Skimming the surface with Burgess Shale arthropod locomotion

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates.     

FOSSIL DIAGENESIS IN THE BURGESS SHALE

Abstract:  Current models for the exceptional preservation of Burgess Shale fossils have focused on either the HF-extractable carbonaceous compressions or the mineral films identified by elemental mapping. BSEM, EDX and microprobe analysis of two-dimensionally preserved Marpolia , Wiwaxia and Burgessia identifies the presence of both carbonaceous and aluminosilicate films for most features, irrespective of original lability. In the light of the deep burial and greenschist facies metamorphism documented for the Burgess Shale, the aluminosilicate films are identified as products of late-stage volatilization and coincident mineralization of pre-existing compression fossils, whereas the three-dimensionally preserved gut-caecal system of Burgessia is interpreted as an aluminosilicate replacement of a pre-existing carbonate phase. The case for late diagenetic emplacement of aluminosilicate minerals is supported by the extensive aluminosilicification of trilobite shell and (originally) calcareous veinlets in the Burgess Shale, as well as documentation of other secondarily aluminosilicified compression fossils. By distinguishing late diagenetic alteration from the early diagenetic processes responsible for exceptional preservation, it is possible to reconcile the range of preservational modes currently expressed in the Burgess Shale.     

From weird wonders to stem lineages: the second reclassification of the Burgess Shale fauna

The Burgess Shale, a set of fossil beds containing the exquisitely preserved remains of marine invertebrate organisms from shortly after the Cambrian explosion, was discovered in 1909, and first brought to widespread popular attention by Stephen Jay Gould in his 1989 bestseller Wonderful life: The Burgess Shale and the nature of history. Gould contrasted the initial interpretation of these fossils, in which they were 'shoehorned' into modern groups, with the first major reexamination begun in the 1960s, when the creatures were perceived as 'weird wonders', possessing unique body plans and unrelated to modern organisms. More recently, a third phase of Burgess Shale studies has arisen, which has not yet been historically examined. This third phase represents a revolutionary new understanding, brought about, I believe, by a change in taxonomic methodology that led to a new perception of the Burgess creatures, and a new way to comprehend their relationships with modern organisms. The adoption of cladistics, and its corollary, the stem group concept, has forged a new understanding of the Burgess Shale ... but has it also changed the questions we are allowed to ask about evolution?     

Dynamic palaeoredox and exceptional preservation in the Cambrian Spence Shale of Utah

Garson, D.E., Gaines, R.R., Droser, M.L., Liddell, W.D. & Sappenfield, A. 2011: Dynamic palaeoredox and exceptional preservation in the Cambrian Spence Shale of Utah. Lethaia, Vol. 45, pp. 164–177. Burgess Shale‐type faunas provide a unique glimpse into the diversification of metazoan life during the Cambrian. Although anoxia has long been thought to be a pre‐requisite for this particular type of soft‐bodied preservation, the palaeoenvironmental conditions that regulated extraordinary preservation have not been fully constrained. In particular, the necessity of bottom water anoxia, long considered a pre‐requisite, has been the subject of recent debate. In this study, we apply a micro‐stratigraphical, ichnological approach to determine bottom water oxygen conditions under, which Burgess Shale‐type biotas were preserved in the Middle Cambrian Spence Shale of Utah. Mudstones of the Spence Shale are characterized by fine scale (mm‐cm) alternation between laminated and bioturbated intervals, suggesting high‐frequency fluctuations in bottom water oxygenation. Whilst background oxygen levels were not high enough to support continuous infaunal activity, brief intervals of improved bottom water oxygen conditions punctuate the succession. A diverse skeletonized benthic fauna, including various polymerid trilobites, hyolithids, brachiopods and ctenocystoids suggests that complex dysoxic benthic community was established during times when bottom water oxygen conditions were permissive. Burgess Shale‐type preservation within the Spence Shale is largely confined to non‐bioturbated horizons, suggesting that benthic anoxia prevailed in intervals, where these fossils were preserved. However, some soft‐bodied fossils are found within weakly to moderately bioturbated intervals (Ichnofabric Index 2 and 3). This suggests that Burgess Shale‐type preservation is strongly favoured by bottom water anoxia, but may not require it in all cases. □Anoxia, Burgess Shale, Burgess Shale type‐preservation, Langston Formation, Spence Shale Member, Utah.     

