Decay of Branchiostoma: implications for soft-tissue preservation in conodonts and other primitive chordates

Decay experiments on the cephalochordate Branchiostoma lanceolatum ('amphioxus') demonstrate that the most decay resistant structures are the notochord sheath and the cartilaginous rods which support the gill bars. However, even more labile soft parts, such as the muscles and skin may survive for at least 124 days under totally anoxic conditions. As the chevron-shaped muscles of the myomeres shrink and collapse, those on opposite sides of the trunk maybe displaced, resulting in pronounced offsetting. Only 1.42% of the initial dry weight of Branchiostoma is resistant to alkali and acid hydrolysis, compared to 46% in the polychaete Nereis virens. Branchiostoma is only likely to be fossilized as a result of decay inhibition and replication by early diagenetic minerals. The results of these experiments cast light on the interpretation of a number of primitive fossil chordates. There is no reason to infer extracellular decay-resistant cuticle in the Burgess Shale Pikaia. The axial lies preserved in the conodont animal specimens from the Carboniferous of Edinburgh, Scotland, represent the notochord. The displacement of the elements to one side of the head reflects the true position of the apparatus - the surrounding tissue has been lost through decay. The chevron-shaped structures in the Carboniferous chordate Conopiscius are the muscles of the myomeres, not external scales. The lines delineating the segments in the Silurian Jamoytius most likely represent the myosepta. There is some doubt about the nature of the only specimen interpreted as a fossil cephalochordate, Palaeobranchiostoma hamatotergum from the Permian of South Africa. □ Taphonomy, decay, softparts, Cephalochordata , Branchiostoma, lancelet, Chordata , Pikaia, conodont , Conopiscius, Jamoytius, Palaeobranchiostoma.     

日本志留纪至三叠纪牙形刺生物地层

本文系统地回顾了日本志留纪至三叠纪牙形刺研究的历史和现在的成果。志留纪牙形刺只有少数零星的报道,没有建立化石带;早泥盆世已建立了５个牙形刺组合,没有中、晚泥盆世的记录;石炭纪有８个牙形刺带,其中晚石炭世３个牙形刺带;二叠纪５个牙形刺带或动物群,其中,中、晚二叠世各１个带（动物群）;三叠纪可划分出１４个牙形刺带。     

Soft anatomy and the affinities of conodonts

Recent claims that conodonts are members of the Craniata or Vertebrata are based in part upon soft tissue features that have been preserved in a small number of specimens. These features include what appear to be radials in the caudal fin and paired structures that have been identified as eye remnants. The evidence for radials is limited, but credible. However, the anatomy of extant cyclostomes suggests that the paired structures are more reasonably interpreted as otic capsules than the remnants of sclerotic eye capsules. Moreover, even if these structures are the remnants of eyes, conodonts might equally well be a sister group to the craniates as a member of that group. Aside from these paired structures, conodont fossils exhibit no features that are suggestive of a cartilaginous skeleton. Given that cyclostome fossils sometimes show evidence of the cartilages of the head, the apparent absence of a similar skeleton in conodont animals calls into question the claim that they are craniates. The simple single chevron shape of conodont myomeres also suggests that they lie outside of the Craniata. All living craniates have double-chevron myomeres as adults, whereas simple myomeres of the conodont type are found in the non-craniate cephalochordates. Thus the available soft tissue evidence suggests that conodonts are best regarded as the sister group of the craniates.     

Conodont anatomy, chordate phylogeny and vertebrate classification

Interpretations of conodont anatomy and affinity continue to generate controversy. Fossilized soft-tissue evidence indicates that conodonts possessed eyes, extrinsic eye muscles, a notochord, myomeres, a differentiated tail with fin radiais, possible otic capsules and possible branchial structures. Indirect evidence suggests a differentiated brain and cartilaginous head skeleton. The multi-component phosphatic tissue complexes of the conodont feeding apparatus cannot be compared to the amorphous apatite of extant agnathan otoliths. By limiting cladistic analysis to a restricted selection of these characters the hypothesis that conodonts are a sister group of the clade comprising extant hagfish, lampreys and gnathostomes can be supported. However, exhaustive analysis of a more complete character-set strongly supports the hypothesis that conodonts are more derived than hagfish. From a taxonomic perspective, these two hypotheses have no effect on how conodonts should be classified. Whether they are a stem group (the former hypothesis) or part of the crown group (the latter), conodonts are clearly part of the total group Vertebrata (=Craniata).     

