Fossils from the deep-sea Ediacaran biotas of Newfoundland are among the oldest architecturally complex soft-bodied macroorganisms on Earth. Most organisms in the Mistaken Point–type biotas of Avalonia—particularly the fractal-branching frondose Rangeomorpha— have been traditionally interpreted as living erect within the water column during life. However, due to the scarcity of documented physical sedimentological proxies associated with fossiliferous beds, Ediacaran paleocurrents have been inferred in some instances from the preferential orientation of fronds. This calls into question the relationship between frond orientation and paleocurrents. In this study, we present an integrated approach from a newly described fossiliferous surface (the “Melrose Surface” in the Fermeuse Formation at Melrose, on the southern portion of the Catalina Dome in the Discovery UNESCO Global Geopark) combining: (1) physical sedimentological evidence for paleocurrent direction in the form of climbing ripple cross-lamination and (2) a series of statistical analyses based on modified polythetic and monothetic clustering techniques reflecting the circular nature of the recorded orientation of Fractofusus misrai specimens. This study demonstrates the reclining rheotropic mode of life of the Ediacaran rangeomorph taxon Fractofusus misrai and presents preliminary inferences suggesting a similar mode of life for Bradgatia sp. and Pectinifrons abyssalis based on qualitative evidence. These results advocate for the consideration of an alternative conceptual hypothesis for position of life of Ediacaran organisms in which they are interpreted as having lived reclined on the seafloor, in the position that they are preserved.

The process of evolution and the structures it produces are best understood in the light of hierarchy theory. The biota traditionally is described by either the genealogical Linnaean hierarchy or economic hierarchies of communities or ecosystems. Here we describe the Bretskyan hierarchy—a hybrid eco-genealogical hierarchy that consists of nested sets of different-sized, usually polyphyletic communities of interacting individuals separated from other such communities in space and time at multiple scales. The Bretskyan hierarchy consists of elements that have both genealogical and economic properties and functions—situated between, and connecting the elements of, the economic hierarchies (Vernadskyan) and the genealogical (Linnaean) hierarchy. The described hierarchy at lower tiers is populated by holobionts, individuals composed of multiple polyphyletic lineages integrated by functional interactions or biotically fabricated structures, such as membranes. At larger spatial tiers and longer time scales, the members of the Bretskyan hierarchy are of a more diffuse nature, partially due to the small size and relatively short duration of us as observers of larger and longer-lasting structures, here described as geobiomes. Their individuality is externally forced and directly tied to the spatial and temporal physical structures of our planet. These are sub-bioprovinces and bioprovinces—large and effectively isolated spatiotemporal structures of biota integrated internally by coevolution and individuated externally by a hierarchy of barriers. Gaia is here understood as the largest eco-genealogical individual compartmentalized by the outer space of the Earth and integrated at long time scales by biotic interactions and plate tectonic mixing of biota. The existence of a hierarchy of barriers and multilevel allopatry suggests that geographic isolation takes part not only in individuating species lineages, but also in producing coherent complexes of separate lineages forming bioprovinces at multiple space and time scales. The sizes, configurations, and durations of Bretskyan units are directly tied to geodynamics, demonstrating the central role of the physical planet in the processes of individuation and merging of geobiomes and the control of coevolution, and all its ramifications, at multiple space and time scales. The Bretskyan hierarchy also allows the integration of previously unconnected themes—“egalitarian” major transitions in individuality (e.g., eukaryogenesis) and some of the megatrajectories in the history of life—into a single theoretical framework of spatial and temporal scaling of eco-genealogy. The pervasive scaling of geodynamical processes and the direct connection of geodynamics to the dynamics of Bretskyan units allows us to formulate conjectures on the scales and limits of spatial and temporal contingency and competitiveness of biotas in evolution.

A thorough understanding of how communities respond to extreme changes, such as biotic invasions, is essential to manage ecosystems today. Here we constructed fossil food webs to identify changes in Late Ordovician (Katian) shallow-marine paleocommunity structure and functioning before and after the Richmondian invasion, a well-documented ancient invasion. Food webs were compared using descriptive metrics and cascading extinction on graphs models. Richness at intermediate trophic levels was underrepresented when using only data from the Paleobiology Database relative to museum collections, resulting in a spurious decrease in modeled paleocommunity stability. Therefore, museum collections and field sampling may provide more reliable sources of data for the reconstruction of trophic organization in comparison to online data repositories. The invasion resulted in several changes in ecosystem dynamics. Despite topological similarities between pre- and postinvasion food webs, species loss occurred corresponding to a minor decrease in functional groups. Invaders occupied all of the preinvasion functional guilds, with the exception of four incumbent guilds that were lost and one new guild, corroborating the notion that invaders replace incumbents and fill preexisting niche space. Overall, models exhibited strong resistance to secondary extinction, although the postinvasion community had a lower threshold of collapse and more variable response to perturbation. We interpret these changes in dynamics as a decrease in stability, despite similarities in overall structure. Changes in food web structure and functioning resulting from the invasion suggest that conservation efforts may need to focus on preserving functional diversity if more diverse ecosystems are not inherently more stable.

