The end of the line: competitive exclusion and the extinction of historical entities.

Identifying competitive exclusion at the macroevolutionary scale has typically relied on demonstrating a reciprocal, contradictory response by two co-occurring, functionally similar clades. Finding definitive examples of such a response in fossil time series has proven challenging, however, as has controlling for the effects of a changing physical environment. We take a novel approach to this issue by quantifying variation in trait values that capture almost the entirety of function for steam locomotives (SL), a known example of competitive exclusion from material culture, with the goal of identifying patterns suitable for assessing clade replacement in the fossil record. Our analyses find evidence of an immediate, directional response to the first appearance of a direct competitor, with subsequent competitors further reducing the realized niche of SLs, until extinction was the inevitable outcome. These results demonstrate when interspecific competition should lead to extinction and suggest that clade replacement may only occur when niche overlap between an incumbent and its competitors is near absolute and where the incumbent is incapable of transitioning to a new adaptive zone. Our findings provide the basis for a new approach to analyse putative examples of competitive exclusion that is largely free of a priori assumptions.

Supernova triggers for end-Devonian extinctions.

The Late Devonian was a protracted period of low speciation resulting in biodiversity decline, culminating in extinction events near the Devonian-Carboniferous boundary. Recent evidence indicates that the final extinction event may have coincided with a dramatic drop in stratospheric ozone, possibly due to a global temperature rise. Here we study an alternative possible cause for the postulated ozone drop: a nearby supernova explosion that could inflict damage by accelerating cosmic rays that can deliver ionizing radiation for up to [Formula: see text] ky. We therefore propose that the end-Devonian extinctions were triggered by supernova explosions at [Formula: see text], somewhat beyond the "kill distance" that would have precipitated a full mass extinction. Such nearby supernovae are likely due to core collapses of massive stars; these are concentrated in the thin Galactic disk where the Sun resides. Detecting either of the long-lived radioisotopes [Formula: see text] or [Formula: see text] in one or more end-Devonian extinction strata would confirm a supernova origin, point to the core-collapse explosion of a massive star, and probe supernova nucleosynthesis. Other possible tests of the supernova hypothesis are discussed.

A new species of the deuterostome Herpetogaster from the early Cambrian Chengjiang biota of South China.

The Cambrian radiation represents a key time period in the history of life. Here, we add to the mounting evidence accumulating on the nature of deuterostomes from this time period through description of a new species of stalked deuterostome, Herpetogaster haiyanensis nov. sp., from the lower Cambrian (series 2, stage 3) Chengjiang biota of China. This represents the first occurrence of the genus in Gondwana, the first juvenile specimen, and the oldest specimens to date. Herpetogaster haiyanensis nov. sp. differs from H. collinsi Caron et al. (2010) in having a stolon that is separated into an outer and inner layer, the segmentation of the body and in the shape and number of branches of the tentacles. The new species reiterates earlier suggestions of deuterostome affinities of the genus-it appears closely related to Phlogites and then successively more distantly related to Cotyledon and Eldonia-and may have fed on hyolithids.

A problematic animal fossil from the early Cambrian Hetang Formation, South China.

The lower-middle Hetang Formation (Cambrian Stage 2-3) deposited in slope-basinal facies in South China is well-known for its preservation of the earliest articulated sponge fossils, providing an important taphonomic window into the Cambrian explosion. However, the Hetang Formation also hosts a number of problematic animal fossils that have not been systematically described. This omission results in an incomplete picture of the Hetang biota and limits its contribution to the understanding of the early evolution of animals. Here we describe a new animal taxon, Cambrowania ovata Tang and Xiao, new genus new species, from the middle Hetang Formation in the Lantian area of southern Anhui Province, South China. Specimens are preserved as carbonaceous compressions, although some are secondarily mineralized. A comprehensive analysis using reflected light microscopy, scanning electron microscopy, energy dispersive X-ray spectroscopy, and micro-CT reveals that the new species is characterized by a spheroidal to fusoidal truss-like structure consisting of rafter-like crossbars, some of which are secondarily baritized and may have been internally hollow. Some specimens have aperture-like structures that are broadly similar to oscula of sponges, whereas others show evidence of a medial split reminiscent of gaping carapaces. While the phylogenetic affinity of Cambrowania ovata Tang and Xiao, new genus new species remains problematic, we propose that it may represent carapaces of bivalved arthropods or more likely sponges in early life stages. Along with other problematic metazoan fossils such as hyolithids and sphenothallids, Cambrowania ovata Tang and Xiao, new genus new species adds to the diversity of the sponge-dominated Hetang biota in an early Cambrian deep-water slope-basinal environment.

