Trace fossils in the Ediacaran-Cambrian transition: Behavioral diversification, ecological turnover and environmental shift

After taxonomic revision, trace fossils show a similarly explosive diversification in the Ediacaran-Cambrian transition as metazoan body fossils. In shallow-marine deposits of Ediacaran age, trace fossils are horizontal, simple and rare, and display feeding strategies related to exploitation of microbial matgrounds. Equally notable is the absence of arthropod tracks and sinusoidal nematode trails. This situation changed in the Early Cambrian, when a dramatic increase in the diversity of distinct ichnotaxa is associated was followed by the onset of vertical bioturbation and the disappearance of a matground-based ecology (‘‘agronomic revolution’’). On deep sea bottoms, animals have been present already in the Ediacaran, but ichnofaunas were poorly diverse and dominated by the horizontal burrows of undermat miners. As shown by the ichnogenus Oldhamia, this life style continued to be predominant into the Early, and to a lesser extent, Middle Cambrian. Nevertheless, there was an explosive radiation of behavioral programs during the Early Cambrian. When exactly the bioturbational revolution arrived in the deep sea is uncertain. In any case, the Nereites ichnofacies was firmly established in the Early Ordovician. The rich ichnofauna in the Early Cambrian Guachos Formation of northwest Argentina probably marks a first step in this ecological onshore-offshore shift.     

Occurrence of the Lower Cambrian anemone-style trace fossils in the Zabuk Formation (Mardin–Derik,SE Turkey)

The Zabuk Formation of the Derik Group exposed over much of south-eastern Turkey presents a succession composed of shallow marine and fluvial siliciclastic sedimentary rocks. The Lower Cambrian assemblages containing abundant anemone-style trace fossils are known from most major palaeocontinents such as Laurentia, Baltica, and Gondwana. These have possible affinities with semi-infaunal dwelling anemones on siliciclastic mid-latitude shelves of West Gondwana in early Fortunian deposits. Among them is Bergaueria, characterized by plug-shaped burrows as exemplified by Bergaueria perata, that is, a characteristic trace fossil of the Cambrian globally.     

Complex behavioural patterns and ethological analysis of the trace fossil Zoophycos: evidence from the Lower Devonian of South China

The trace fossil Zoophycos is abundant in the shallow‐marine deposits (tempestites) of the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by deposit‐feeding behaviour, where the animal constructed the trace upwards without leaving faeces in the spreiten. The width of the marginal tube in different whorls is almost constant. The scratches are observed on the wall of the marginal tubes. The Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos‐maker might have been a vermiform polychaete instead of a predator such as a decapod crustacean (Spongeliomorpha producer). Based on stratigraphical and ichnological features, the complex trace fossils resulted from the complex activity of different opportunistic organisms (r‐strategist) that quickly occupied and thrived within the quiet, nutrient‐rich environment after storm events.     

A new rhynchonelliform brachiopod Longtancunella with soft-part preservation from the Hongjingshao Formation (Cambrian Stage 3) in Yunnan,South China

Rhynchonelliform brachiopods made their first appearance in early Cambrian, and became a major group within the palaeozoic evolutionary fauna since late Cambrian. Exceptionally preserved fossils from the early Cambrian Lagerstätten provide valuable chances to investigate their phylogeny and ecology. Longtancunella is one of the most interesting early rhynchonelliforms, and has been mainly recovered from the Chengjiang Lagerstätte (Series 2, Stage 3). Here, we report a new rhynchonelliform Longtancunella xiazhuangensis n. sp. from the lower Hongjingshao Formation (upper Stage 3) in Yunnan Province, China. These specimens were well preserved with soft parts, including pinnate mantle canal system and a pedicle. It is identified as a new species based mainly on its difference in shell ornamentations, pinnate mantle canals and pedicle morphology from the type species. Its pedicle looks unusually stout with distinct annulated lamellae on the surface, and reveals crucial evidence in illustrating its ecology and settling strategy as an early marine epifauna. The ecological interaction between L. xiazhuangensis and other marine animals also provides insights into the food web structure in the early Cambrian.     

