Palaeontological data and identifying mass extinctions

It is often assumed that mass extinctions may be read directly from the fossil record. However, recent work on the Cretaceous-Tertiary (K-T) boundary has shown the difficulty of doing this. For example, it is hard to tell whether the stratigraphic ranges of taxa are complete or not, and what the shape of an extinction really is. Range completeness may be assessed by (1) a statistical approach to the relative completeness of ranges of taxa, and (2) tests based on collecting effort near the ends of ranges. Tests carried out recently suggest that the record is good in parts and getting better. Hence, palaeontologists ought to be able to document the nature of extinction events ever more precisely.     

Evolutionary patterns from mass originations and mass extinctions

The Fossil Record 2 database gives a stratigraphic range of most known animal and plant families. We have used it to plot the number of families extant through time and argue for an exponential fit, rather than a logistic one, on the basis of power spectra of the residuals from the exponential. The times of origins and extinctions, when plotted for all families of marine and terrestrial organisms over the last 600 Myr, reveal different origination and extinction peaks. This suggests that patterns of biological evolution are driven by its own internal dynamics as well as responding to upsets from external causes. Spectral analysis shows that the residuals from the exponential model of the marine system are more consistent with 1/f noise suggesting that self-organized criticality phenomena may be involved.     

The evolutionary significance of mass extinctions

Local extinctions of populations, species or groups of species in a particular area are commonly observed by biologists. There are also historical records of the total extinction of single species such as the Dodo, the Great Auk and the Tasmanian Wolf. Mass extinctions are on a much larger scale, and their study is based on the fossil record. The aims of this review are to explore the nature of mass extinctions and their evolutionary significance. The key questions are: what is mass extinction, what are the causes of mass extinctions, do mass extinctions follow a regular pattern, and how do mass extinctions affect our understanding of evolutionary processes?     

Stepwise mass extinctions and impact events: Late Eocene to early Oligocene

Species ranges and relative abundances of dominant planktonic foraminifers of eight late Eocene to early Oligocene deep-sea sections are discussed to determine the nature and magnitude of extinctions and to investigate a possible cause-effect relationship between impact events and mass extinctions.Late Eocene extinctions are neither catastrophic nor mass extinctions, but occur stepwise over a period of about 1–2 million years. Four stepwise extinctions are identified at the middle/late Eocene boundary, the upperGlobigerapsis semiinvoluta zone, theG. semiinvoluta/Globorotalia cerroazulensis zone boundary and at the Eocene/Oligocene boundary. Each stepwise extinction event represents a time of accelerated faunal turnover characterized by generally less than 15% species extinct and in itself is not a significant extinction event. Relative species abundance changes at each stepwise extinction event, however, indicate a turnover involving > 60% of the population implying major environmental changes.There microtektite horizons are present in late Eocene sediments; one in the upperG. semiinvoluta zone (38.2 Ma) and two closely spaced layers only a few thousand years apart in the lower part of theGloborotalia cerroazulensis zone (37.2 Ma). Each of the three impact events appears to have had some effect on microplankton communities. However, the overriding factor that led to the stepwise mass extinctions may have been the result of multiple causes as there is no evidence of impacts associated with the step preceding, or the step following the deposition of the presently known microtektite horizons.     

Signatures of random and selective mass extinctions in phylogenetic tree balance

Current models of diversification with evolving speciation rates have trouble mimicking the extreme imbalance seen in estimated phylogenies. However, these models have not incorporated extinction. Here, we report on a simple simulation model that includes heritable and evolving speciation rates coupled with mass extinctions, Random (but not selective) mass extinctions, coupled with evolving among-lineage variation in speciation rates, increase imbalance of postrecovery clades. Thus, random mass extinctions are plausible contributors to the imbalance of modern clades. Paleontological evidence suggests that mass extinctions are often random with respect to ecological and morphological traits, consistent with our simulations. In contrast, evidence that the current anthropogenic mass extinction is phylogenetically selective suggests that the current extinction episode may be qualitatively different from past ones in the way it reshapes future biotas.     