Using GIS to examine biogeographic and macroevolutionary patterns in some late Paleozoic cephalopods from the North American Midcontinent Sea.

Geographic range is an important macroevolutionary parameter frequently considered in paleontological studies as species' distributions and range sizes are determined by a variety of biotic and abiotic factors well known to affect the differential birth and death of species. Thus, considering how distributions and range sizes fluctuate over time can provide important insight into evolutionary dynamics. This study uses Geographic Information Systems (GIS) and analyses of evolutionary rates to examine how in some species within the Cephalopoda, an important pelagic clade, geographic range size and rates of speciation and extinction changed throughout the Pennsylvanian and early Permian in the North American Midcontinent Sea. This period is particularly interesting for biogeographic and evolutionary studies because it is characterized by repetitive interglacial-glacial cycles, a global transition from an icehouse to a greenhouse climate during the Late Paleozoic Ice Age, and decelerated macroevolutionary dynamics, i.e. low speciation and extinction rates. The analyses presented herein indicate that cephalopod species diversity was not completely static and actually fluctuated throughout the Pennsylvanian and early Permian, matching findings from other studies. However, contrary to some other studies, the mean geographic ranges of cephalopod species did not change significantly through time, despite numerous climate oscillations; further, geographic range size did not correlate with rates of speciation and extinction. These results suggest that pelagic organisms may have responded differently to late Paleozoic climate changes than benthic organisms, although additional consideration of this issue is needed. Finally, these results indicate that, at least in the case of cephalopods, macroevolution during the late Paleozoic was more dynamic than previously characterized, and patterns may have varied across different clades during this interval.

Soft-bodied fossils from the upper Valongo Formation (Middle Ordovician: Dapingian-Darriwilian) of northern Portugal.

Soft-bodied preservation is common in the Cambrian but comparatively rare in the Ordovician. Here, a new deposit preserving soft-bodied fossils is reported from the Middle Ordovician (Dapingian-Darriwilian) upper Valongo Formation of northern Portugal. The deposit contains the first known occurrences of soft-bodied fossils from the Middle Ordovician (Dapingian-Darriwilian) of Portugal and is the first Ordovician example of soft-tissue preservation involving carbonaceous films from the Iberian Peninsula. It also represents the lone deposit of soft-bodied fossils from the Middle Ordovician of northern Gondwana. Thus temporally, it lies between the exceptional deposits of the Lower Ordovician of Fezouata (Morocco) and the Upper Ordovician of the Soom Shale (South Africa); it also serves as a biogeographic link between these and the various Ordovician soft-bodied deposits in Laurentia. The soft-bodied fossils come from the deep-water slates of the upper part of the Valongo Formation and include a discoidal fossil questionably referable to Patanacta, wiwaxiid sclerites, and a possible pseudoarctolepid arthropod.

Metabolic rates, climate and macroevolution: a case study using Neogene molluscs.

Basal metabolic rate (BMR) is posited to be a fundamental control on the structure and dynamics of ecological networks, influencing organism resource use and rates of senescence. Differences in the maintenance energy requirements of individual species therefore potentially predict extinction likelihood. If validated, this would comprise an important link between organismic ecology and macroevolutionary dynamics. To test this hypothesis, the BMRs of organisms within fossil species were determined using body size and temperature data, and considered in the light of species' survival and extinction through time. Our analysis focused on the high-resolution record of Pliocene to recent molluscs (bivalves and gastropods) from the Western Atlantic. Species-specific BMRs were calculated by measuring the size range of specimens from museum collections, determining ocean temperature using the HadCM3 global climate model, and deriving values based on relevant equations. Intriguingly, a statistically significant difference in metabolic rate exists between those bivalve and gastropod taxa that went extinct and those that survived throughout the course of the Neogene. This indicates that there is a scaling up from organismic properties to species survival for these communities. Metabolic rate could therefore represent an important metric for predicting future extinction patterns, with changes in global climate potentially affecting the lifespan of individuals, ultimately leading to the extinction of the species they are contained within. We also find that, at the assemblage level, there are no significant differences in metabolic rates for different time intervals throughout the entire study period. This may suggest that Neogene mollusc communities have remained energetically stable, despite many extinctions.