辽河断陷盆地西部晚寒武世至早奥陶世牙形石生物地层

辽河断陷盆地西部凹陷古潜山顶部地层的时代 ,自 70年代末期以来 ,一直被认为属于中、上元古代。本文通过在总面积为 60 0 km2 的西部凹陷中段系统、详尽的牙形石生物地层研究 ,并结合疑源类、小壳化石以及岩石地层学资料 ,确认研究区内古潜山顶部地层的时代主要为晚寒武世至早奥陶世。中、上元古代地层在研究区内虽然存在 ,但分布极其局限。从而 ,彻底改变了2 0年以来对该地区古潜山地层划分和对比上的传统认识。     

Conchodontus,Mitrellataxis and Fungulodus: Conodonts,fish or both?

Conchodontus, Mitrellataxis and Fungulodus are phosphatic microfossils from the Late Devonian of China and North America, alternatively interpreted as conodont elements or fish scales. The histology and microornament of these sclerites have been studied in an attempt to resolve their affinity, and to determine characters for distinguishing between conodont elements and the ichthyoliths of other lower vertebrates. The histology of all three genera is directly comparable to conodont elements, dispelling the notion that conodonts are histologically indistinguishable from the teeth and scales of other vertebrates. Microornament is found not to be useful in discriminating between high-level taxonomic groupings. White matter and thickness of prismless enamel are suggested as apomorphies of the Conodonta.     

Zur Biologie, Taxonomie und Chronologie der Conodonten

The actual state of knowledge concerning the conodont animal is critically reviewed. All soft tissue documented by the specimens of the conodont animal were interpreted by the British collectors as Craniota (Vertebrata) features. Opinions on the precise affinities, however, remain controverse: The alleged chorda could have been a gut, bilateral symmetry, myomery, apatitic biomineralisation occur also in Invertebrata, caudal fins do in Chaetognatha. Even the lobes in the capital area interpreted as sensory organs could have served as aiding organs for food intake. Therefore, the originally sensational findings are reduced to the core issue that the conodont animal had an eel-like body and at the blunt end of it, behind peculiar lobes, the mineralized conodont elements were arranged. As a consequence, a relationship to the Protochordata cannot be excluded. A systematic position of the conodont animal as possible Vertebrata is likely only if organized below the Ostracodermata which is particularly so because conodonts occur so early in the Cambrian. In the second part of the article conodonts are discussed as guide fossils. The comprehensive, empirically grown taxonomy of the single elements is the prerequisite for recognizing outstanding, high resolution stratigraphy. In comparison, a multielement taxonomy remains as long problematical, as it is based on assumed, mostly statistically reconstructed rather than on natural multielement apparatuses. Taxonomical and nomenclatorial operations built on assumed apparatus affinity can bear heavily on nomenclature and stratigraphy of platform elements. It can be shown at the example of the Devonian that the platform elements have provided particularly in the pelagic facies a Standard Conodont Zonation that covers time almost without any gaps. The basis for this is that platform single element species and their phylogenetic development are extra-well known. In a special paragraph, history and applicability of the Standard Zonation is discussed in details. Different conodont biofacies are developed between the pelagic realm where the Standard Zonation originated, and the coastal areas so that conodonts can indicate temporal and bathymetric relationships of the mother rocks. In addition alternative conodont zonations were introduced mainly for the shallow water. Because of their conspicuous stratigraphic significance in the Devonian, conodonts were employed by International Committees to redefine Series and Stage Boundaries. For the boundary selection species were utilized from phylognetic lineages as only from tight evolution their point of origination could be well identified and recognized in worldwide correlation. The GSSPs (Global Stratotype Section and Point) within the Devonian are discussed in detail. In recent times, attempts are made at calibrating the duration of conodont zones on the basis of their well known rate of evolution and high temporal resolution. These analyses have to be continued in order to demonstrate the absolute time relationship of condont zones in more and continuous detail. Currently the phyletically controlled Standard Conodont Zonation is the authoritative stratigraphic method in the Devonian. It has provided together with the GSSPs precise and valuable fix points for a Global Time Scale (GTS). In combination with additional stratigraphie methods the Standard Conodont Zonation may provide further refinements in the future.     

Late Devonian Conodonts from the Zeravshan-Gissar Mountainous Area,Uzbekistan

Late Devonian conodonts, recovered from the siliceous and carbonate rocks of the upper part of the Akbasay Formation on the left side of the Kule Gorge, Zeravshan-Gissar mountainous area (Uzbekistan), are studied. Seven conodont associations are defined for the Frasnian–Famennian interval.     

