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.     

RHYNCHONELLIFORMEAN BRACHIOPODS WITH SOFT-TISSUE PRESERVATION FROM THE EARLY CAMBRIAN CHENGJIANG LAGERSTÄTTE OF SOUTH CHINA

Abstract:  Cosmopolitan kutorginates, the most abundant Early Cambrian rhynchonelliformean brachiopods, became extinct at the end of the Middle Cambrian. Consequently, any information concerning the anatomy of this peculiar lineage of brachiopods has great phylogenetic significance with regard to their extant relatives for analogies with the stem-group clade. Such data have been supplied from fossils of which the soft parts have been preserved in exceptional detail. A new brachiopod, Kutorgina chengjiangensis sp. nov., from the Early Cambrian Chengjiang Lagerstätte of southern China, is described here. It is the first articulated brachiopod species collected from this deposit. The specimens preserve a set of soft-body parts, i.e. lophophore, digestive tract and pedicle, all previously poorly known in almost all Palaeozoic calcareous brachiopod taxa. The lophophore attains an early spirolophe stage, clearly homologous to that in the coeval lingulids. The digestive tract consists of a mouth, oesophagus, swollen stomach, intestine and a terminal anus. The pedicle protruding between the valves is stout and elongate, with annulated lamellae on the surface, and contains a putative cœlomic cavity. K. chengjiangensis displays the characteristics of the stem group of calcareous brachiopods, and illustrates anatomical similarities between Cambrian phosphatic- and calcareous-shelled brachiopods, thus corroborating that the Brachiopoda are a monophyletic group.     

Evidence of lophophore diversity in early Cambrian brachiopoda

The lophophore, an essential organ of the Brachiopoda, has been used widely in evolutionary and advanced phylogenetic studies, but is hitherto unknown in the fossil record. Here, the extraordinarily well-preserved lophophores of two inarticulated brachiopods Lingulella chengjiangensis and Heliomedusa orienta, from the Lower Cambrian Chengjiang fauna (Yunnan, China) are described. These primitive lophophores, respectively, trocholophous and schizolophous, have some key characters that may be plesiomorphies inherited by their recent descendants. This discovery provides direct evidence regarding the taxonomy, ecosystems and early evolution of inarticulated brachiopods.     

寒武纪早期澄江生物群中受创伤的中间型始莱德利基三叶虫(Eoredlichia intermedia)

三叶虫是寒武纪海洋中的常见动物,其受伤愈合现象的化石记录可以反映生态系统内物种之间竞争关系。中国华南寒武纪早期澄江化石生物群因保存软躯体构造化石而成为研究寒武纪大爆发时期古海洋生态群落的绝佳窗口,然而,澄江生物群中三叶虫受伤愈合现象的化石至今未见报道。中间型始莱德利基虫(Eoredlichia intermedia Lu,1940)是澄江生物群的常见物种,也是我国寒武系第二个三叶虫化石带的标准带化石。在2 000余枚中间型始莱德利基虫标本中,发现两枚成体标本的头甲侧边缘、颊刺和胸甲肋节保存了不同程度的残缺。据此推测,Eoredlichia intermedia在生命活动过程中曾受到碎壳型捕食动物的攻击或其他伤害而形成伤口。伤口部位边缘加厚且光滑,显示了明显的愈合痕迹。这些特征表明始莱德利基虫在受到非致死的伤害后,具备自身修复损伤的能力。这是迄今所知最为古老的,也是在澄江生物群中首次发现的三叶虫受伤并且愈合的标本,反映了寒武纪第二世第三期底栖生态系统中碎壳型捕食者和三叶虫猎物之间的军备竞赛生态关系。     

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.     

贵州麻江地区寒武纪杷榔动物群的群落组成和生态空间分析*

寒武纪特异埋藏化石库保存了海洋生物群落最原始的生态结构, 为群落生态研究提供了宝贵材料。贵州寒武纪斜坡相区的杷榔动物群(第4期)具有较高的生物多样性, 但定量生态学研究相对薄弱。本研究对贵州麻江地区杷榔动物群的新产地平定剖面进行化石采集和分类统计, 并运用多元数理统计方法开展定量化分析。鉴定出腕足动物、海绵动物、刺胞动物、节肢动物、软舌螺动物、古虫动物、开腔骨动物和曳鳃动物8个化石门类, 计31属33种, 划分出12种生态类型。其中, 腕足动物具有最高的个体丰度(占比61.5%), 节肢动物物种分异度最高(占比60%), 群落以底栖固着生态类型为主, 游泳捕食为辅。对比分析结果显示, 寒武纪海洋群落具有空间异质性, 且节肢动物在影响群落组成和生态空间利用上起决定性作用。随水体深度增加, 海洋生态群落中的表栖类群出现明显扩张, 也体现了环境对群落面貌的塑造作用。     