Getting somewhere with the Red Queen: chasing a biologically modern definition of the hypothesis.

The Red Queen hypothesis (RQH) is both familiar and murky, with a scope and range that has broadened beyond its original focus. Although originally developed in the palaeontological arena, it now encompasses many evolutionary theories that champion biotic interactions as significant mechanisms for evolutionary change. As such it de-emphasizes the important role of abiotic drivers in evolution, even though such a role is frequently posited to be pivotal. Concomitant with this shift in focus, several studies challenged the validity of the RQH and downplayed its propriety. Herein, we examine in detail the assumptions that underpin the RQH in the hopes of furthering conceptual understanding and promoting appropriate application of the hypothesis. We identify issues and inconsistencies with the assumptions of the RQH, and propose a redefinition where the Red Queen's reign is restricted to certain types of biotic interactions and evolutionary patterns occurring at the population level.

Disc-shaped fossils resembling porpitids or eldonids from the early Cambrian (Series 2: Stage 4) of western USA.

The morphology and affinities of newly discovered disc-shaped, soft-bodied fossils from the early Cambrian (Series 2: Stage 4, Dyeran) Carrara Formation are discussed. These specimens show some similarity to the Ordovician Discophyllum Hall, 1847; traditionally this taxon had been treated as a fossil porpitid. However, recently it has instead been referred to as another clade, the eldonids, which includes the enigmatic Eldonia Walcott, 1911 that was originally described from the Cambrian Burgess Shale. The status of various Proterozoic and Phanerozoic taxa previously referred to porpitids and eldonids is also briefly considered. To help ascertain that the specimens were not dubio- or pseudofossils, elemental mapping using energy dispersive X-ray spectroscopy (EDS) was conducted. This, in conjunction with the morphology of the specimens, indicated that the fossils were not hematite, iron sulfide, pyrolusite, or other abiologic mineral precipitates. Instead, their status as biologic structures and thus actual fossils is supported. Enrichment in the element carbon, and also possibly to some extent the elements magnesium and iron, seems to be playing some role in the preservation process.

New olenelloid trilobites from the Northwest Territories, Canada.

The Olenelloidea are a superfamily of early Cambrian trilobites, which have been the subject of several phylogenetic analyses and also used to address macroevolutionary questions regarding the nature and timing of the Cambrian radiation. The Sekwi Formation of the Mackenzie Mountains, Northwest Territories, Canada, has yielded numerous species from this clade, and here we present new information that expands on the diversity known from this biogeographically and biostratigraphically important region. In particular, we describe seven new species, (Olenellus baileyi, Mesonacis wileyi, Elliptocephala jaredi, Holmiella taurus, H. domackae, Mummaspis delgadoae, and Bristolia colberti). Also recovered are specimens of Elliptocephala logani, specimens that shared affinities with Olenellus clarki, O. getzi, O. fowleri, and Frizolenellus hanseni, and one partial specimen, which appears to be a new species of Bolbolenellus.

Unraveling the phylogenetic relationships of the Eccoptochilinae, an enigmatic array of ordovician cheirurid trilobites.

The Cheiruridae are a diverse group of trilobites and several subfamilies within the clade have been the focus of recent phylogenetic studies. This paper focuses on the relationships of one of those subfamilies, the Ordovician Eccoptochilinae. We analyze sixteen species from six genera within the traditionally defined group, using the pilekiid Anacheirurus frederici as an outgroup. To assess the monophyly of the Eccoptochilinae seven sphaerexochine species, Kawina arnoldi, Sphaerexochus arenosus, S. atacius, S. latifrons, S. mirus, S. parvus, and S. scabridus were included in the analysis as well. The results of this analysis show that the genus Eccoptochile represents a paraphyletic grade and species traditionally assigned to Parasphaerexochus and Skelipyx plot within Pseudosphaerexochus. Also, representative species of Sphaerexochinae plot within the traditionally defined Eccoptochilinae, suggesting Eccoptochilinae itself is paraphyletic. To resolve this, we propose all species of Pseudosphaerexochus be placed within Sphaerexochinae and Eccoptochilinae be restricted to a monotypic Eccoptochile clavigera.

Phylogenetic and biogeographic analysis of sphaerexochine trilobites.