The logarithmic helicospiral has been the most widely accepted model of regularly coiled molluscan form since it was proposed by Moseley and popularized by Thompson and Raup. It is based on an explicit assumption that shells are isometric and grow exponentially, and an implicit assumption that the external form of the shell follows the internal shape, which implies that the parameters of the spiral could be reconstructed from the external whorl profile. In this contribution, we show that these assumptions fail on all 25 gastropod species we examine. Using a dataset of 176 fossil and modern gastropod shells, we construct an empirical morphospace of coiling using the parameters of whorl expansion rate, translation rate, and rate of increasing distance from coiling axis, plus rate of aperture shape change, from their best-fit models. We present a case study of change in shell form through geologic time in the austral family Struthiolariidae to demonstrate the utility of our approach for evolutionary paleobiology. We fit various functions to the shell-coiling parameters to demonstrate that the best morphological model is not the same for each parameter. We present a set of R routines that will calculate helicospiral parameters from sagittal sections through coiled shells and allow workers to compare models and choose appropriate sets of parameters for their own datasets. Shell-form parameters in the Struthiolariidae highlight a hitherto neglected hypothesis of relationship between Antarctic Perissodonta and the enigmatic Australian genus Tylospira that fits the biogeographic and stratigraphic distribution of both genera.

Researchers often interpret the presence of tortoises in Pleistocene assemblages as evidence of an interglacial age, based on an assumption that these fossils indicate thermic climates, as modern giant tortoises require. Since the Paleocene, tortoises have been common components of terrestrial fossil assemblages and have repeatedly evolved species of giant size. Whereas extant giant tortoises are found only on islands off the coasts of South America and Africa, at least two species persisted in North America until the terminal Pleistocene. These tortoises, Hesperotestudo crassiscutata and Gopherus “hexagonatus,” both of which reached carapace lengths of >1 m, were distributed across the southern United States. This study provides new metrics to derive quantitative weight estimates from measurements of the tortoise shell. The linear measurements of 69 anatomical features of the shells of 108 live tortoises indicate that the regression between straight carapace length and weight is most significant, with a maximum r2 > 0.99. This regression is useful for tortoises that weigh between 1.8 and 339 kg. This mass estimate, coupled with a heat dissipation rate derived from thermoregulation modeling, provides estimates of how long tortoises can maintain a viable body temperature at low ambient temperatures. Depending on size, a tortoise can survive a maximum of 2.3 to 33 hours of freezing temperatures, which corresponds to a mean annual temperature ≥22°C and a mean winter low temperature ≥7.5°C. This analysis infers warmer temperatures at Pleistocene sites with fossil tortoise occurrences than previous qualitative estimates.

Brachiopods dominated the seafloor as a primary member of the Paleozoic fauna. Despite the devastating effects of the end-Permian extinction, the group recovered during the early Mesozoic only to gradually decline from the Jurassic to today. This decline likely had multiple causes, including increased predation and bioturbation-driven substrate disruption, but the role of changing substrate is not well understood. Given the importance of substrate for extant brachiopod habitat, we documented Mesozoic–Cenozoic lithologic preferences and morphological changes to assess how decreasing firm-substrate habitat may have contributed to the brachiopod decline. Compared with bivalves, Mesozoic brachiopods occurred more frequently and were disproportionately abundant in carbonate lithologies. Although patterns in glauconitic or ferruginous sediments are equivocal, brachiopods became more abundant in coarser-grained carbonates and less abundant in fine-grained siliciclastics. During the Jurassic, brachiopod species rarely had abraded beaks but tended to be more convex with a high beak, potentially consistent with a non-analogue lifestyle resting on the seafloor. However, those highly convex morphotypes largely disappeared by the Cenozoic, when more terebratulides had abraded beaks, suggesting closer attachment to hard substrates. Rhynchonellides disproportionately declined to become a minor component of Cenozoic faunas, perhaps because of less pronounced morphological shifts. Trends in lithologic preferences and morphology are consistent with bioturbation-driven substrate disruption, with brachiopods initially using firmer carbonate sediments as refugia before adapting to live primarily attached to hard surfaces. This progressive habitat restriction likely played a role in the final brachiopod decline, as bioturbating ecosystem engineers transformed benthic habitats in the Mesozoic and Cenozoic.

