Franz T. Fürsich

Phanerozoic trends in the global diversity of marine invertebrates.

It has previously been thought that there was a steep Cretaceous and Cenozoic radiation of marine invertebrates. This pattern can be replicated with a new data set of fossil occurrences representing 3.5 million specimens, but only when older analytical protocols are used. Moreover, analyses that employ sampling standardization and more robust counting methods show a modest rise in diversity with no clear trend after the mid-Cretaceous. Globally, locally, and at both high and low latitudes, diversity was less than twice as high in the Neogene as in the mid-Paleozoic. The ratio of global to local richness has changed little, and a latitudinal diversity gradient was present in the early Paleozoic.

Effects of sampling standardization on estimates of Phanerozoic marine diversification.

Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.

A fossil record full of holes: The Phanerozoic history of drilling predation

The evolutionary history of drilling predation, despite a long and rich fossil record (Precambrian–Holocene), contains a 120 m.y. gap (Late Triassic–Early Cretaceous). Drilled bivalve and brachiopod shells from Jurassic deposits of Hungary, India, and four localities documented in the literature indicate that drillers may have existed continuously throughout the Mesozoic. They may have been descendants of Paleozoic predators, unknown Mesozoic carnivores, or precursors of modern drillers. A literature database suggests three major phases in the Phanerozoic history of drilling predators: (1) the Paleozoic phase (latest Precambrian–Carboniferous) dominated by rare to moderately frequent drillings in brachiopods and sessile echinoderms; (2) the Mesozoic phase (Permian–Early Cretaceous) with very rare, or even facultative, drillers that had little impact on marine benthic communities, but nevertheless may have been present continuously; and (3) the Cenozoic phase (Late Cretaceous –Holocene) dominated by frequent gastropod drillings in mollusks.

Large colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr ago

The evidence for macroscopic life during the Palaeoproterozoic era (2.5–1.6 Gyr ago) is controversial. Except for the nearly 2-Gyr–old coil-shaped fossil Grypania spiralis, which may have been eukaryotic, evidence for morphological and taxonomic biodiversification of macroorganisms only occurs towards the beginning of the Mesoproterozoic era (1.6–1.0 Gyr). Here we report the discovery of centimetre-sized structures from the 2.1-Gyr-old black shales of the Palaeoproterozoic Francevillian B Formation in Gabon, which we interpret as highly organized and spatially discrete populations of colonial organisms. The structures are up to 12 cm in size and have characteristic shapes, with a simple but distinct ground pattern of flexible sheets and, usually, a permeating radial fabric. Geochemical analyses suggest that the sediments were deposited under an oxygenated water column. Carbon and sulphur isotopic data indicate that the structures were distinct biogenic objects, fossilized by pyritization early in the formation of the rock. The growth patterns deduced from the fossil morphologies suggest that the organisms showed cell-to-cell signalling and coordinated responses, as is commonly associated with multicellular organization. The Gabon fossils, occurring after the 2.45–2.32-Gyr increase in atmospheric oxygen concentration, may be seen as ancient representatives of multicellular life, which expanded so rapidly 1.5 Gyr later, in the Cambrian explosion.

Biostratinomy and palaeoecology of the cassian formation (Triassic) of the Southern Alps

Abstract The sediments forming the Cassian Formation (Middle—Upper Triassic) of the Central Dolomites represent the following environments: back-reef areas, shallow marginal basins, slope, and central basin. In each of these major environments faunal assemblages were found whose composition, mode of preservation and occurrence proved to be characteristics. In the back-reef areas, algal/foraminifera- and calcareous sponge/coral-patch reefs with a highly diverse fauna of frame-builders and reef-dwellers, in some localities excellently preserved, form the most conspicuous feature. In the shallow marginal basins autochthonous algal meadow/soft-bottom associations dominated by gastropods and deposit-feeding bivalves prevail. Slope deposits do not contain any autochthonous associations but either remains of pelagic faunas or transported faunal elements of shallow-water origin in the form of Cipit boulders (subaerially cemented parts from the outer edge of the carbonate platforms), worn Pachycardia rugosa-dominated, or echinoderm debris-dominated assemblages. Sediments of the central basin lack benthic faunas but contains pelagic assemblages of low diversity dominated by Daonella, Posidonia, and some ammonites.