BACKGROUND: Sphaerexochinae is a speciose and widely distributed group of cheirurid trilobites. Their temporal range extends from the earliest Ordovician through the Silurian, and they survived the end Ordovician mass extinction event (the second largest mass extinction in Earth history). Prior to this study, the individual evolutionary relationships within the group had yet to be determined utilizing rigorous phylogenetic methods. Understanding these evolutionary relationships is important for producing a stable classification of the group, and will be useful in elucidating the effects the end Ordovician mass extinction had on the evolutionary and biogeographic history of the group. METHODOLOGY/PRINCIPAL FINDINGS: Cladistic parsimony analysis of cheirurid trilobites assigned to the subfamily Sphaerexochinae was conducted to evaluate phylogenetic patterns and produce a hypothesis of relationship for the group. This study utilized the program TNT, and the analysis included thirty-one taxa and thirty-nine characters. The results of this analysis were then used in a Lieberman-modified Brooks Parsimony Analysis to analyze biogeographic patterns during the Ordovician-Silurian. CONCLUSIONS/SIGNIFICANCE: The genus Sphaerexochus was found to be monophyletic, consisting of two smaller clades (one composed entirely of Ordovician species and another composed of Silurian and Ordovician species). By contrast, the genus Kawina was found to be paraphyletic. It is a basal grade that also contains taxa formerly assigned to Cydonocephalus. Phylogenetic patterns suggest Sphaerexochinae is a relatively distinctive trilobite clade because it appears to have been largely unaffected by the end Ordovician mass extinction. Finally, the biogeographic analysis yields two major conclusions about Sphaerexochus biogeography: Bohemia and Avalonia were close enough during the Silurian to exchange taxa; and during the Ordovician there was dispersal between Eastern Laurentia and the Yangtze block (South China) and between Eastern Laurentia and Avalonia.

Sharks that pass in the night: using Geographical Information Systems to investigate competition in the Cretaceous Western Interior Seaway.

One way the effects of both ecology and environment on species can be observed in the fossil record is as changes in geographical distribution and range size. The prevalence of competitive interactions and species replacements in the fossil record has long been investigated and many evolutionary perspectives, including those of Darwin, have emphasized the importance of competitive interactions that ultimately lead one species to replace another. However, evidence for such phenomena in the fossil record is not always manifest. Here we use new quantitative analytical techniques based on Geographical Information Systems and PaleoGIS tectonic reconstructions to consider this issue in greater detail. The abundant, well-preserved fossil marine vertebrates of the Late Cretaceous Western Interior Seaway of North America provide the component data for this study. Statistical analysis of distributional and range size changes in taxa confirms earlier ideas that the relative frequency of competitive replacement in the fossil record is limited to non-existent. It appears that typically, environmental gradients played the primary role in determining species distributions, with competitive interactions playing a more minor role.

Tracing the trilobite tree from the root to the tips: a model marriage of fossils and phylogeny.

Trilobites are a highly diverse group of extinct arthropods that persisted for nearly 300 million years. During that time, there was a profusion of morphological form, and they occupied a plethora of marine habitats. Their diversity, relative abundance, and complex morphology make them excellent candidates for phylogenetic analysis, and partly as a consequence they have been the subject of many cladistic studies. Although phylogenetic knowledge is certainly incomplete, our understanding of evolutionary patterns within the group has dramatically increased over the last 30 years. Moreover, trilobites have formed an important component of various studies of macroevolutionary processes. Here, we summarize the phylogenetic breadth of knowledge on the Trilobita, and present various hypotheses about phylogenetic patterns within the group, from the highest to the lowest taxonomic levels. Key topics we consider include the question of trilobite monophyly, the phylogenetic position of trilobites vis à vis extant arthropod groups, and inter- and intra-ordinal relationships.

Exceptionally preserved jellyfishes from the Middle Cambrian.

Cnidarians represent an early diverging animal group and thus insight into their origin and diversification is key to understanding metazoan evolution. Further, cnidarian jellyfish comprise an important component of modern marine planktonic ecosystems. Here we report on exceptionally preserved cnidarian jellyfish fossils from the Middle Cambrian (approximately 505 million years old) Marjum Formation of Utah. These are the first described Cambrian jellyfish fossils to display exquisite preservation of soft part anatomy including detailed features of structures interpreted as trailing tentacles and subumbrellar and exumbrellar surfaces. If the interpretation of these preserved characters is correct, their presence is diagnostic of modern jellyfish taxa. These new discoveries may provide insight into the scope of cnidarian diversity shortly after the Cambrian radiation, and would reinforce the notion that important taxonomic components of the modern planktonic realm were in place by the Cambrian period.