Possible ctenophoran affinities of the Precambrian "sea-pen" Rangea

The Namibian Kuibis Quartzite fossils of Rangea are preserved three-dimensionally owing to incomplete collapse of the soft tissues under the load of instantaneously deposited sand. The process of fossilization did not reproduce the original external morphology of the organism but rather the inner surface of collapsed organs, presumably a system of sacs connected by a medial canal. The body of Rangea had tetraradial symmetry, a body plan shared also by the White Sea Russian fossil Bomakellia and possibly some other Precambrian frond-like fossils. They all had a complex internal anatomy, smooth surface of the body, and radial membranes, making their alleged colonial nature unlikely. Despite a different style of preservation, the Middle Cambrian Burgess Shale frond-like Thaumaptilon shows several anatomical similarities to Rangea. The body plan of the Burgess Shale ctenophore Fasciculus, with its numerous, pinnately arranged comb organs, is in many respects transitional between Thaumaptilon and the Early Cambrian ctenophore Maotianoascus from the Chengjiang fauna of South China. It is proposed that the irregularly distributed dark spots on the fusiform units of the petaloid of Thaumaptilon represent a kind of macrocilia and that the units are homologous with the ctenophoran comb organs. These superficial structures were underlain by the complex serial organs, well represented in the fossils of Rangea. The Precambrian "sea-pens" were thus probably sedentary ancestors of the ctenophores.     

贵州台江凯里动物群中的非钙质藻类化石

贵州台江凯里组的非钙质藻类化石在我国中寒武统尚属首次发现。国外主要见于北美。描述的非钙质藻类化石Ｍａｒｐｏｌｉａ ｓｐｉｓｓａ． Ｂｏｓｗｏｒｔｈｉａ ｓｉｍｕｌａｎｓ ,Ａｌｇａ ｇｅｎ． ｅｔ ｓｐ． ｉｎｄｅｔ, Ａ,Ａｌｇａ ｅｔｓｐ． ｉｎｄｅｔ, Ｂ等常见于北美布尔吉斯页岩动物群。当前中武寒武非钙质藻类化石的发现不仅填补了我国中寒武统非钙质藻类的空白,而且对于凯里动物群与布尔吉斯页斯     

The bivalved arthropods Isoxys and Tuzoia with soft‐part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island,Australia)

Abstract: Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft‐bodied features previously unknown from this Burgess Shale‐type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better‐known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.     

Interpreting ‘shelly’ fossils preserved as organic films: the case of hyolithids

Most studies of Burgess Shale‐type preservation have focussed on soft‐bodied organisms, but ‘shelly’ fossils are also preserved as carbonaceous films. These films are usually interpreted as coherent organic layers – often external sheaths or periostracal layers – that were present in the original mineralized elements. The example of hyolithids shows that the organic films of skeletal parts do not represent original ‘layers’, but a composite resulting from the coalescence, into a single carbonaceous film, of all the preservable organic matter present in the skeletal element. The diagenetic processes that led to Burgess Shale‐type preservation, which involve the polymerization of organic matter and the loss of original internal structure and chemical integrity of the original tissues, are entirely compatible with – and could account for – the characteristics observed in the fossil films of hyolithid skeletal elements. These observations have general implications for the interpretation of other organisms preserved as carbonaceous films, such as the diverse and often problematic Cambrian sponges.     

A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale

Wiwaxia corrugata and the indisputable polychaetes of the Middle Cambrian Burgess Shale, particularly Canadia spinosa, have figured prominently in recent hypotheses about the early evolution of polychaete annelids. Based on similarities between the sclerites of Wiwaxia and the notochaetae of Canadia with the broad notochaetae (paleae) of Recent chrysopetalid polychaetes, these two fossil taxa have been variously treated as closely related to the most highly derived stem forms of the polychaete (and annelid) crown group or as members of a specific, Recent subgroup within Polychaeta, the order Phyllodocida. Chrysopetalidae is a member of Phyllodocida, which is part of the major polychaete clade Aciculata; the latter two taxa are distinguished by four and six well defined autapomorphic characters, respectively. The best preserved or otherwise appropriate fossils of Wiwaxia corrugata, Canadia spinosa and the other polychaetes of the Burgess Shale have been studied in detail in order to determine whether they possess any characters that could support the homology of wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae. Most of these fossil taxa have significant autapomorphies but the specific characters of the Aciculata and Phyllodocida are entirely absent. It is demonstrated that constraining cladograms in such a manner that wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae are homologous leads to results that are markedly unparsimonious. Furthermore, Canadia and the other polychaetes of the Burgess Shale cannot be referred to any extant subgroup within the Polychaeta and cannot be used to polarize character evolution within the annelid crown group. Apart from its dubious sclerites, Wiwaxia has no characters that could indicate any close relationship with Polychaeta or Annelida.     