Conodont element function

There are close similarities between conodont elements and teeth both in general shape and in that they possess pointed tips and have expanded bases with more porous tissue. A number of examples of conodont elements which parallel specific kinds of tooth shapes and organizations are added to earlier known similarities. Both in teeth and in conodont elements the surface structures include cutting edges, striations, and barbs. The change in strength of the conodont denticles caused by the evolution of white matter is also shown to agree with a tooth function. On the other hand, the elements grew throughout the life of the animal by lamellae added to the surface. The solution of this paradox is found in the elements alternating between a growth phase and a functional phase. During growth the oral surface of the elements was enveloped in folds of secreting soft tissue. Structures henceforth termed burrs were formed at the contacts between the folds. Parts of the burrs evolved into cutting edges. Three different bite types occurred among the conodont elements. Many (all?) conodonts were predators which used their elements to seize and to process the food mechanically. The shape of the conodont elements cannot be used for conclusions regarding the affinities of the conodonts. Similarly, an identification of a fossil as belonging to the conodonts must be supported by other evidence than just shape.     

广西桂林泥盆纪牙形刺组合序列与海平面变化

本文以桂林灵川岩山圩中,晚泥盆世界线剖面,桂林沙河能源疗养院背后信都组－唐家湾组剖面及桂林额头村Ｄ／Ｃ界线剖面的牙形刺资料为基础,结合前人在桂林地区所做的牙形刺生物地层工作,识别出该区中,晚泥盆世牙形刺化石的１６个组合序列,并以此为尺度结合层序地层格架,层序界面及层序中的微相叠加形式,讨论该区中,晚泥盆世高分辨的海平面变化。     

Evolution of Permian conodont provincialism and its significance in global correlation and paleoclimate implication

The main components of Asselian through Artinskian conodont faunas found around the world are basically the same, and the provincialism is indicated only by less common endemic elements such as Gondolelloides and New Genus A Henderson in North Pangea, Sweetognathus bucaramangus around the equator and Vjalovognathus in eastern Gondwana. Provincialism is marked by differences at the species level of Mesogondolella, Neostreptognathodus and Sweetognathus during the Kungurian, and becomes very distinct with differences at the genus level during the Guadalupian and Lopingian. Three provinces of Permian conodonts, referred to as the North Cool Water Province (NCWP), the Equatorial Warm Water Province (EWWP) and the peri-Gondwana Cool Water Province (GCWP), are recognized. The NCWP is marked by Gondolelloides in the early Cisuralian, dominance of Neostreptognathodus and no or rare Sweetognathus in the late Cisuralian, dominance of Merrillina and Mesogondolella and absence of Sweetognathus in the Guadalupian, and dominance of Merrillina and Mesogondolella and absence of Iranognathus in the Lopingian. The EWWP is characterized by the absence of Gondolelloides and Vjalovognathus in the Cisuralian, abundance of Sweetognathus and Pseudosweetognathus in the Kungurian (late Cisuralian), Jinogondolella and Sweetognathus in the Guadalupian, and Clarkina and Iranognathus in the Lopingian. The GCWP is marked by Vjalovognathus, Merrillina in the Cisuralian, Vjalovognathus, Merrillina and Mesogondolella in the Guadalupian, and Vjalovognathus and Merrillina in the Lopingian. Mixed faunas are recognized in regions bordering between the EWWP and GCWP including Western Timor during the Artinskian, Pamirs during the Kungurian and the Salt Range during the Guadalupian and Lopingian.

Three different conodont zonations are proposed, one for each of the three conodont provinces. Four potential horizons for inter-provincial correlation of Permian conodonts are recognized. They are in ascending order: (1) the first appearance of Sweetognathus whitei, which is closely related to the last occurrence of Carboniferous-type conodonts such as Streptognathodus and Adetognathus; (2) the first appearance of Neostreptognathodus pequopensis; (3) the base of the Jinogondolella nankingensis Zone; and (4) the base of the Clarkina postbitteri–Iranognathus erwini Zone.

The spatial and temporal distribution pattern of Permian conodonts suggests that temperature is the primary controlling factor. Evolution of Permian conodont provincialism reveals a glaciation during the Asselian and Sakmarian, a global warming during the Artinskian, a climate cooling in North Pangea during the Kungurian, a continuation of Kungurian climate trends in the Guadalupian, a relatively minor warming during the Wuchiapingian, a returned cooling in the Changhsingian and Lower and Middle Griesbachian, and a global warming in the Late Griesbachian, which ended the Permian conodont lineage.     

