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The 2.0–1.8-billion-year-old Stirling Range Formation in southwestern Australia preserves the deposits of a siliciclastic shoreline formed under the influence of storms, longshore currents, and tidal currents. Sandstones contain a megascopic fossil biota represented by discoidal fossils similar to the Ediacaran Aspidella Billings, 1872, as well as ridge pairs preserved in positive hyporelief on the soles of channel-fill sandstones bounded by mud drapes. The ridges run parallel or nearly parallel for most of their length, meeting in a closed loop at one end and opening with a slight divergence at the opposite end. The ridges are interpreted as casts of sediment-laden mucus strings formed by the movement of multicellular or syncytial organisms along a muddy surface. The taxa Myxomitodes stirlingensis n. igen., n. isp., are introduced for these traces. The Stirling biota was roughly coeval with other presumed multicellular eukaryotes appearing after a long period of profound environmental changes involving a rise in ambient oxygen levels, similar to that which preceded the Cambrian explosion. The failure of multicellular life to diversify during most of the Proterozoic may be due to environmental constraints related to the comparatively low level of oxidation of the world oceans.
By digitally imaging colonies with more than a hundred cells, the distributions of cell size and shape are determined for four examples of 2-Ga microfossils: bacillus-shaped Eosynechococcus moorei and three dyads or diplococci (Sphaerophycus parvum and two forms of Eoentophysalis belcherensis). By assuming that each colony obeys steady-state growth, the measured distributions can be inverted to infer the time evolution of the individual cell shape. The time evolution can also be predicted analytically from rate-based models of cell growth, permitting the data to distinguish among different postulates for the physical principles governing growth. The cell cycles are found to be best described by the exponential growth of cell volume, although linear volume growth is not ruled out. However, the measured dyad cycles are inconsistent with several growth models based on surface area or the behavior of the septum at the division plane. Where they have been measured, modern bacilli obey exponential growth whereas eukaryotics obey linear growth, which implies that these 2-Ga microfossils are likely prokaryotic.
The end-Permian mass extinction is commonly portrayed not only as a massive biodiversity crisis but also as the time when marine benthic faunas changed from the Paleozoic Fauna, dominated by rhynchonelliform brachiopod taxa, to the Modern Fauna, dominated by gastropod and bivalve taxa. After the end-Permian mass extinction, scenarios involving the Mesozoic Marine Revolution portray a steady increase in numerical dominance by these benthic molluscs as largely due to the evolutionary effects of an “arms race.” We report here a new global paleoecological database from study of shell beds that shows a dramatic geologically sudden earliest Triassic takeover by bivalves as numerical dominants in level-bottom benthic marine communities, which continued through the Early Triassic. Three bivalve genera were responsible for this switch, none of which has any particular morphological features to distinguish it from many typical Paleozoic bivalve genera. The numerical success of these Early Triassic bivalves cannot be attributed to any of the well-known morphological evolutionary innovations of post-Paleozoic bivalves that characterize the Mesozoic Marine Revolution. Rather, their ability to mount this takeover most likely was due to the large extinction of rhynchonelliform brachiopods during the end-Permian mass extinction and aided by their environmental distribution and physiological characteristics that enabled them to thrive during periods of oceanic and atmospheric stress during the Permian/Triassic transition.
Ecology is thought to be of crucial importance in determining taxonomic turnover at geological time scales, yet general links between ecology and biodiversity dynamics are still poorly explored in deep time. Here we analyze the relationships between the environmental affinities of Triassic–Jurassic marine benthic genera and their biodiversity dynamics, using a large, taxonomically vetted data set of Triassic–Jurassic taxonomic occurrences.
On the basis of binomial probabilities of proportional occurrence counts, we identify environmental affinities of genera for (1) carbonate versus siliciclastic substrates, (2) onshore versus offshore depositional environments, (3) reefs versus level-bottom communities, and (4) tropical versus non-tropical latitudinal zones. Genera with affinities for carbonates, onshore environments, and reefs have higher turnover rates than genera with affinities for siliciclastic, offshore, and level-bottom settings. Differences in faunal turnover are largely due to differences in origination rates. Whereas previous studies have highlighted the direct influence of physical and biological factors in exploring environmental controls on evolutionary rates, our analyses show that the patterns can largely be explained by the partitioning of higher taxa with different evolutionary tempos among environments. The relatively slowly evolving bivalves are concentrated in siliciclastic rocks and in level-bottom communities. Furthermore, separate analyses on bivalves did not produce significant differences in turnover rates between environmental settings. The relationship between biodiversity dynamics and environments in our data set is thus governed by the partitioning of higher taxa within environmental categories and not directly due to greater chances of origination in particular settings. As this partitioning probably has ecological reasons rather than being a simple sampling artifact, we propose an indirect environmental control on evolutionary rates.
Affinities for latitudinal zones are not linked to systematically different turnover rates, possibly because of paleoclimatic fluctuations and latitudinal migrations of taxa. However, the strong extinction spike of tropical genera in the Rhaetian calls for an important paleoclimatic component in the end-Triassic mass extinction.
Understanding what drives global diversity requires knowledge of the processes that control diversity and turnover at a variety of geographic and temporal scales. This is of particular importance in the study of mass extinctions, which have disproportionate effects on the global ecosystem and have been shown to vary geographically in extinction magnitude and rate of recovery.