Decline of the Burgess Shale fauna: ecologic or taphonomic restriction?

Aronson, R. B. 1992 07 15: Decline of the Burgess Shale fauna: ecologic or taphonomic restriction? An important alternative has not been tested in the debate over the origin, significance, and decline of Burgess Shale taxa: their observed stratigraphic distribution could be an artifact of increasing bioturbation. Based on information currently available, comparisons of the observed and expected stratigraphic distributions of Burgess Shale-type faunas falsify bioturbation as the exclusive cause. Ecological factors and/or taphonomic controls other than bioturbation were also responsible for the post-Cambrian decline of Burgess Shale-type faunas. Bioturbation, Burgess Shale fauna, soft-bodied animals, taphonomy.     

Phylogenetic Significance of the Burgess Shale Crustacean Canadaspis

The crustaceans, like the other major living groups of arthropods, have a long evolutionary history. The earliest examples occur in the Cambrian, and fossils of this age are a critical source of evidence of relationships both within the Crustacea, and between the Crustacea and other major arthropod groups. Canadaspis perfecta, from the Middle Cambrian Burgess Shale, is important as one of the oldest well-documented crustaceans. The evidence for reconstructing its remarkable combination of primitive and derived characters is reviewed, and its possible phylogenetic significance re-assessed.     

Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) ‘The enigmatic animal Opabinia regalis,Middle Cambrian,Burgess Shale,British Columbia’

Harry Whittington''s 1975 monograph on Opabinia was the first to highlight how some of the Burgess Shale animals differ markedly from those that populate today''s oceans. Categorized by Stephen J. Gould as a ‘weird wonder’ (Wonderful life, 1989) Opabinia, together with other unusual Burgess Shale fossils, stimulated ongoing debates about the early evolution of the major animal groups and the nature of the Cambrian explosion. The subsequent discovery of a number of other exceptionally preserved fossil faunas of Cambrian and early Ordovician age has significantly augmented the information available on this critical interval in the history of life. Although Opabinia initially defied assignment to any group of modern animals, it is now interpreted as lying below anomalocaridids on the stem leading to the living arthropods. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Cambrian stem-group annelids and a metameric origin of the annelid head

The oldest fossil annelids come from the Early Cambrian Sirius Passet and Guanshan biotas and Middle Cambrian Burgess Shale. While these are among the best preserved polychaete fossils, their relationship to living taxa is contentious, having been interpreted either as members of extant clades or as a grade outside the crown group. New morphological observations from five Cambrian species include the oldest polychaete with head appendages, a new specimen of Pygocirrus from Sirius Passet, and an undescribed form from the Burgess Shale. We propose that the palps of Canadia are on an anterior segment bearing neuropodia and that the head of Phragmochaeta is formed of a segment bearing biramous parapodia and chaetae. The unusual anatomy of these taxa suggests that the head is not differentiated into a prostomium and peristomium, that palps are derived from a modified parapodium and that the annelid head was originally a parapodium-bearing segment. Canadia, Phragmochaeta and the Marble Canyon annelid share the presence of protective notochaetae, interpreted as a primitive character state subsequently lost in Pygocirrus and Burgessochaeta, in which the head is clearly differentiated from the trunk.     

Review

Book reviews in this article:
Phyla, phylogeny and fantasy
Wonderful life. The Burgess Shale and the nature of history Stephen J. Gould.
Wonderful life. The Burgess Shale and the nature of history .—Stephen Jay Gould.     

贵州剑河中寒武世中期甲劳组水母状化石的发现

贵州剑河革东镇八郎村南甲劳组中部钙质白云岩中新近发现的具有粗强放射状辐管的水母状化石和其下凯里组中具细辐管及同心环的拟轮盘水母(Pararotadiscus)有明显区别,属于伊尔东钵科(Eldoniidae)。其层位晚于中寒武世早期凯里生物群中拟轮盘水母(Pararotadiscus)层位1个化石带,属中寒武世中期,是该类化石在中国的最高层位,相当于北美布尔吉斯页岩生物群中Eldonia化石层位。     