Evolution of morphogenesis in 360‐million‐year‐old conodont chordates calibrated in days

SUMMARY Highly rhythmic increments of crown tissue are identifiable in conodont oral apparatus elements from the Late Devonian of the Holy Cross Mountains, Poland; individual laminae being of thickness comparable with daily increments of vertebrate tooth enamel and fish otoliths. Abundant occurrence of such specimens enables bed‐by‐bed (stratophenetic) studies of the process of evolution at the population level and quantitative presentation of the evolution of ontogeny in the sampled geological section covering several million years. The morphologic transformation is expressed as expansion of a juvenile asymmetry to later stages of the ontogeny and in decrease of the mature element width, which was due to a change of the mineral tissue secretion rate. It was not just a simple extension of a juvenile character into the later stage of the ontogeny (heterochrony) but rather a true developmental novelty. The evolution was gradual and very slow. The proposed quantitative approach to growth increments in the mineral skeleton of ancient chordates introduces real‐time units to evolutionary developmental studies connected with direct paleontological evidence on the course of evolution.     

Evidence of parallel evolution in the dental elements of Sweetognathus conodonts

The repeated emergence of similar morphologies in the dental elements of Permian Sweetognathus conodonts has been a hypothesized example of parallel evolution. To test if morphological parallelisms occur between isolated Sweetognathus lineages, this study uses two-dimensional-based geometric morphometrics combined with a revised and expanded phylogeny of Permian Sweetognathus conodonts to quantify dental element trait distributions and compare the phenotypic trajectories between lineages. A hierarchical clustering method was used to identify recurrent species pairs based on principal component scores describing their morphological variation, with the further incorporation of widely used ecological metrics such as limiting similarity and morphological overlap. Our research implies that a major contributor to conodont diversity in Palaeozoic marine trophic networks is the emergence of recurrent parallel morphologies via disruptive and directional selection. This study illustrates the mechanisms through which conodonts achieved their status as hyper-diverse predators and scavengers, contributing substantially to the complexity of Palaeozoic marine communities.     

The sharpest tools in the box? Quantitative analysis of conodont element functional morphology

Conodonts have been considered the earliest skeletonizing vertebrates and their mineralized feeding apparatus interpreted as having performed a tooth function. However, the absence of jaws in conodonts and the small size of their oropharyngeal musculature limits the force available for fracturing food items, presenting a challenge to this interpretation. We address this issue quantitatively using engineering approaches previously applied to mammalian dentitions. We show that the morphology of conodont food-processing elements was adapted to overcome size limitations through developing dental tools of unparalleled sharpness that maximize applied pressure. Combined with observations of wear, we also show how this morphology was employed, demonstrating how Wurmiella excavata used rotational kinematics similar to other conodonts, suggesting that this occlusal style is typical for the clade. Our work places conodont elements within a broader dental framework, providing a phylogenetically independent system for examining convergence and scaling in dental tools.     

Traces of microboring organisms in Palaeozoic conodont elements

Conodont elements from Palaeozoic strata (Ordovician, Devonian and Carboniferous) contain abundant evidence for presumed post-mortem colonisation by endolithic organisms. It is possible to distinguish several morphotypes of microcavities in conodont apatite, which show some morphological similarities to microborings in calcareous substrates. Some of them are believed to be produced by microendoliths, some others are yet unknown from calcareous substrates. The present report deals with the various kinds of microendolithic traces in conodont apatite. The significance of microborings in conodont elements is discussed.     

Step-wise morphological trends in fluctuating environments: Evidence in the Late Devonian conodont genus Palmatolepis

Long-term morphological changes were investigated in the conodont genus Palmatolepis, using a Fourier analysis of the outline of platform elements. Three time-slices and four Late Frasnian - Early Famennian sections on North Gondwana were studied. No difference existed between locations, but shape variations were recorded associated with the successive Kellwasser events. The Middle Famennian period was analyzed in a section located at the Baltica margin across the annulata event. Some morphological changes may be related to the deposition of the corresponding black shales, but the main feature is a heterogeneity of the populations, which can be split into two morphological types. Latest Famennian conodonts from another North Gondwanan section show only minor shape changes during this time interval, which is environmentally stable prior to the global Hangenberg event. Based on synthetic shape variations across the whole period an morphological trend towards narrower elements is observed. This morphological change happens step-wise in response to global events. The amplitude of the shape shift is proportionally related to the extend of the successive Late Devonian global events.     