Here, we analyze regional diversity and turnover patterns for the paleocontinents of Laurentia, Baltica, and Avalonia spanning the Late Ordovician mass extinction and Early Silurian recovery. Using a database of genus occurrences for inarticulate and articulate brachiopods, bivalves, anthozoans, and trilobites, we show that sampling-standardized diversity trends differ for the three regions. Diversity rebounded to pre-extinction levels within 5 Myr in the paleocontinent of Laurentia, compared with 15 Myr or longer for Baltica and Avalonia. This increased rate of recovery in Laurentia was due to both lower Late Ordovician extinction rates and higher Early Silurian origination rates relative to the other continents. Using brachiopod data, we dissected the Rhuddanian recovery into genus origination and invasion. This analysis revealed that standing diversity in the Rhuddanian consisted of a higher proportion of invading taxa in Laurentia than in either Baltica or Avalonia. Removing invading genera from diversity counts caused Rhuddanian diversity to fall in Laurentia. However, Laurentian diversity still rebounded to pre-extinction levels within 10 Myr of the extinction event, indicating that genus origination rates were also higher in Laurentia than in either Baltica or Avalonia. Though brachiopod diversity in Laurentia was lower than in the higher-latitude continents prior to the extinction, increased immigration and genus origination rates made it the most diverse continent following the extinction. Higher rates of origination in Laurentia may be explained by its large size, paleogeographic location, and vast epicontinental seas. It is possible that the tropical position of Laurentia buffered it somewhat from the intense climatic fluctuations associated with the extinction event, reducing extinction intensities and allowing for a more rapid rebound in this region. Hypotheses explaining the increased levels of invasion into Laurentia remain largely untested and require further scrutiny. Nevertheless, the Late Ordovician mass extinction joins the Late Permian and end-Cretaceous as global extinction events displaying an underlying spatial complexity.
We assessed selective extinction patterns in bivalves during a late Neogene mass extinction event observed along the temperate Pacific coast of South America. The analysis of 99 late Neogene and Quaternary fossil sites (recorded from 7°S to 55°S), yielding ∼2800 occurrences and 118 species, revealed an abrupt decline in Lyellian percentages during the late Neogene–Pleistocene, suggesting the existence of a mass extinction that decimated ∼66% of the original assemblage. Using the late Neogene data set (n = 59 species, 1346 occurrences), we tested whether the extinction was nonrandom according to taxonomic structure, life habit, geographic range, and body size. Our results showed that the number of higher taxa that went extinct was not different than expected by random. At first sight, extinction was selective only according to life habit and geographic range. Nevertheless, when phylogenetic effects were accounted for, body size also showed significant selectivity. In general, epifaunal, small-sized (after phylogenetic correction), and short-ranged species tended to have increased probability of extinction. This is verified by strong interactions between the variables herein analyzed, suggesting the existence of nonlinear effects on extinction chances. In the heavily decimated epifaunal forms, survival was not enhanced by widespread ranges or larger body sizes. Conversely, the widespread and large-sized infaunal forms tended to have lower probability of extinction. Overall, the ultimate extinction of late Neogene bivalve species along the Pacific coast of South America seems to have been determined by a complex interplay of ecological and historical (phylogenetic) effects.
Most evolutionary innovations—power-enhancing phenotypes previously absent in a lineage—have arisen multiple times within major clades. This repetition permits a comparative approach to ask how, where, when, in which clades, and under which circumstances adaptive innovations are acquired and secondarily lost. I use new and literature-based data on the phylogeny, functional morphology, and fossil record of gastropods to explore the acquisition and loss of the siphonal canal and its variations in gastropods. The siphonal indentation, canal, notch, or tube at the front end of the shell is associated in living gastropods with organs that detect chemical signals directionally and at a distance in an anteriorly restricted inhalant stream of water.
Conservative estimates indicate that the siphonate condition arose 23 times and was secondarily lost 17 times. Four siphonate clades have undergone prodigious diversification. All siphonate gastropods have a shell whose axis of coiling lies at a low angle above the plane of the aperture (retroaxial condition). In early gastropods, the siphonal canal was short and more or less confined to the apertural plane. Later (mainly Cretaceous and Cenozoic) variations include a dorsally deflected canal, a long canal, and a closed canal. The closed siphonal canal, in which the edges join to form a tube, arose 15 times, all in the adult stages of caenogastropods with determinate growth.
Gastropods in which the siphonate condition arose were mobile, bottom-dwelling, microphagous animals. Active predaceous habits became associated with the siphonate condition in the Mesozoic and Cenozoic Purpurinidae-Latrogastropoda clade. Loss of the siphonate condition is associated with nonmarine habits, miniaturization, and especially with a sedentary or slow-moving mode of life.
The siphonate condition arose seven times each during the early to middle Paleozoic, the late Paleozoic, and the early to middle Mesozoic, and only once each during the Late Cretaceous and Cenozoic. Well-adapted incumbents prevented most post-Jurassic clades from evolving a siphonal indentation and its associated organs. Dorsally deflected, long, and closed canals are known only from Cretaceous and Cenozoic marine gastropods, and represent improvements in sensation and passive armor.
In a discussion of contrasting ecologies of clades that gained and lost the siphonate condition, I argue that macroevolutionary trends in the comings and goings of innovations and clades must incorporate ecological and functional data. Acquisitions of energy-intensive, complex innovations that yield greater power have a greater effect on ecosystems, communities, and their resident clades than do reversals, which generally reflect energy savings.