The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group

Opabinia regalis Walcott is an enigmatic fossil from the Middle Cambrian Burgess Shale of uncertain affinities. Recent suggestions place it in a clade with Anomalocaris Whiteaves from the Burgess Shale and Kerygmachela Budd from the Greenlandic Sirius Passet Fauna; these taxa have been interpreted as 'lobopods'. Consideration of available Opabinia specimens demonstrates that reflective extensions from the axial region, previously thought to be either gut diverticula or musculature, can be accommodated in neither the trunk nor the lateral lobes that arise from it. They must therefore be external structures independent of the lateral lobes. On the basis of their sub-triangular appearance, size and taphonomy, they are considered here to represent lobopod limbs. Some evidence for the existence of terminal claws is also presented. The question of whether Kerygmachela, Opabinia and Anomalocaris constitute a monophyletic or paraphyletic grouping is considered. While they share several characters, most of these are plesiomorphies. Further, Opabinia and Anomalocaris share several arthropod-like characters not possessed by Kerygmachela. It is concluded that these three taxa probably form a paraphyletic grouping at the base of the arthropods. Retention of lobopod-like characters within the group provides important documentation of the lobopod-arthropod transition. A proper understanding of Opabinia and its close relatives, which may include the tardigrades, opens the way for a reconstruction of the arthropod stem-group. This in turn allows the construction of a speculative but satisfying scenario for the evolution of major arthropod features, including the origin of the biramous limb, tergites and arthropod segmentation. 'Arthropodization' may thus be seen not to be a single event but a series of adaptive innovations. OPABINIA, ANOMALOCARIS, KERYGMACHELA, Burgess Shale, problematica, Lobopodia, Arthropoda.     

The macro‐ and microfossil record of the Cambrian priapulid Ottoia

The stem‐group priapulid Ottoia Walcott, 1911, is the most abundant worm in the mid‐Cambrian Burgess Shale, but has not been unambiguously demonstrated elsewhere. High‐resolution electron and optical microscopy of macroscopic Burgess Shale specimens reveals the detailed anatomy of its robust hooks, spines and pharyngeal teeth, establishing the presence of two species: Ottoia prolifica Walcott, 1911, and Ottoia tricuspida sp. nov. Direct comparison of these sclerotized elements with a suite of shale‐hosted mid‐to‐late Cambrian microfossils extends the range of ottoiid priapulids throughout the middle to upper Cambrian strata of the Western Canada Sedimentary Basin. Ottoiid priapulids represented an important component of Cambrian ecosystems: they occur in a range of lithologies and thrived in shallow water as well as in the deep‐water setting of the Burgess Shale. A wider survey of Burgess Shale macrofossils reveals specific characters that diagnose priapulid sclerites more generally, establishing the affinity of a wide range of Small Carbonaceous Fossils and demonstrating the prominent role of priapulids in Cambrian seas.     

英文贵州中寒武世凯里生物群中宏观藻类化石新材料(英文)

贵州省台江县中寒武世凯里生物群含有丰富的非钙质藻类和具有软躯体后生动物化石 ,它为布尔吉斯页岩型生物群在世界广泛分布提供了更有力的证据。在生物群的宏观藻类中描述了 5个属 5个种 ,包括 2个新属。它们是MarpoliaspissaWalcott、AcinocricusstichusConwayMorrisandRobison、UdotealgaerectaYang、EosargassumsawataYang和RhizophytonzhaoyuanlongiiYang ,并且将凯里生物群中的宏观藻类化石组合与加拿大布尔吉斯页岩生物群中的宏观藻类进行了对比 ,发现两个生物群不仅具有相似的动物化石组成 ,而且宏观藻类化石组成也很相似。     

New Macroalgal Fossils from Middle Cambrian Kaili Biota in Guizhou Province,China (in English)

Abundant and well-preserved remains of noncalcareous algae and soft-bodied metazoans were collected from Middle Cambrian Kaili biota in Taijiang county, Guizhou Province, China. These remains provide further evidence for the wide geographic distribution of many Burgess Shale taxa. Among the algae, 5 genera (including two new genera) and 5 species are described. They are Marpolia spissa Walcott, Acinocricus stichus Conway Morris and Robison, Udotealga erecta Yang, Eosargassum sawata Yang, and Rhizophyton zhaoyuanlongii Yang. Contrasting the macroalgal fossil assemblage in the Kaili biota with one in the Burgess Shale biota, it is clear that similarity of the Kaili biota and the Burgess Shale biota is reflected by the same content of not only the soft-bodied metazoans, but also the noncalcareous algae.