Late Devonian dimpled phosphatic microspherules associated with conodonts are possible otoliths of short‐lived and opportunistic organisms

Dimpled phosphatic microspherules, contradictorily associated with conodonts, are widely distributed in strata ranging in age from the Cambrian to Carboniferous. These microspherules have attracted much attention from palaeobiologists and were suggested to be ‘conodont pearls’, ‘conodont otoliths’ or ‘fish otoliths’ due to their similar chemical composition and co‐occurrence with conodonts or fish teeth. However, these hypotheses are still highly controversial. Here, we report ‘checks’, ‘rhythmic growth patterns’ and ‘sub‐diurnal increments’ from growth annuli of the Late Devonian phosphatic microspherules from South China, on the basis of quantitative microstructure analysis. The annulus width of phosphatic microspherules becomes narrower with increasing radius. These microstructural characteristics of growth annuli are most typical indicators of modern animal otoliths. In addition, a maximum value of about 90 annuli is encountered from all the specimens. We propose that these microspherules are essentially phosphatic otoliths, which might have been secreted by a specific kind of marine organisms with very short lifespans (less than 90 days). Furthermore, the sudden enrichment of phosphatic microspherules in the late Frasnian may represent a biological response of short‐lived animals to ecological crisis such as ocean eutrophication.     

Euconodont hard tissue: preservation patterns of the basal body

Conodont elements, consisting of crown and basal tissue are the well-known fossilized hard parts of Conodonta (extinct marine chordates), but the taphonomic processes leading to decomposition or remineralization of the basal tissue are not well understood. Here we focus on the taphonomy of basal tissue, reviewing the published record and describing new material from Asia and Europe (248 occurrences globally). These include crown and basal tissue in conjunction, and isolated basal bodies showing different stages of preservation. Some isolated specimens resemble phosphate rings similar to those assigned to Phosphannulus universalis. High-resolution biostratigraphy indicates that the lamellar type of conodont basal tissue is found in all facies and depositional environments. Other basal tissue types, described in the literature as tubular, mesodentine, spherulitic or lamellar with canalules, are limited to the early Palaeozoic and found exclusively in siliciclastic deposits (with the exception of spherulitic tissue). Although the stratigraphic record of basal tissue spans the range of Euconodonta (Cambrian–Triassic), this study shows that most of the isolated plate and ring-like structures are derived from early Palaeozoic coniform conodonts. Basal tissue of platform-type elements has a much more fragile shape and is therefore rarely preserved as a recognizable isolated unit.     

Finite element,occlusal, microwear and microstructural analyses indicate that conodont microstructure is adapted to dental function

Conodonts constitute the earliest evidence of skeletal biomineralization in the vertebrate evolutionary lineage, manifest as a feeding apparatus of tooth‐like elements comprised of enamel‐ and dentine‐like tissues that evolved in parallel with these canonical tissues in other total‐group gnathostomes. As such, this remarkable example of evolutionary parallelism affords a natural experiment in which to explore the constraints on vertebrate skeletal evolution. Using finite element analysis, informed by occlusal and microwear analyses, we tested the hypothesis that coincidence of complex dental function and microstructural differentiation in the enamel‐like tissues of conodonts and other vertebrates is a consequence of functional adaptation. Our results show topological co‐variation in the patterns of stress distribution and crystallite orientation. In regions of high stress, such as the apex of the basal cavity and inner parts of the platform, the crown tissue comprises interwoven prisms, discontinuities between which would have acted to decussate cracks, preventing propagation. These results inform a general occlusal model for platform conodont elements and demonstrate that the complex microstructure of conodont crown tissue is an adaptation to the dental functions that the elements performed.     

New Middle Devonian conodont data from the Dong Van area,NE Vietnam (South China Terrane)

Middle Devonian conodonts from the Si Phai section in NE Vietnam are described. The section ranges from the Middle Devonian ensensis to timorensis conodont zones to the Late Devonian rhomboidea conodont Zone. A rich overall assemblage is described, including 27 taxa of species or subspecies rank and 11 taxa described in an open nomenclature. Among the dominant Polygnathus forms, four new taxa are described: Polygnathus linguiformis saharicus subsp. nov., Polygnathus linguiformis vietnamicus subsp. nov., Polygnathus rhenanus siphai subsp. nov., and Polygnathus xylus bacbo subsp. nov. Conodont assemblages are attributed to polygnathid, polygnathid-klapperinid, and klapperinid conodont biofacies representing hemipelagic to pelagic environments. The klapperinid biofacies, unreported in the previous literature, are here attributed to offshore areas of the external shelf. The taxonomic compositions of the studied conodont assemblages, as well as their CAI characteristics (CAI 4–5), suggest a palaeogeographic affinity of the studied strata to the Chinese Devonian Guangxi Basin, and the South China Terrane in general. Furthermore, the conodont biofacies and the palaeogeographic distribution of the fauna are discussed.