Jean Vannier

TUZOIA: MORPHOLOGY AND LIFESTYLE OF A LARGE BIVALVED ARTHROPOD OF THE CAMBRIAN SEAS

Abstract For almost 30 years, paleontologists have analyzed evolutionary sequences in terms of simple null models, most commonly random walks. Despite this long history, there has been little discussion of how model parameters may be estimated from real paleontological data. In this paper, I outline a likelihood-based framework for fitting and comparing models of phyletic evolution. Because of its usefulness and historical importance, I focus on a general form of the random walk model. The long-term dynamics of this model depend on just two parameters: the mean (μstep) and variance (σ2step) of the distribution of evolutionary transitions (or “steps”). The value of μstep determines the directionality of a sequence, and σ2step governs its volatility. Simulations show that these two parameters can be inferred reliably from paleontological data regardless of how completely the evolving lineage is sampled. In addition to random walk models, suitable modification of the likelihood function permits consideration of a wide range of alternative evolutionary models. Candidate evolutionary models may be compared on equal footing using information statistics such as the Akaike Information Criterion (AIC). Two extensions to this method are developed: modeling stasis as an evolutionary mode, and assessing the homogeneity of dynamics across multiple evolutionary sequences. Within this framework, I reanalyze two well-known published data sets: tooth measurements from the Eocene mammal Cantius, and shell shape in the planktonic foraminifera Contusotruncana. These analyses support previous interpretations about evolutionary mode in size and shape variables in Cantius, and confirm the significantly directional nature of shell shape evolution in Contusotruncana. In addition, this model-fitting approach leads to a further insight about the geographic structure of evolutionary change in this foraminiferan lineage.

Early Cambrian Food Chain: New Evidence from Fossil Aggregates in the Maotianshan Shale Biota, SW China

Abstract Three categories of fossil aggregates are recognized in the Lower Cambrian Maotianshan Shale biota from SW China: (1) elliptical aggregates with randomly distributed exoskeletal remains of typically small- to medium-size bivalved arthropods (e.g., ostracode-like bradoriids, phyllocarid-like waptiids), hyoliths, and trilobites are interpreted as coprolites, possibly produced by anomalocaridids (e.g., waptiid-rich coprolites) and/or unknown epibenthic predators; (2) elongate, ribbon-like aggregates composed of oriented small hyolith shells, interpreted as the feces of infaunal carnivorous worms such as priapulids; and (3) concentric aggregates, typically with a central nucleus (e.g., remains of medusoid eldoniids or bivalved arthropod carapaces) and peripheral exoskeletal fragments, possibly generated by bottom currents whirling around carcasses. These new coprolite data add to morphofunctional information obtained from fossil organisms and indicate that predation occurred at different levels of the water column with: (1) endobenthic predators (diverse priapulid fauna) feeding near the sediment-water interface; (2) epibenthic predators/scavengers (almost exclusively arthropods); (3) predators living in the lower levels of the water column (e.g., anomalocaridids); and (4) mid-water predators exploiting upper levels in the water column (e.g., eldoniids, ctenophores, chaetognaths). Communities living at or close to the water-sediment interface (epibenthic sensu stricto, meiobenthic, and demersal animals) were exposed to a multidirectional predatorial pressure from infaunal, epifaunal, and mid-water predators. Although predation was diverse, nothing indicates that the food chain extended beyond the level of primary carnivores. Animals already had acquired complex behaviors such as hunting (e.g., anomalocaridids, priapulids) and predator avoidance in which sensory systems were involved. The example of the Maotianshan Shale indicates that the burst of anatomical innovations (new body plans) that characterizes the early Cambrian also was accompanied by the rapid development of new feeding strategies and by an unprecedented expansion of ecological interactions (prey-predator relationships).

