钉类和疑似的寒武钉类为主, 共4属9种。相较于前人研究, 本文新发现了织金钉类: Zhijinites tumourifomis、Z. lubricus、Z. clavus, 和疑似的寒武钉类: ?Deiradoclavus sp.、?Cambroclavus sp.。排除埋藏学假象后, 分析了钉形类不同化石属种保存方式及形态结构, 发现Zhijinites的多数种都存在盘体顶面凹陷结构, 并且Zhijinites部分种的锥体内壁有收缩现象或具有十字交错纤维结构, 这些发现能较好地揭示出Zhijinites各形态种间的相似性和差异性。分析钉形化石时空分布特点发现, 目前钉形类骨片化石最早在织金地区有化石记录, 之后在扬子地台内部辐射扩散, 并几乎同时辐射扩散至邻近陆块。此外, 本文讨论的疑似寒武钉类化石时代可下延至寒武纪幸运期, 可能是目前该类化石在全球的最早记录, 为探索寒武钉类化石的系统演化和地层学意义提供了新信息。     

The morphology and affinity of anabaritids revealed by their internal molds from the Cambrian Fortunian,southern Shaanxi,China

Anabaritids are a group of tubular skeletal fossils known from the lower Cambrian worldwide. Many previously collected specimens of anabaritids lack both the apical and apertural regions, and thus their complete morphology and growth patterns are not fully understood. Here we describe a number of internal molds of Anabarites from the lower Cambrian (Fortunian Stage) Zhangjiagou Lagerstätte, southern Shaanxi Province. Our new specimens preserved the apex, the aperture with three lobes, and three longitudinal grooves initiating apically rather than adapically as previously reported. This work enriches the morphology of anabaritids, casting doubt on the previous proposition that anabaritids originated from a cylindrical tubular ancestor. It is proposed here that a triradial symmetry may have been established in very early ontogenetic stages of some anabaritids; furthermore, the aperture with three lobes resembles those of olivooids and thus supports a possible close relationship between anabaritids and olivooid scyphozoans.     

A new scyphozoan from the Cambrian Fortunian Stage of South China

Animals with radial symmetry are abundant in the Cambrian Fortunian Stage of South China, but with relatively low diversity: representatives include Olivooides, Quadrapyrgites, carinachitiids, hexangulaconulariids and Pseudooides. Here, we report a new radial animal, Qinscyphus necopinus gen. et sp. nov., from the Fortunian small shelly fauna of southern Shaanxi Province, South China. Qinscyphus necopinus has a cup‐shaped profile, with slightly raised annuli and five groups of triangular thickenings in pentaradial symmetry. This organism has a comparable morphology to, and thus a close affinity with, Olivooides and Quadrapyrgites, and is interpreted as a coronate scyphozoan. This discovery adds a new crown‐group cnidarian to the Cambrian Explosion.     

Ecological innovations in the Cambrian and the origins of the crown group phyla

Simulation studies of the early origins of the modern phyla in the fossil record, and the rapid diversification that led to them, show that these are inevitable outcomes of rapid and long-lasting radiations. Recent advances in Cambrian stratigraphy have revealed a more precise picture of the early bilaterian radiation taking place during the earliest Terreneuvian Series, although several ambiguities remain. The early period is dominated by various tubes and a moderately diverse trace fossil record, with the classical ‘Tommotian’ small shelly biota beginning to appear some millions of years after the base of the Cambrian at ca 541 Ma. The body fossil record of the earliest period contains a few representatives of known groups, but most of the record is of uncertain affinity. Early trace fossils can be assigned to ecdysozoans, but deuterostome and even spiralian trace and body fossils are less clearly represented. One way of explaining the relative lack of clear spiralian fossils until about 536 Ma is to assign the various lowest Cambrian tubes to various stem-group lophotrochozoans, with the implication that the groundplan of the lophotrochozoans included a U-shaped gut and a sessile habit. The implication of this view would be that the vagrant lifestyle of annelids, nemerteans and molluscs would be independently derived from such a sessile ancestor, with potentially important implications for the homology of their sensory and nervous systems.