The changing face of the deep: Colonization of the Early Ordovician deep-sea floor, Puna, northwest Argentina

An Upper Tremadocian deep-sea ichnofauna from the Chiquero Formation of Puna, northwest Argentina, represents a link between Ediacaran and Cambrian microbial-mat dominated ecosystems and younger Ordovician deep-marine trace-fossil assemblages. This ichnofauna is preserved at the base of thin-bedded turbidites formed in the lobe fringe of a back-arc deep-sea fan. While Ediacaran–Cambrian deep-marine trace fossils are typically linked to matground grazing and feeding, microbial textures in the Chiquero Formation are rare and not associated with trace fossils. Morphologic patterns (e.g. radial trace fossils and networks) of the Chiquero ichnofauna indicate the onset of novel trophic types, recording trapping of microorganisms and bacterial farming. However, in comparison with younger Ordovician deep-sea ichnofaunas, graphoglyptids are relatively rare, poorly diverse, and geometrically simpler. This study indicates that the Early Ordovician was a pivotal point in the ecology of deep-sea infaunal communities. This Upper Tremadocian ichnofauna records the arrival of the Agronomic Revolution to the deep sea. Comparisons with slightly older and younger deep-sea ichnofaunas demonstrate that the colonization of the deep sea was a protracted process spanning the Early Paleozoic, lagging behind colonization of nearshore and offshore substrates.     

湖北三峡地区埃迪卡拉系灯影组“蝌蚪状”遗迹化石*

遗迹化石是埃迪卡拉纪存在两侧对称动物最有力的证据。但多数埃迪卡拉纪遗迹化石为简单、水平的表面爬迹或潜穴。在湖北三峡地区灯影组石板滩段含典型埃迪卡拉软躯体化石的地层中新发现一类形态特别的化石,呈蝌蚪状,一端膨大,一端细管状。通过对化石形态、同位素分析以及沉积学特征的研究,说明该化石为遗迹化石,而不是实体化石。蝌蚪状化石为一种复合迹,垂向活动形成球状的膨大端,平行藻席层活动形成近于水平的潜穴,反映了造迹生物垂向切穿藻席层并沿藻席层进行觅食的行为。该发现说明了在埃迪卡拉纪晚期已有两侧对称动物开始形成较为复杂的潜穴。     

扬子区晚震旦世动物化石新材料

扬子区晚震旦世地层中已有多种类型的动物及遗迹化石的报道。文中补充了不同地区和层位的骨骼及遗迹化石新材料,包括：江西上饶王家院陡山沱组顶部磷块岩中的磷酸盐化“十”字形骨针和云南晋宁王家湾灯影组上段隧石条带中的单轴双射硅质骨针,它们可能分属于钙质海绵和普通海绵类;贵州瓮安陡山沱组上段磷质白云岩中的可疑壳状化石;宜昌三斗坪金瓜墩陡山沱组下部（第二段）遗迹化石;瓮安陡山沱组上段及湖北南漳灯影组上段底部磷质硬底中的生物扰动构造。另外还报道了贵州清镇桃子冲剖面桃子冲组最底部的瓶状动物（Pro-tolagena limbata）化石,并认定桃子冲组与灯影组之间为假整合接触,其底界并无跨入震旦系的证据。     

Oldhamia radiata: the trace of an undermat miner from the Cambrian Series 2 Xinji Formation,western Henan,North China

Oldhamia trace fossils represent a complex horizontal burrow system usually associated with microbial mats, occurring in the upper part of the Cambrian Stage 2 of the Terreneuvian Series, with a peak in abundance during Stage 3 and 4 of Series 2, and then rapidly disappearing in the Wuliuan Stage of the Miaolingian Series. We here report Oldhamia radiata from the Cambrian Series 2 Xinji Formation in western Henan of North China, associated with microbial mats, as evidenced by the presence of wrinkle structures. The distinctive patterns indicate the exploitation of microbial mats by the tracemakers of O. radiata, featuring a specific feeding strategy under the mats where possibly anoxic and sulfidic conditions prevailed during the early Cambrian.     

First occurrence of the Ichnogenus Selenichnites from the Middle Jurassic Strata of the Skoura Syncline (Middle Atlas,Morocco); Palaeoecological and palaeoenvironmental context

Mesozoic strata of North Africa yield the first occurrence of the ichnogenus Selenichnites. The trace fossils occur on the top surface of a sandy carbonate deposit in the axis of a Middle Atlas syncline (Skoura Syncline, NE Morocco). The ichnofossil-bearing horizon belongs to the Late Bajocian–Early Bathonian Ich Timellaline/Bou Akrabene Formation. The trace fossils are crescent-shaped and the best preserved exhibits a posterior central axial impression (possible telson tail impression). They are interpreted as feeding burrows (fodinichnia) or hiding depressions of Xiphosurids or Limulids (horseshoe crabs) on a sandy carbonate substrate beneath a veneer of muddy deposits. The sedimentological character suggests a relatively protected shallow water subtidal palaeoenvironment preceding the Bathonian regression of the Atlas domain. This discovery provides the first evidence of xiphosurans or xiphosuran-like organisms inhabiting the southern shores of the Tethys in the Middle Jurassic.     