Priapulid worms: Pioneer horizontal burrowers at the Precambrian-Cambrian boundary

The major evolutionary events that characterize the Precambrian–Cambrian transition are accompanied by profound ecological changes in the composition of benthic communities, the nature of the substrate, and the occupation of marine ecospace. The increased animal activity on and within the substrate is attested to by numerous trace fossils, such as the cosmopolitan Treptichnus pedum whose fi rst appearance is used as the global stratotype section and point (GSSP) to mark the base of the Cambrian. In spite of its major importance in biostratigraphy, the maker of Treptichnus trace fossils, and more generally of treptichnids, has long remained an enigma. Treptichnids were subhorizontal burrow systems produced in the subsurface and had a worldwide distribution throughout the Cambrian. Here we show, by using experimental ichnology, that the treptichnid burrow systems were most probably produced by priapulid worms or by worms that used the same locomotory mechanisms as the Recent priapulids (e.g., Priapulus). Their typical three-dimensional morphology with repeated arcuate probing branches suggests that their function was related to the feeding strategy of the worm such as predation or scavenging upon small epibenthic or endobenthic invertebrates. This interpretation is strongly supported by the preserved gut contents of Cambrian priapulids from the Burgess Shale Lagerstatte that contain effectively a variety of small epibenthic prey. The antiquity of treptichnids would designate priapulids as one of the earliest infaunal colonizers of the substrate that possibly interacted with epibenthic communities, thus playing a leading role (1) in the construction of the early marine food chain, and (2) as important subhorizontal bioturbators in the early stages of the “Cambrian Substrate Revolution.”

Silurian Myodocopes: Pioneer pelagic ostracods and the chronology of an ecological shift

Analysis of all relevant palaeontological and global geological data strongly supports the notion that representatives of Silurian myodocope ostracods had pelagic lifestyles and habitats and that they may well be, within the Ostracoda, pioneer colonisers of such environments. Morphological evidence (from fossil and Recent myodocopes) combined with facies distributional and concomitant faunal evidence (from the Silurian of, for example, Britain, France, Czechoslovakia, Sardinia, Australia and China) endorses the idea that myodocope ostracods may have undergone a benthic to pelagic ecological shift during mid Silurian times. Lower Silurian myodocopes lived, with benthic associates, on well oxygenated shelves. Upper Silurian ostracods lived, typically with low diversity, largely pelagic faunas in outer shelf topographic lows or off-shelf basin slopes, and are characteristically associated with deposits which are in part suggestive of lowered oxygen levels or even anoxic conditions. A pre-adaptation for swimming may have allowed Silurian myodocopes to respond to environmental forcing (negative oxygen levels; positive trophic and nutrient incentives; rises in sea levels) by migrating, through time, up the water column.

The Early Cambrian colonization of pelagic niches exemplified by Isoxys (Arthropoda)

The anatomy of the bivalved arthropod Isoxys (Early and Middle Cambrian) is reconstructed, based on new evidence from soft parts and exoskeletal design and on a critical review of previous work. Isoxys had a long segmented body flanked with a pair of short antennules, followed by a series of 14 biramous appendages provided with long paddle-like exopods concealed under a widely open bivalved carapace folded dorsally and bearing long cardinal spines. The close resemblance between Isoxys and Recent pelagic crustaceans (halocyprid ostracods, larval stages of malacostracans) indicates that Isoxys was probably an active epipelagic swimmer (evidence from soft parts, carapace design and distributional pattern). Some species (e.g. I. auritus and I. paradoxus from the Maotianshan Shale biota; Early Cambrian) may have lived in the vicinity of the bottom either permanently or temporarily, whereas others may have had ecological preferences for more open-marine settings. The spinosity of Isoxys had a possible role in predatorial deterrence rather than in buoyancy control or in retarding sinking within the water column. The presence of Isoxys in the Maotianshan Shale of S. China indicates that arthropods had already colonized midwater niches by the Early Cambrian. The midwater communities of the Maotianshan Shale comprised numerous other invertebrates, such as abundant medusiform eldonids, vetulicolids, chordates and possibly early vertebrates. This contradicts the opinion that pelagic communities remained poorly developed until late Cambrian/Ordovician times and that the occupation of the midwater niches largely post-dates the initial diversification of the benthic faunas.