Because teeth are the most easily preserved part of the vertebrate skeleton and are particularly morphologically variable in mammals, studies of fossil mammals rely heavily on dental morphology. Dental morphology is used both for systematics and phylogeny as well as for inferences about paleoecology, diet in particular. We analyze the influence of evolutionary history on our ability to reconstruct diet from dental morphology in the mammalian order Carnivora, and we find that much of our understanding of diet in carnivorans is dependent on the phylogenetic constraints on diet in this clade. Substantial error in estimating diet from dental morphology is present regardless of the morphological data used to make the inference, although more extensive morphological datasets are more accurate in predicting diet than more limited character sets. Unfortunately, including phylogeny in making dietary inferences actually decreases the accuracy of these predictions, showing that dietary predictions from morphology are substantially dependent on the evolutionary constraints on carnivore diet and tooth shape. The “evolutionary ratchet” that drives lineages of carnivorans to evolve greater degrees of hypercarnivory through time actually plays a role in allowing dietary inference from tooth shape, but consequently requires caution in interpreting dietary inference from the teeth fossil carnivores. These difficulties are another reminder of the differences in evolutionary tempo and mode between morphology and ecology.

Numerical and taxonomic resolution of compositional data sets affects investigators’ abilities to detect and measure relationships between communities and environmental factors. We test whether varying numerical (untransformed, square-root- and fourth-root-transformed relative abundance and presence–absence data) and taxonomic (species, genera, families) resolutions reveals different insights into early to middle Miocene molluscan communities along bathymetric and salinity gradients. The marine subtidal has a more even species-abundance distribution, a higher number of rare species, and higher species:family and species:genus ratios than the three habitats—marine and estuarine intertidal, estuarine subtidal—with higher fluctuations in salinity and other physical parameters. Taxonomic aggregation and numerical transformation of data result in very different ordinations, although all habitats differ significantly from one another at all taxonomic and numerical levels. Rank correlations between species-level and higher-taxon, among-sample dissimilarities are very high for proportional abundance and decrease strongly with increasing numerical transformation, most notably in the two intertidal habitats. The proportion of variation explained by depth is highest for family-level data, decreases gradually with numerical transformation, and is higher in marine than in estuarine habitats. The proportion of variation explained by salinity is highest for species-level data, increases gradually with numerical transformation, and is higher in subtidal than in intertidal habitats. Therefore, there is no single best numerical and taxonomic resolution for the discrimination of communities along environmental gradients: the “best” resolution depends on the environmental factor considered and the nature of community response to it. Different numerical and taxonomic transformations capture unique aspects of metacommunity assembly along environmental gradients that are not detectable at a single level of resolution. We suggest that simultaneous analyses of community gradients at multiple taxonomic and numerical resolutions provide novel insights into processes responsible for spatial and temporal community stability.

The complex morphological evolution of the bivalve Ptychomya throughout the well-studied Agrio Formation in the Neuquén Basin (west-central Argentina, lower/upper Valanginian–lowest Barremian) constitutes an ideal opportunity to study evolutionary patterns and processes occurring at geological timescales. Ptychomya is represented in this unit by four species, the morphological variation of which needs to be temporally assessed to obtain a thorough picture of the evolution of the group. Here we use geometric morphometrics to measure variation in shell outline, ribbing pattern, and shell size in these species. We bracket the ages of our samples using a combination of ammonoid biostratigraphy and absolute ages and study the anagenetic pattern of evolution of each trait by means of paleontological time-series analysis and change tracking. We find that evolution in Ptychomya is mostly speciational, as the majority of traits show stasis, with the exceptions of shell size in P. coihuicoensis and shell outline in P. windhauseni, which seem to evolve directionally toward larger and higher shells, respectively. Ptychomya displays changes in its average morphology and disparity, which are the result of a mixture of taxonomic turnover and mosaic evolution of traits. Pulses of speciation would have been triggered by ecological opportunity, as they occur during the recovery of shallow-burrowing bivalve faunas after dysoxic events affecting the basin. On the other hand, the presence of directional patterns of evolution in P. coihuicoensis and P. windhauseni seems to be the result of a general shallowing-upward trend observed in the basin during the upper Hauterivian–lowest Barremian, as opposed to the cyclical paleoenvironmental stability inferred for the early/late Valanginian–early Hauterivian, which would have prompted stasis in P. koeneni and P. esbelta.