北京昌平青白口系痕迹化石

北京昌平青白口系长龙山组和景儿峪组发育有丰富的痕迹化石,其中长龙山组产有Ｈｅｌｍｉｎｔｈｏｐｓｉｓ ｓｐ．,Ｃｏｃｈｌｉｃｈｎｕｓ ｓｐ．,代表潮下低能环境;景儿峪组产有Ｓｋｏｌｉｔｈｏｓ ｓｐ．和Ｍｏｎｏｃｒａｔｅｒｉｏｎ ｓｐ．,代表高能潮间环境。     

Silurian trace fossils in carbonate turbidites from the Alexander arc of southeastern Alaska

Early to Late Silurian (Llandovery‐Ludlow) body and trace fossils from the Heceta Formation of southeastern Alaska are preserved in the oldest widespread carbonates in the Alexander terrane. These fossils represent the earliest benthos to inhabit diverse shallow and deep subtidal environments in the region and are important indicators of early stages in benthic community development within the evolving Alexander arc. The ichnofossils are particularly significant because they add to a small but growing body of knowledge about trace fossils in deep‐water carbonates of Paleozoic age.

Carbonate turbidites that originated along a deep marine slope within the arc yield a low‐diversity suite of trace fossils consisting of five distinct biogenic forms. Simple burrows (Planolites, two forms), ramifying tunnels (Chondrites), and tiny cylindrical burrows (?Chondrites) represent the feeding activities (fodinichnia) of pre‐turbidite animals that burrowed in the lime mud before the influx of coarser sediment deposited by turbidity currents. These trace fossils are associated locally with cross‐cutting burrows created as domichnia (Palaeophycus). Rarer hypichnial burrows and endichnial traces were created by post‐turbidite animals that fed soon after the deposition of coarse detritus from turbidity flows.

Trace fossils in these deposits reflect much lower diversity levels than in Paleozoic siliciclastic turbidites. This difference may represent unfavorable environmental conditions for infaunas, differential preservation, or significant paleogeographic isolation of the Alexander terrane during the Silurian. Greater utilization of trace fossils in terrane analysis may help to resolve this issue and provide new data for reconstructing the paleogeography of circum‐Pacific terranes.     

Trace fossils from the Lower Triassic of North China — a potential signature of the gradual recovery of a terrestrial ecosystem

The Permian–Triassic Mass Extinction (PTME) resulted in the most severe loss of life in the history of the earth, both marine and terrestrial ecosystems suffered heavy losses simultaneously, and the biotic recovery process was difficult for both. There are many studies on the timing and pattern of this mass extinction, however, the consistency of the recovery between the marine and terrestrial facies remains controversial. In particular, few metazoans survived and the deposits were dominated by the microbialites during the Early Triassic. The reappearance of the metazoans signified the beginning of the biotic recovery. In this paper, we measured the trace fossils of the Heshanggou Formation in western Henan, North China, which plays an important role in the transformation from the flourishing of the microorganisms to the recovery of metazoans. We quantitatively described the characteristics and measured the diameter, depth, abundance and diversity of the trace fossils. Based on the acquired data, the lower part of the Heshanggou Formation is confined to simple, small, vertical burrows, suggesting that the ecological environment was suitable for the survival of special species. In the middle part, the vertical burrows become deeper, which may be due to the adaptation of organisms to the harsh environment. Disturbance is more intense and the diversity clearly increases in the upper part. Through the increasing bioturbation and diversity of the trace fossils, we found a gradual biotic recovery pattern in the Early Triassic, and the recovery mode is similar to marine ichnofossils in South China.     