Arthropod visual predators in the early pelagic ecosystem: evidence from the Burgess Shale and Chengjiang biotas

Exceptional fossil specimens with preserved soft parts from the Maotianshan Shale (ca 520 Myr ago) and the Burgess Shale (505 Myr ago) biotas indicate that the worldwide distributed bivalved arthropod Isoxys was probably a non-benthic visual predator. New lines of evidence come from the functional morphology of its powerful prehensile frontal appendages that, combined with large spherical eyes, are thought to have played a key role in the recognition and capture of swimming or epibenthic prey. The swimming and steering of this arthropod was achieved by the beating of multiple setose exopods and a flap-like telson. The appendage morphology of Isoxys indicates possible phylogenetical relationships with the megacheirans, a widespread group of assumed predator arthropods characterized by a pre-oral ‘great appendage’. Evidence from functional morphology and taphonomy suggests that Isoxys was able to migrate through the water column and was possibly exploiting hyperbenthic niches for food. Although certainly not unique, the case of Isoxys supports the idea that off-bottom animal interactions such as predation, associated with complex feeding strategies and behaviours (e.g. vertical migration and hunting) were established by the Early Cambrian. It also suggests that a prototype of a pelagic food chain had already started to build-up at least in the lower levels of the water column.

Gut Contents as Direct Indicators for Trophic Relationships in the Cambrian Marine Ecosystem

Present-day ecosystems host a huge variety of organisms that interact and transfer mass and energy via a cascade of trophic levels. When and how this complex machinery was established remains largely unknown. Although exceptionally preserved biotas clearly show that Early Cambrian animals had already acquired functionalities that enabled them to exploit a wide range of food resources, there is scant direct evidence concerning their diet and exact trophic relationships. Here I describe the gut contents of Ottoia prolifica, an abundant priapulid worm from the middle Cambrian (Stage 5) Burgess Shale biota. I identify the undigested exoskeletal remains of a wide range of small invertebrates that lived at or near the water sediment interface such as hyolithids, brachiopods, different types of arthropods, polychaetes and wiwaxiids. This set of direct fossil evidence allows the first detailed reconstruction of the diet of a 505-million-year-old animal. Ottoia was a dietary generalist and had no strict feeding regime. It fed on both living individuals and decaying organic matter present in its habitat. The feeding behavior of Ottoia was remarkably simple, reduced to the transit of food through an eversible pharynx and a tubular gut with limited physical breakdown and no storage. The recognition of generalist feeding strategies, exemplified by Ottoia, reveals key-aspects of modern-style trophic complexity in the immediate aftermath of the Cambrian explosion. It also shows that the middle Cambrian ecosystem was already too complex to be understood in terms of simple linear dynamics and unique pathways.

The earliest ostracods: the geological evidence

The oldest assumed ostracods appear in the fossil record from the TremadocianPaltodus deltifer conodont Biozone. Although geographically widespread these early ostracods have no obvious Cambrian antecedents. Their first appearance at ca. 485 Ma contrasts with molecular evidence that suggests a much earlier (latest Proterozoic or Cambrian) origin for ostracods. Some Cambrian bivalved arthropods such asAltajanella andVojbokalina, conventionally referred to the Bradoriida, have carapace morphologies that resemble Ordovician palaeocopid ostracods, though such a relationship is unproven without soft part anatomy. Evidence from preserved soft anatomy demonstrates that Bradoriida, such asKunmingella, and Phosphatocopida, essentially the Cambrian ‘ostracod’ record of traditional usage, belong outside the Eucrustacea. Early Ordovician ostracods appeared first in shallow marine, oxygenated environments on shelf margins, in a similar setting to other elements of the ‘Paleozoic fauna’. Their biodiversity was low (3 named genera and ca. 12 species), though some taxa such asNanopsis andEopilla achieved widespread dispersal between major Ordovician palaeocontinents. As bradoriids were largely extinct by the Late Cambrian, ostracods do not appear to have directly competed with them for shallow marine environments. The rapid colonisation of these settings by ostracods may have been facilitated by the available ecospace vacated by Bradoriida.