Considering the case for biodiversity cycles: re-examining the evidence for periodicity in the fossil record.

We re-examine the evidence for a 62 million year (Myr) periodicity in biodiversity throughout the Phanerozoic history of animal life reported by, as well as related questions of periodicity in origination and extinction. We find that the signal is robust against variations in methods of analysis, and is based on fluctuations in the Paleozoic and a substantial part of the Mesozoic. Examination of origination and extinction is somewhat ambiguous, with results depending upon procedure. Origination and extinction intensity as defined by may be affected by an artifact at 27 Myr in the duration of stratigraphic intervals. Nevertheless, when a procedure free of this artifact is implemented, the 27 Myr periodicity appears in origination, suggesting that the artifact may ultimately be based on a signal in the data. A 62 Myr feature appears in extinction, when this same procedure is used. We conclude that evidence for a periodicity at 62 Myr is robust, and evidence for periodicity at approximately 27 Myr is also present, albeit more ambiguous.

Random walks in the history of life.

The simplest null hypothesis for evolutionary time series is that the observed data follow a random walk. We examined whether aspects of Sepkoski's compilation of marine generic diversity depart from a random walk by using statistical tests from econometrics. Throughout most of the Phanerozoic, the random-walk null hypothesis is not rejected for marine diversity, accumulated origination or accumulated extinction, suggesting that either these variables were correlated with environmental variables that follow a random walk or so many mechanisms were affecting these variables, in different ways, that the resultant trends appear random. The only deviation from this pattern involves rejection of the null hypothesis for roughly the last 75 million years for the diversity and accumulated origination time series.

Taking the pulse of the cambrian radiation.

The Cambrian radiation is that key episode in the history of life when a large number of animal phyla appeared in the fossil record over a geologically short period of time. Over the last 20 years, scientific understanding of this radiation has increased significantly. Still, fundamental questions remain about the timing of the radiation and also the tempo of evolution. Trilobites are an excellent group to address these questions because of their rich abundance and diversity. Moreover, their complex morphology makes them readily amenable to phylogenetic analysis, and deducing the nature of macroevolutionary processes during the Cambrian radiation requires an understanding of evolutionary patterns. Phylogenetic biogeographic analysis of Early Cambrian olenellid trilobites, based on a modified version of Brooks Parsimony Analysis, revealed the signature of the breakup of Pannotia, a tectonic event that most evidence suggests is constrained to the interval 600 to 550 Ma. As trilobites are derived metazoans, this suggests the phylogenetic proliferation associated with the Cambrian radiation was underway tens of millions of years before the Early Cambrian, although not hundreds of millions of years as some have argued.Phylogenetic information from Early Cambrian olenellid trilobites was also used in a stochastic approach based on two continuous time models to test the hypothesis that rates of speciation were unusually high during the Cambrian radiation. No statistical evidence was found to support this hypothesis. Instead, rates of evolution during the Cambrian radiation, at least those pertaining to speciation, were comparable to those that have occurred during other times of adaptive or taxic radiation throughout the history of life.

Documenting a significant relationship between macroevolutionary origination rates and Phanerozoic pCO2 levels.

We show that the rates of diversification of the marine fauna and the levels of atmospheric CO(2) have been closely correlated for the past 545 million years. These results, using two of the fundamental databases of the Earth's biota and the Earth's atmospheric composition, respectively, are highly statistically significant (P < 0.001). The strength of the correlation suggests that one or more environmental variables controlling CO(2) levels have had a profound impact on evolution throughout the history of metazoan life. Comparing our work with highly significant correlations described by D. H. Rothman [Rothman, D. H. (2001) Proc. Natl. Acad. Sci. USA 98, 4305-4310] between total biological diversity and a measure of stable carbon isotope fractionation, we find that the rates of diversification rather than total diversification correlate with environmental variables, and that the rate of diversification follows the record of CO(2) projected by R. A. Berner and Z. Kothavala [Berner, R. A. & Kothavala, Z. (2001) Am. J. Sci. 301, 182-204] more closely than that predicted by Rothman.