Infaunal augurs of the Cambrian explosion: An Ediacaran trace fossil assemblage from Nevada,USA

Morphologically complex trace fossils, recording the infaunal activities of bilaterian animals, are common in Phanerozoic successions but rare in the Ediacaran fossil record. Here, we describe a trace fossil assemblage from the lower Dunfee Member of the Deep Spring Formation at Mount Dunfee (Nevada, USA), over 500 m below the Ediacaran–Cambrian boundary. Although millimetric in scale and largely not fabric‐disruptive, the Dunfee assemblage includes complex and sediment‐penetrative trace fossil morphologies that are characteristic of Cambrian deposits. The Dunfee assemblage records one of the oldest documented instances of sediment‐penetrative infaunalization, corroborating previous molecular, ichnologic, and paleoecological data suggesting that crown‐group bilaterians and bilaterian‐style ecologies were present in late Ediacaran shallow marine ecosystems. Moreover, Dunfee trace fossils co‐occur with classic upper Ediacaran tubular body fossils in multiple horizons, indicating that Ediacaran infauna and epifauna coexisted and likely formed stable ecosystems.     

Lithostratigraphy of the Lower Cambrian metaclastics and their age based on trace fossils in the Sand?kl? region, southwestern Turkey

In the Sand?kl? region of the Taurus Range of Turkey, greater than 3000 m in thickness metamorphosed siliciclastics and volcanics (Kocayayla Group) underlies the trilobite-and conodont-bearing Middle-Late Cambrian Hudai Quartzite and Çaltepe Formation.The Kocayayla Group, previously regarded as Infracambrian or Precambrian, is dated for the first time as Early Cambrian on the basis of trace fossils. Cruziana ?fasciculata, C. ?salomonis, ?Cruziana isp., ?Diplichnites isp., Monomorphichnus isp., Petalichnus isp., Rusophycus ?avalonensis, R. ?latus, Arenicolites isp., cf. Altichnus foeyni, Planolites isp., Skolithos isp., and ?Treptichnus isp. have been recognised. These trace fossils are considered Tommotian or younger in age but older than the overlying, trilobite and conodont bearing Middle Cambrian limestones of the Çaltepe Formation. The trace fossils were likely produced by trilobites, suspension feeding annelids and deposit feeding “worms”, probably polychaetes. Sections bearing abundant Skolithos represent the Skolithos ichnofacies, which is typical of high energy environments with loose sandy, well sorted to slightly muddy substrates in intertidal to shallow subtidal zones. The other trace fossils represent the Cruziana ichnofacies, which is typical of subtidal, poorly sorted and soft substrates, from moderate energy to low energy environments between the fairweather and storm wave base.The Kocayayla Group was deposited at an early stage in a shallow marine stable shelf condition. The shelf subsided in a later stage and was affected by normal faults along which mafic and felsic volcanic rocks erupted. The volcanic activity had ceased and a shallow marine clastic sedimentation took place in the final stage of the shelf development. The Kocayayla Group was deformed and metamorphosed before the deposition of the trilobite-bearing Middle-Upper Cambrian succession.     

陕西镇巴下寒武统水井沱组小壳化石新属种

本文描述了陕西镇巴下寒武统水井沱组中的小壳化石３新属５新种。样品采自镇巴小洋坝曾家坡,共生化石有三叶虫、古介形虫、海绵骨针和大量软舌螺等。化石组合面貌表明其地质时代属筇竹寺期。另外,在峡东水井沱组中发现了微网虫（Ｍｉｃｒｏｄｉｃｔｙｏｎｓｐ．）化石骨片（图版Ｉ,图１８,１９）,可与云南筇竹寺组及陕南水井沱组对比,说明Ｍｉｃｒｏｄｉｃｔｙｏｎ广泛分布于扬子地台区。     

A comparison of crayfish burrow morphologies: Triassic and Holocene fossil,paleo‐ and neo‐ichnological evidence,and the identification of their burrowing signatures

The architectural and surficial morphologies of crayfish burrows from the Upper Triassic Chinle Formation and Holocene sediments were compared in order to determine: 1) if Triassic burrows could truly be attributed to crayfish activity; 2) how comparable the burrowing mechanisms are; and 3) whether or not a common set of burrowing signatures could be identified for both ancient and modern freshwater crayfish. Materials used in this study include burrows from the members of the Upper Triassic Chinle Formation, casts of modern burrows constructed by Procambarus clarkii Hobbs and Procambarus acutus acutus (Girard) in the laboratory, and casts of naturally constructed modern burrows of Cambarus diogenes di‐ogenes (Girard).