Despite the rich fossil record of Neogene chondrichthyans (chimaeras, sharks, rays, and skates) from Europe, little is known about the macroevolutionary processes that generated their current diversity and geographical distribution. We compiled 4368 Neogene occurrences comprising 102 genera, 41 families, and 12 orders from four European regions (Atlantic, Mediterranean, North Sea, and Paratethys) and evaluated their diversification trajectories and paleobiogeographic patterns. In all regions analyzed, we found that generic richness increased during the early Miocene, then decreased sharply during the middle Miocene in the Paratethys, and moderately during the late Miocene and Pliocene in the Mediterranean and North Seas. Origination rates display the most significant pulses in the early Miocene in all regions. Extinction rate pulses varied across regions, with the Paratethys displaying the most significant pulses during the late Miocene and the Mediterranean and North Seas during the late Miocene and early Pliocene. Overall, up to 27% and 56% of the European Neogene genera are now globally and regionally extinct, respectively. The observed pulses of origination and extinction in the different regions coincide with warming and cooling events that occurred during the Neogene globally and regionally. Our study reveals complex diversity dynamics of Neogene chondrichthyans from Europe and their distinct biogeographic composition despite the multiple marine passages that connected the different marine regions during this time.

In their “alternative interpretation” Marshall and Hecht raise two interesting questions:

1. I) Is “Taxocene Analysis” a permissible approach to community evolution, and if so under what conditions?

2. II) What really happened to the ten South American ungulate genera that vanished during the Early Pleistocene?

As these are fundamentally different questions, I shall consider each separately.

The multibrachiate grade of organization in camerate crinoids was principally attained through pinnule differentiation: the accelerated development of a proximal pinnule that ultimately became a pinnule-bearing arm. Although pinnule differentiation is not the only means of achieving a multibrachiate organization in modern crinoids, it was the primary mode in producing both dichotomous and exotomous branching patterns in multibrachiate camerates. Whereas most of the proximal branchings, commonly fixed in the cup of camerates, were due to pinnule differentiaton, the first dichotomy and distal dichotomies above the cup apparently developed from direct modification of a brachial plate at the growing tip of the immature crinoid arm.

According to when they attained high diversity, major taxa of marine animals have been clustered into three groups, the Cambrian, Paleozoic, and Modern Faunas. Because the Cambrian Fauna was a relatively minor component of the total fauna after mid-Ordovician time, the Phanerozoic history of marine animal diversity is largely a matter of the fates of the Paleozoic and Modern Faunas. The fact that most late Cenozoic genera belong to taxa that have been radiating for tens of millions of years indicates that the post-Paleozoic increase in diversity indicated by fossil data is real, rather than an artifact of improvement of the fossil record toward the present.

Assuming that ecological crowding produced the so-called Paleozoic plateau for family diversity, various workers have used the logistic equation of ecology to model marine animal diversification as damped exponential increase. Several lines of evidence indicate that this procedure is inappropriate. A plot of the diversity of marine animal genera through time provides better resolution than the plot for families and has a more jagged appearance. Generic diversity generally increased rapidly during the Paleozoic, except when set back by pulses of mass extinction. In fact, an analysis of the history of the Paleozoic Fauna during the Paleozoic Era reveals no general correlation between rate of increase for this fauna and total marine animal diversity. Furthermore, realistically scaled logistic simulations do not mimic the empirical pattern. In addition, it is difficult to imagine how some fixed limit for diversity could have persisted throughout the Paleozoic Era, when the ecological structure of the marine ecosystem was constantly changing. More fundamentally, the basic idea that competition can set a limit for marine animal diversity is incompatible with basic tenets of marine ecology: predation, disturbance, and vagaries of recruitment determine local population sizes for most marine species. Sparseness of predators probably played a larger role than weak competition in elevating rates of diversification during the initial (Ordovician) radiation of marine animals and during recoveries from mass extinctions. A plot of diversification against total diversity for these intervals yields a band of points above the one representing background intervals, and yet this band also displays no significant trend (if the two earliest intervals of the initial Ordovician are excluded as times of exceptional evolutionary innovation). Thus, a distinctive structure characterized the marine ecosystem during intervals of evolutionary radiation—one in which rates of diversification were exceptionally high and yet increases in diversity did not depress rates of diversification.