Origin and dispersal of the earliest brachiopods

Brachiopods first appeared at the very beginning of the Phanerozoic together with the first skeletal organisms. Most brachiopod taxa that arose in the first half of the Cambrian had a short temporal range and became completely extinct by the middle of the Middle Cambrian. Rigid articulation of the valves of brachiopods was provided by various structures, which also appeared in the Early Cambrian. This fact points to the importance of this feature for the formation of the whole group and at the same time testifies to the high variability of rigid articulation at the early stages of brachiopod evolution. This is a typical manifestation of archaic diversity in this animal phylum, which appeared very early in the Phanerozoic. Another important property of the archaic diversity of the early brachiopods was the large number of centers of diversification. As for the majority of groups, climatic zonality was the main factor determining the distribution of brachiopods at the beginning of the Phanerozoic. The main ecological types of brachiopods also appeared in the Early Cambrian.     

Shell structure of <Emphasis Type="Italic">Kutorgina</Emphasis> Billings (Brachiopoda,Kutorginida)

Shell structure in members of the genus Kutorgina, Lower Cambrian brachiopods with calcareous shells and primitive articulation, is described for the first time. This type of shell structure resembles that of the Lower Cambrian mollusks and hyoliths rather than of brachiopods.     

Drill holes in Australian Cainozoic brachiopods

The fossil record of drill holes in marine invertebrates has received a considerable amount of interest from paleontologists, primarily due to its importance for reconstructing the history of interactions between drilling predators and their prey. Such drill holes have been described in numerous studies of Paleozoic brachiopods but rarely in those focusing on brachiopods of the post-Paleozoic, a striking pattern given that in the late Mesozoic and Cainozoic drilling gastropods diversified and frequencies of drilled molluscs increased dramatically. During the past several years, however, drilled brachiopods were reported in several studies of the Mesozoic and Cainozoic, suggesting that this phenomenon may be more common than has been previously assumed. Here we report on 10 genera of brachiopods from four Cainozoic basins in Australia of which 7 shows evidence of having been drilled by predators. Of 298 specimens examined, 38 contain a single complete hole. Drilled specimens were identified in all 4 basins and in all stratigraphic units. When considered in the context of recent reports of drilled Cainozoic brachiopods, these Australian brachiopods further imply that drilling predation on these invertebrates was geographically, taxonomically and temporally widespread.     

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.     

A new family of Cambrian rhynchonelliformean brachiopods (Order Naukatida) with an aberrant coral‐like morphology

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian (Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian (Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral‐like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube‐like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian (Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef‐building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well‐developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle.     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

First record of a bivalved larval shell in Early Cambrian tommotiids and its phylogenetic significance

Abstract: Brachiopods are marine Lophotrochozoa whose soft parts are enclosed in a bivalved shell. Although brachiopods are represented by a rich record from the Early Cambrian to the present, the origin of their bivalved body plan remains controversial. The Early Cambrian organophosphatic tommotiids Micrina and Paterimitra from Australia have been proposed as stem brachiopods. Here, we describe their earliest ontogeny, indicating that tommotiids possessed bivalved planktotrophic larvae. The curious combinations of characters in Micrina and Paterimitra indicate that they may belong to the stems of the Linguliformea and Rhynchonelliformea, respectively. The bivalved shell of adult living brachiopods may represent a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan of the Brachiopoda may have evolved through the paedomorphic retention of a bivalved larval state.     

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.     

Fulu biota,a new exceptionally-preserved Cambrian fossil assemblage from the Longha Formation in southeastern Yunnan

A new exceptionally-preserved Cambrian biota, with fossils belonging to multiple phyla, has recently been found from the middle part of the Longha Formation, near Fulu Village, southeastern Yunnan, South China, and is named the Fulu biota. Groups recovered so far include polymerid trilobites, agnostoids, large bivalved arthropods, bradoriid arthropods, echinoderms, brachiopods, priapulids (worms), hyoliths, macroscopic algae, and trace fossils. The arthropods dominate the biota and are highly diverse with echinoderms second in diversity. The associated agnostoids provide a precise late Guzhangian Age (late Miaolingian Epoch) for the biota, indicating that the biota occurs stratigraphically about halfway between the occurrences of the Kaili and the Guole biotas of South China, from both of which the new Fulu biota differs significantly in faunal components. Guzhangian biotas preserving poorly mineralized tissues and complete echinoderms have been previously unknown in Gondwana until now; thus, the discovery of Fulu biota provides a unique window into life at this critical time interval.     