Hardgrounds, reworked concretion levels and condensed horizons in the Jurassic of western India: their significance for basin analysis

Jurassic sediments in the shallow pericratonic basins of Kachchh and Rajasthan, western India, exhibit numerous signs of reduced sedimentation, omission, erosion and in situ reworking, in combination with synsedimentary cementation. Hardgrounds developed on carbonate shoals in the Bathonian of Rajasthan, whilst reworked concretion levels are characteristic of offshore siliciclastic sediments of the Callovian of Kachchh. A prominent marker horizon, the Oxfordian Dhosa Oolite Member, occurs throughout much of the Kachchh sub-basin and is a highly condensed unit characterized by hardgrounds, intraformational cobbles, reworked concretions, stromatolitic iron crusts, iron oncoids, and shell lags. Hardgrounds, reworked concretion levels, and condensation horizons are interpreted as the preserved relicts of transgressive pulses. Such pulses were possibly controlled by eustatic rise of sea level. Of at least equal importance, however, was a tectonic control which is demonstrated by the presence of small neptunian dykes, boulder beds derived from small submarine cliffs and rapid lateral facies and thickness changes in the Dhosa Oolite Member. These indications of extensional tectonics are thought to be connected to rifting and initial sea floor spreading between Africa and India.

Testing the role of biological interactions in the evolution of mid-Mesozoic marine benthic ecosystems

Abstract Evaluating the relative importance of biotic versus abiotic factors in governing macroevolutionary patterns is a central question of paleobiology. Here, we analyzed patterns of changes in global relative abundances and diversity of ecological groups to infer the role of biological interactions as driving evolutionary forces in mid-Mesozoic macrobenthic marine ecosystems. Specifically, we tested the hypothesis of escalation, which states that macroevolutionary patterns were controlled by an increasing pressure exerted by enemies on their victims. Associated with evidence of increasing levels of predation and biogenic sediment reworking (bulldozing) is an increasing representation of predation- and disturbance-resistant groups in the fossil record. In particular, we observe increasing proportions of mobile organisms; a decline of vulnerable epifauna living freely on the substrate; and a trend toward infaunalization of the benthos. These trends were most pronounced in the paleotropics, i.e., the region where biological activity is thought to have been highest. The observation that these changes affected several biotic traits and occurred within independent clades argues against the overriding role of a single key adaptive innovation in causing shifts in ecological abundance. Also, changes in the abiotic environment cannot explain these faunal patterns because of lacking cross-correlations with physico-chemical parameters such as global sea level, climate, and seawater chemistry. We conclude that in marine benthic ecosystems of the mid Mesozoic, enemy-driven evolution, or escalation, was a plausible and important factor.

Ecological, taxonomic, and taphonomic components of the post-Paleozoic increase in sample-level species diversity of marine benthos

Abstract Biological veracity of the sharp diversity increase observed in many analyses of the post-Paleozoic marine fossil record has been debated vigorously in recent years. To assess this question for sample-level (“alpha”) diversity, we used bulk samples of shelly invertebrates, representing three major fossil groups (brachiopods, bivalves, and gastropods), to compare the Jurassic and late Cenozoic sample-level diversity of marine benthos. After restricting the data set to single-bed, whole-fauna, bulk samples (n ≥ 30 specimens) from comparable open marine siliciclastic facies, we were able to retain 427 samples (255 Jurassic and 172 late Cenozoic), with most of those samples originating from our own empirical work. Regardless of the diversity metric applied, the initial results suggest that standardized sample-level species (or genus) diversity, driven by evenness and/or richness of the most common taxa, increased between the Jurassic and late Cenozoic by at least a factor of 1.6. When the data are partitioned into the three dominant higher taxa, it becomes clear that (1) the bivalves, which dominated the samples for both time intervals, increased in sample-level diversity between the Jurassic and the late Cenozoic by a much smaller factor than the total fauna; (2) the removal of brachiopods, which were a noticeable component of the Jurassic samples, did not significantly affect standardized sample-level diversity estimates; and (3) the gastropods, which were rare in the Jurassic but common in many late Cenozoic samples, contributed notably to the increase in sample-level diversity observed between the two time intervals. Parallel to these changes, the samples revealed secular trends in ecological structure, including Jurassic to late Cenozoic increases in proportion of (1) infauna, (2) mobile forms, and (3) non-suspension-feeding organisms. These trends mostly persist when data are restricted to bivalves. Supplementary analyses indicate that these patterns cannot be attributed to sampling heterogeneities in paleolatitudinal range, lithology, or paleoenvironment of deposition. Likewise, when data are restricted to samples dominated by species with originally aragonitic shells, the observed temporal changes persist at a comparable magnitude, suggesting that the pervasive loss of aragonite in the older fossil record is unlikely to have been the primary cause of the observed patterns. The comparable ratio of identified to unidentified species and genera, observed when comparing the Jurassic and late Cenozoic samples, indicates that the relatively poorer (mold/cast) preservation of Jurassic aragonite species also is unlikely to have been responsible for the observed patterns. However, the diagenesis-related taphonomic and methodological artifacts cannot be ruled out as an at least partial contributor to the observed post-Paleozoic changes in diversity, taxonomic composition, and ecology (the outcomes of the three tests of the diagenetic bias available to us are incongruent). The study demonstrates that the post-Paleozoic trends in the sample-level diversity, ecology, and taxonomic structure of common taxa can be replicated across multiple studies. However, the diversity increase estimated here is much less prominent than suggested by many previous analyses. The results also narrow the list of causative explanations down to two testable hypotheses. The first is diagenetic bias—a spurious trend driven by either (a) increasing taphonomic loss of small specimens in the older fossil record or (b) a shift in sampling procedures between predominantly lithified rocks of the Mesozoic and predominately unlithified, and therefore sievable, sediments of the late Cenozoic. The second hypothesis is genuine biological changes—macroevolutionary trends in the structure of marine benthic associations through time, consistent with predictions of several related models such as evolutionary escalation, increased ecospace utilization, and the Mesozoic marine revolution. Future studies should focus on testing these two rival models, a key remaining challenge for identifying the primary causative mechanism for the long-term changes in sample-level diversity, ecology, and taxonomic structure observed in the Phanerozoic marine fossil record.