Particular marine taxa exhibit background rates of origination and extinction that rank similarly when compared with those of other taxa. Rates are correlated in this way because certain heritable traits influence probability of speciation and probability of extinction in similar ways. Background rates of origination and extinction were depressed during the late Paleozoic ice age for all major marine invertebrate taxa, but remained correlated. Also, taxa with relatively high background rates of extinction experienced exceptionally heavy losses during biotic crises because background rates of extinction were intensified in a multiplicative manner; decimation of a large group of taxa of this kind in the two Permian mass extinctions established their collective identity as the Paleozoic Fauna.

Characteristic rates of origination and extinction for major taxa persisted from Paleozoic into post-Paleozoic time. Because of the causal linkage between rates of origination and extinction, pulses of extinction tended to drag down overall rates of origination as well as overall rates of extinction by preferentially eliminating higher taxa having relatively high background rates of extinction. This extinction/origination ratchet depressed turnover rates for the residual Paleozoic Fauna during the Mesozoic Era. A decline of this fauna's extinction rate to approximately that of the Modern Fauna accounts for the nearly equal fractional losses experienced by the two faunas in the terminal Cretaceous mass extinction.

Viewed arithmetically, the fossil record indicates slow diversification for the Modern Fauna during Paleozoic time, followed by much more rapid expansion during Mesozoic and Cenozoic time. When viewed more appropriately as depicting geometric—or exponential—increase, however, the empirical pattern exhibits no fundamental secular change: the background rate of increase for the Modern Fauna—the fauna that dominated post-Paleozoic marine diversity—simply persisted, reflecting the intrinsic origination and extinction rates of constituent taxa. Persistence of this overall background rate supports other evidence that the empirical record of diversification for marine animal life since Paleozoic time represents actual exponential increase. This enduring rate makes it unnecessary to invoke environmental change to explain the post-Paleozoic increase of marine diversity.

Because of the resilience of intrinsic rates, an empirically based simulation that entails intervals of exponential increase for the Paleozoic and Modern Faunas, punctuated by mass extinctions, yields a pattern that is remarkably similar to the empirical pattern. It follows that marine animal genera and species will continue to diversify exponentially long into the future, barring disruption of the marine ecosystem by human-induced or natural environmental changes.

Understanding the mechanisms that prevent or promote the coexistence of taxa at local scales is critical to understanding how biodiversity is maintained. Competitive exclusion and environmental filtering are two processes thought to limit which taxa become established in a community. However, determining the relative importance of the two processes is a complex task, especially when the critical initial stages of colonization cannot be directly observed. Here, we explore the use of phylogenetic community structure for identifying filtering mechanisms in a fossil community. We integrated a time-calibrated molecular phylogeny of bivalve genera with a spatial dataset of late Cenozoic bivalves from the Pacific coast of North America to characterize how the community that was present in the semirestricted San Joaquin Basin (SJB) embayment of present-day California was phylogenetically structured. We employed phylogenetic distance-based metrics across six time bins spanning 27–2.5 Ma and found no evidence of significant clustering or evenness in the SJB community when compared with communities randomly assembled from the regional source pool. Additionally, we found that new colonizers into the SJB were not significantly more or less closely related to native taxa than expected by chance. These findings suggest that neither competitive exclusion nor environmental filtering were overwhelmingly influential factors shaping the composition of the SJB community over time. We further discuss interpretations of these patterns in light of current understandings in community phylogenetics and reiterate the critical role historical perspectives play in how community assembly rules are assessed.

Vertebrate microfossil assemblages in a stratigraphic sequence often yield similar assortments of taxa but at different relative abundances, potentially indicative of marginal paleocommunity changes driven by paleoenvironmental change over time. For example, stratigraphically younger assemblages in the Dinosaur Park Formation (DPF) yield proportionally more aquatic taxa, consistent with marine transgression. However, individual deposits may have received specimens from multiple source paleocommunities over time, limiting our ability to confidently identify ecologically significant, paleocommunity differences through direct assemblage comparisons. We adapted a three-source, two-tracer Bayesian mixing model to quantify proportional contributions from different source habitats to DPF microfossil assemblages. Prior information about the compositions of separate, relatively unmixed terrestrial, freshwater, and marine assemblages from the Belly River Group allowed us to define expected taxon percent abundances for the end-member habitats likely contributing specimens to the mixed deposits. We compared the mixed assemblage and end-member distributions using 21 different combinations of vertebrate taxa. Chondrichthyan, dinosaur, and amphibian occurrence patterns ultimately allowed us to parse the contributions from the potential sources to 14 of the 15 mixed assemblages. The results confirmed a significant decline in terrestrial contributions at younger DPF sites, driven primarily by increased freshwater specimen inputs—not incursions from the adjacent marine paleocommunity. A rising base level likely increased lateral channel migration and the prevalence of freshwater habitats on the landscape, factors that contributed to increased paleocommunity mixing at younger channel deposit sites. Bayesian methods can account for source-mixing bias, which may be common in assemblages associated with major paleoenvironmental changes.