BILLENGSELLIDE AND ORTHIDE BRACHIOPODS: NEW INSIGHTS INTO EARLIEST ORDOVICIAN EVOLUTION AND BIOGEOGRAPHY FROM NORTHERN IRAN

Abstract:  The eastern Alborz Mountains of Iran comprise a significant peri-Gondwanan terrane relevant to the early evolution of late Cambrian – early Ordovician brachiopods incorporated into the emerging benthic biota of the Paleozoic Evolutionary Fauna. A low diversity brachiopod assemblage from the late Tremadocian unit of the Lashkarak Formation contains six new species including the polytoechioideans Polytoechia and Protambonites and the orthoideans Paralenorthis, Ranorthis, Tarfaya and Xianorthis . The fauna preserves the earliest records of Polytoechia , unknown previously outside Laurentia and the Uralian margin of Baltica, and of Paralenorthis and Ranorthis , which were widespread along Gondwanan margins and in Baltica from the Floian (Arenig), plus Xianorthis , known hitherto only from the Floian of South China. The enigmatic Tarfaya has an impunctate shell fabric and setigerous perforations along the posterior margin, indicating placement within the Orthoidea in a new Family Tarfayidae. New species of Polytoechia , Protambonites , Paralenorthis, Ranorthis , Tarfaya , Xianorthis are described.     

Evidence of predation damage in Pliocene Apletosia maxima (Brachiopoda)

Abstract:  Little is known about predation of Mesozoic and Cenozoic articulated brachiopods, but it is far from clear whether this is because they suffered very little predation pressure or because there have been few attempts to search for evidence of it. A study of 248 museum specimens of the large Pliocene terebratulid Apletosia maxima from the Coralline Crag (UK) has revealed that more than 16 per cent of them show evidence of having been attacked by predators. The styles of damage can be attributed to drilling muricid gastropods (most of which were successful) and failed crushing attacks probably by decapods. Brachiopods are usually thought to offer a poor tissue yield to potential predators, but in this instance it appears that A. maxima was attractive to predators even though they were living with a rich molluscan fauna. It is suggested that the mass of adductor and diductor muscles (likely to be spicule-free) of these particularly large brachiopods may have made them profitable. Further studies of post-Palaeozoic brachiopod faunas are required, particularly those from mixed shallow-water communities, before it can be established whether articulated brachiopods have or have not been driven into refugia by increasing predation pressure.     

The origin and geometry of radial ribbing patterns in articulate brachiopods

Ackerly, S. C. 1992 07 15: The origin and geometry of radial ribbing patterns in articulate brachiopods.
Geometric models for simple. radial ribbing in articulate brachiopods include (1) ribs radiating isometrically from the shell umbo. (2) divergence of thc ribs from some 'point' within the shell, and (3) reorientation of the ribs at right angles to the shell margin. Analyses of the Orthida, the ancestral taxon of articulate brachiopods, indicate that rib geometries are isometric in Early Cambrian taxa (model 1). but that by the Early Ordovician rib orientations are generally perpendicular to the shell margin (model 3). A combination of functional and morphogenetic Factors explains the ribbing geometries observed in orthide brachiopods.     

The monophyletic origin of the Brachiopoda

Although it is commonly acepted that the brachiopods descended from phoronid-like ancestors there is dispute over their origin. Traditionally they have been regarded as a monophyletic group, a clade. More recently it has been claimed that brachiopods are polyphyletic and that several of the orders arose independently from separate phoronid-like stocks. The latter point of view implies that brachiopods are not a taxon but merely a grade of organization. Traditional stratophenetic approaches do little to resolve the problem, which may be outside their domain. It is possible, even probable, that the initial radiation involved organisms that lacked mineralized shells. Cladistic analysis of both living forms and Lower Paleozoic taxa strongly supports the contention that brachiopods are monophytetic and closely related to the phoronids. It suggests, however, that the 'inarticulate' Paterinida and Kutorginida are genealogically more closely related to the Articulata than they are to the remaining Inarticulata. □ Brachiopoda, Lophophorata, cladistics, Cambrian.