Sequence stratigraphic significance of sedimentary cycles and shell concentrations in the Upper Jurassic–Lower Cretaceous of Kachchh, western India

Abstract Upper Jurassic and Lower Cretaceous siliciclastic shallow water sediments of the Kachchh Basin, western India, form strongly asymmetric coarsening-upward cycles, which are interpreted as recording changes in relative sea level (deepening–shallowing cycles). These cycles correspond to depositional sequences, in which deposits of the lowstand systems tract are not present, the sequence boundary coinciding with the transgressive surface. Shell concentrations are found in the transgressive lags at the base of the transgressive systems tract (TST), in the maximum flooding zone (MFZ), and at or close to the top of the highstand systems tract. They belong to six assemblages, five of them dominated by large bivalves such as Seebachia, Herzogina, Gryphaea, Gervillella, Megacucullaea, Pisotrigonia and Indotrigonia, the sixth by the coral Amphiastraea. Three types of shell concentrations can be distinguished that differ from each other in a number of ecological and taphonomic features, such as species diversity, preservation quality, orientation in cross-section, percentage of disarticulation, and degree of biogenic alteration. Characteristic features of concentrations at the base of the TSTs are moderate time-averaging, sorting, a preferred convex-up orientation, and nearly total disarticulation of shells. They are suggestive of an environment in which reworking and local transport were frequent events. Similar features are shown by concentrations near the tops of the HSTs, except that there shells were largely concentrated in lenses and in pavements rather than in beds as in the transgressive lags. Associated sedimentary structures indicate deposition above fair weather wave base in a high-energy environment. Concentrations occurring in the MFZ, in contrast, are autochthonous and highly time-averaged, having accumulated during times of low rates of sedimentation below storm wave base. This is supported by their high preservation quality (comparatively high percentage of articulated shells, shells of infaunal organisms commonly preserved in life position), biogenic alteration being the most important taphonomic agent. The dominant elements of these shell concentrations, i.e. Seebachia, Megacuccullaea, and Indotrigonia in the Upper Jurassic, and Pisotrigonia in the Lower Cretaceous, are endemic to the Ethiopean faunal province and belong to lineages that rapidly evolved during this time period.

Faunal response to transgressive−regressive cycles : example from the Jurassic of western India

Abstract Upper Callovian to Oxfordian shelf sediments at Ler in the Kachchh Basin, western India, are interpreted in terms of transgressive-regressive cycles. The transgressive phases are represented by thin layers of reworked and bored concretions, sometimes in association with skeletal concentrations, the regressive phases are documented by much thicker units of largely fine-grained sediments. The benthic fauna of transgressive and regressive phases differes markedly and thus mirrors the sedimentary cycles: During transgressive phases non-sedimentation produced hard substrates colonized mainly by byssate, cemented, or pedicle-attached suspension-feeding epifaunal species, whilst sediment input during regressive phases lead to soft substrate conditions characterized by infaunal deposit- and suspension-feeders as well as by epifaunal opportunists. Whereas the composition of the “regressive fauna” remains fairly constant through time, that of successive “transgressive faunas” often differs drastically. This is possibly due to a greater environmental sensitivity of epifaunal taxa as compared to infaunal taxa.