The Cenozoic genus Terebratula seems to be an exception to the post-Permian trend in brachiopod retreat to offshore habitats, because it was species rich and numerically abundant in warm-temperate shallow-water environments in the Mediterranean and the Paratethys realms. This was so despite the general dominance of bivalves and the pervasive bioturbation and predation pressure during the Neogene. Terebratula, however, went extinct in the Calabrian (Pleistocene). The optimal environmental conditions for Terebratula during its prime are poorly known. The Águilas Basin (SE Spain) is an ideal study area to investigate the habitat of Terebratula, because shell beds of this brachiopod occur there cyclically in early Pliocene deposits. We evaluate the paleoecological boundary conditions controlling the distribution of Terebratula by estimating its environmental tolerances using benthic and planktic foraminiferal and nannoplanktic assemblages and oxygen isotopes of the secondary layer brachiopod calcite. Our results suggest that Terebratula in the Águilas Basin favored oligotrophic to mesotrophic, well-oxygenated environments at water depths of 60–90 m. Planktic foraminiferal assemblages and oxygen isotopes point to sea-surface temperatures between ~16°C and 22°C, and bottom-water temperatures between 17°C and 24°C. The analyzed proxies indicate that Terebratula tolerated local variations in water depth, bottom temperature, oxygenation, productivity, and organic enrichment. Terebratula was probably excluded by grazing pressure from well-lit environments and preferentially occupied sediment-starved, current-swept upper offshore habitats where coralline red algae were absent. Narrow temperature ranges of Terebratula species might have been a disadvantage during the high-amplitude seawater temperature fluctuations that started about 1 Ma, when the genus went extinct.

The number of individuals of species varies, but estimating abundance, given incomplete and biased sampling in both contemporary and fossilized communities, is challenging. Here, we describe a new occupancy model in a hierarchical Bayesian framework with random effects, in which multispecies occupancy and detection are modeled as a means to estimate relative species abundance and relative population densities. The modeling framework is suited for temporal samples of fossil communities with repeated sampling including multiple species with similar preservation potential. We demonstrate our modeling framework using a fossil community of benthic organisms to estimate relative species abundance dynamics and changing relative population densities of focal species in nine (geological) time intervals over 2.3 Myr. We also explore potential explanatory factors (paleoenvironmental proxies) and temporal autocorrelation that could provide extra information on unsampled time intervals. The modeling framework is applicable across a wide range of questions on species-level dynamics in paleoecological community settings.

The Burgess Shale-type Lagerstätten of the Early Cambrian Maotianshan Shale record an apparently sudden conquest of pelagic niches by ten phyla of metazoans. One of these phyla is Chaetognatha, a group of predatory marine worms. Given their role as major predators in modern planktonic ecosystems, the chaetognaths discovered in the Maotianshan Shale (Yunnan Province, South China) suggest that the pelagos at the time was already quite complex. Modern chaetognaths, however, can be divided into benthic and pelagic forms; the pelagic nature of Eognathacantha ercainella should therefore be validated by strong morphological evidence.

Knowing that planktonic animals present morphological adaptations that increase their buoyancy, we studied the drag produced during the active phase of chaetognath locomotion for the modern forms Paraspadella gotoi (benthic) and Sagitta elegans (pelagic). By using a motion model developed by Jordan in 1992, we could calculate the resistive force produced by the undulatory movement of chaetognaths' bodies.

This mechanistic approach evaluates the effect of three motion parameters (relative speed, total length, and tail ratio) on the drag force produced during locomotion. Our results show that the increase of size contributes to higher drag while the shorter tail of the pelagic form balances this effect by reducing the wetted surface subject to friction. For chaetognaths, therefore, a bigger body (both in length and width) and a shorter tail indicate a pelagic lifestyle, a finding that can be applied to the study of the fossil Eognathacantha ercainella. A discriminant analysis can confirm that the Early Cambrian chaetognath presents a pelagic morphology with similarities to modern bathypelagic and mesopelagic species.