Triassic and Holocene crayfish burrow morphologies exhibit simple to complex architectures, varying degrees of branching, chamber, and chimney development. They also exhibit relatively textured surficial morphologies (bioglyphs) such as scrape and scratch marks, mud‐ and lag‐liners, knobby and hummocky surfaces, pleopod striae, and body impressions. Holocene crayfish construct distinctive burrows due to their conservative limb arrangement, functional morphology, and behavior with respect to environmental stimuli. Similarities between Holocene and Triassic crayfish burrows suggest that extant and Triassic crayfish employed identical burrowing mechanisms. Features of the surficial and architectural morphologies impart a distinctive signature to burrows of both ancient and modern freshwater burrowing crayfish.

Burrowing signatures of crayfish can be used to identify new and previously misinterpreted continental trace fossils. These are useful in studies of the paleohydrogeology, paleoclimatology and paleoecol‐ogy of burrow‐bearing deposits.     

华中西南区一条国际前寒武系与寒武系界线层型补充剖面

1992年,全球前寒武系与寒武系界线层型剖面和层型点确定在加拿大纽芬兰东南幸运角剖面,并以遗迹化石带Trichophycus pedium的底作为前寒武系与寒武系的分界点。但是,幸运角剖面主要是以硅质碎屑岩相为主,难以与含有丰富小壳化石和具有可对比的稳定同位素资料的碳酸盐相界线剖面进行对比。为此,提出我国云南会泽大海附近的一条以碳酸盐相为主的剖面作为全球前寒武系与寒武系界线层型剖面的补充剖面。理由是（1）在大海剖面上,震旦系灯影组白岩哨段、待补段和下寒武统牛家菁组的中谊村段,大海段之间为整合接触;（2）早寒武世早期地层单元含有丰富的小壳化石并且可划分出4个小壳化石组合带;（3）利用碳同位素资料可以将大海剖面与许多其它剖面相当地层对比;（4）著名的梅树村剖面离大海剖面不远,它们均含有丰富的小壳化石和遗迹化石,两者易于对比。文章进一步认为寒武系的下界放在第I小壳化石组合带（Anabarites trisulcatus-Protohertzina anabarica）的底,相当于大海剖面11层的底。     

云南澄江早寒武世地层并论中国的Parabadiella,摩洛哥的Abadiella及澳大利亚的Abadiella huoi

根据近来的发现,区域地层的阐明及生物在寒武系界线上下的变化,澄江早寒武世地层划分作了修订。早寒武世岩石地层名称的更换,年代地层及生物地层单位的分带这里作了简述.目前的岩石地层单位从上到上是渔户村组（梅树阶）,黑林铺组（筇竹寺阶）,红井哨组、乌龙箐组（沧浪铺阶）及山邑村组（龙王庙阶）。此层序可分为2个亚带。所有的组合带在中国西南地台区都有存在,但两个亚带在澄江是区域性的。云南东部震旦系与寒武系的界线暂时置于渔户村且小歪头山段的底界,位于小壳化石最早出现的下部。中国的Parabadiella,摩洛哥的Abadiella及澳大利亚的Abadiella huoi这里重新作了研究。Parabadiella及Abadiella是两个不同的属。模式标本的图片在这里再次刊出。澳大利亚的Abadiella huoi一种应是Wutingaspis的一个新种（Wutingaspis jelli Chang sp.nov)。Opik(1975）的Dolerolenus(?)sp.nov.一种应归属Parabadiella huoi一种。Parabadiella是中国也是澳大利亚寒武系最古老的三叶虫。澳大利亚的Wutingaspis jelli的层位与摩洛哥Abadiella的层位相当,可与中国的Wutingaspis-Eoredichia带的上部对比。     

Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca,Spain

Gibert, J.M. de, Mas, G. & Ekdale, A.A. 2012: Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca, Spain. Lethaia, Vol. 45, pp. 574–585. Unusual helical trace fossils occur in Tortonian shelfal calcarenites in the island of Mallorca. Their morphology may be either simple (ichnogenus Gyrolithes) or, more commonly, consist of two concentric helical burrows (ichnogenus Lapispira). They bear a very characteristic pelleted lining and are associated and probably connected to Thalassinoides and Ophiomorpha burrow systems very abundant in the same unit. These features allow the interpretation that the tracemaker was a thalassinidean shrimp. The complex and compound nature of these trace fossils is comparable to that seen in other modern and fossil crustacean burrow systems, and it reflects the behavioural plasticity of the architects. □Trace fossils, ichnology, ichnofabrics, crustacean burrows, Miocene, shallow marine, Lapispira.