LEPERDITICOPID ARTHROPODS (ORDOVICIAN-LATE DEVONIAN): FUNCTIONAL MORPHOLOGY AND ECOLOGICAL RANGE

Abstract The functional morphology and autecology of leperditicopid arthropods (Ordovician-uppermost Devonian) are analyzed in the light of well-preserved specimens from the Devonian of China and detailed comparisons with recent ostracodes. Leperditicopids were large, bivalved arthropods (adults ranging from 5 to about 50 mm in length) typically with an asymmetric carapace (strong ventral overlap), a complex muscular system (powerful adductors, extrinsic muscles, tendinous structures) whose insertions are preserved as scars on the inner surface of the exoskeleton and steinkerns, and an extensive radiating network of integumental sinuses probably involved in gaseous and ionic exchanges (oxygen uptake and transport, osmoregulation). The conspicuous chevron scars adjacent to the adductor scars are interpreted as the anchoring spots of mandibular tendinous structures possibly involved in the opening mechanism of the valves. The ultrastructure of the carapace is comparable to that of thick-shelled recent myodocopid ostracodes. A review of leperditicopid occurences (depositional environment, associated faunas and floras) shows that the group preferentially occupied very shallow marginal habitats (tidal flats, reef-flats, lagoons, embayments, or estuarian complexes) that were subjected to environmental stress (salinity, temperature, moisture). This ecological range implies specific adaptations (osmoregulation, resistance to desiccation) supported by morphological evidence (e.g., circulatory system, carapace closing system, thick shell). Most leperditicopids had epibenthic lifestyles and were probably detritus feeders. They may have been adapted (powerful mandibles) to scrape food on algal/microbial mats. Their typical pattern of occurrence (monospecificity, large numerical abundance) displays some of the characteristics of opportunistic populations (e.g., recent ostracodes, branchiopods) living in variable environments. Morphological similarities with Ostracoda are important (e.g., muscular and tendinous features, circulatory system, valve overlap, carapace ultrastructure) but taxonomic relationships with that group remain inconclusive because of the lack of evidence from soft parts.

Patterns of ostracod migration for the ‘North Atlantic’ region during the Ordovician

Abstract A review of Ordovician neritic ostracods from the ‘North Atlantic’ region including Europe and North America identifies over 100 genera (including 44 palaeocopes and 31 binodicopes) which show a complex pattern of migration between two or more of the palaeocontinents Gondwana, Ibero-Armorica, Perunica, Avalonia, Baltica and Laurentia. Many dispersals were relatively slow, and the migration of a genus between palaeocontinents often took the duration of one or more graptolite biozones. Over 70 migrations appear to have occurred more rapidly, including those of Pseudulrichia, a genus which dispersed to five palaeocontinents within the duration of three graptolite biozones. Longevity clearly facilitated the chances of migration, as the most widespread genera such as Vannieria, Platybolbina, Medianella and Euprimites, are often the most long-ranging. Low migration rates prior to the Llanvirn are, at least in part, related to low ostracod taxonomic diversity. Greatly increased diversity from the late Llanvirn coincided with a much higher rate of migration. Coupled with the spread of carbonate–mudstone shelf marine facies in Laurentia during the early and mid Caradoc, this resulted in the migration of up to 18 Baltic-origin genera to Laurentia. Relative to overall ostracod diversity, migration rates were generally higher during periods of lower global sea level, suggesting that ostracod dispersal may have been aided by mid-ocean islands or outer-shelf carbonate platforms, which provided more extensive island-hopping routes during periods of low sea level. The palaeogeographical convergence of Avalonia, Perunica and Baltica, and subsequently of Avalonia and Baltica with Laurentia, in low latitudes and warm surface waters, is suggested by increasing ostracod migration between these palaeocontinents from the late Llanvirn onwards. This culminated, during the Ashgill, in numerous species–level links. Baltica may have been the source area for more than 40 migrant genera, reflecting its high-diversity faunas and its intermediate palaeogeographical position between Laurentia and Avalonia. Several ostracod genera used Baltica as a staging-post in migrations between Avalonia and Laurentia. Migrations continued during the late Ashgill Hirnantian Stage (24 migrations), especially between Laurentia, Baltica and Avalonia (up to 19 migrations of genera), suggesting close geographical proximity for these palaeocontinents. Some ostracods, particularly the binodicopes Pseudulrichia, Klimphores, Kinnekullea, Aechmina and Spinigerites, could occupy outer-shelf and cooler-water benthic palaeoenvironments. They were part of a widespread deep-shelf fauna from the mid Caradoc onwards, for which distances or climatic barriers were less of an obstacle for trans-oceanic migration. None of these ostracods were bathyal.