Long-term evolution of an ecosystem with spontaneous periodicity of mass extinctions

Twenty years ago, after analysing palaeontological data, Raup and Sepkoski suggested that mass extinctions on Earth appear cyclically in time with a period of approximately 26 million years (My). To explain the 26 My period, a number of proposals were made involving, e.g., astronomical effects, increased volcanic activity, or the Earth's magnetic field reversal, none of which, however, has been confirmed. Here we study a spatially extended discrete model of an ecosystem and show that the periodicity of mass extinctions might be a natural feature of the ecosystem's dynamics and not the result of a periodic external perturbation. In our model, periodic changes of the diversity of an ecosystem and some of its other characteristics are induced by the coevolution of species. In agreement with some palaeontological data, our results show that the longevity of a species depends on the evolutionary stage at which the species is created. Possible further tests of our model are also discussed.     

Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates

The fossil record is our only direct means for evaluating shifts in biodiversity through Earth''s history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic–Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent ‘Lagerstätten effects’. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous–Palaeogene extinction.     

Human influence on distribution and extinctions of the late Pleistocene Eurasian megafauna

Late Pleistocene extinctions are of interest to paleontological and anthropological research. In North America and Australia, human occupation occurred during a short period of time and overexploitation may have led to the extinction of mammalian megafauna. In northern Eurasia megafaunal extinctions are believed to have occurred over a relatively longer period of time, perhaps as a result of changing environmental conditions, but the picture is much less clear. To consider megafaunal extinction in Eurasia, we compare differences in the geographical distribution and commonness of extinct and extant species between paleontological and archaeological localities from the late middle Pleistocene to Holocene. Purely paleontological localities, as well as most extinct species, were distributed north of archaeological sites and of the extant species, suggesting that apart from possible differences in adaptations between humans and other species, humans could also have a detrimental effect on large mammal distribution. However, evidence for human overexploitation applies only to the extinct steppe bison Bison priscus. Other human-preferred species survive into the Holocene, including Rangifer tarandus, Equus ferus, Capreolus capreolus, Cervus elaphus, Equus hemionus, Saiga tatarica, and Sus scrofa. Mammuthus primigenius and Megaloceros giganteus were rare in archaeological sites. Carnivores appear little influenced by human presence, although they become rarer in Holocene archaeological sites. Overall, the data are consistent with the conclusion that humans acted as efficient hunters selecting for the most abundant species. Our study supports the idea that the late Pleistocene extinctions were environmentally driven by climatic changes that triggered habitat fragmentation, species range reduction, and population decrease, after which human interference either by direct hunting or via indirect activities probably became critical.     

Long-term evolution of an ecosystem with spontaneous periodicity of mass extinctions

Twenty years ago, after analysing palaeontological data, Raup and Sepkoski suggested that mass extinctions on Earth appear cyclically in time with a period of approximately 26 million years (My). To explain the 26 My period, a number of proposals were made involving, e.g., astronomical effects, increased volcanic acitivity, or the Earth's magnetic field reversal, none of which, however, has been confirmed. Here we study a spatially extended discrete model of an ecosystem and show that the periodicity of mass extinctions might be a natural feature of the ecosystem's dynamics and not the result of a periodic external perturbation. In our model, periodic changes of the diversity of an ecosystem and some of its other characteristics are induced by the coevolution of species. In agreement with some palaeontological data, our results show that the longevity of a species depends on the evolutionary stage at which the species is created. Possible further tests of our model are also discussed.     

Earliest Triassic microbialites in the South China block and other areas: controls on their growth and distribution

Earliest Triassic microbialites (ETMs) and inorganic carbonate crystal fans formed after the end-Permian mass extinction (ca. 251.4 Ma) within the basal Triassic Hindeodus parvus conodont zone. ETMs are distinguished from rarer, and more regional, subsequent Triassic microbialites. Large differences in ETMs between northern and southern areas of the South China block suggest geographic provinces, and ETMs are most abundant throughout the equatorial Tethys Ocean with further geographic variation. ETMs occur in shallow-marine shelves in a superanoxic stratified ocean and form the only widespread Phanerozoic microbialites with structures similar to those of the Cambro-Ordovician, and briefly after the latest Ordovician, Late Silurian and Late Devonian extinctions. ETMs disappeared long before the mid-Triassic biotic recovery, but it is not clear why, if they are interpreted as disaster taxa. In general, ETM occurrence suggests that microbially mediated calcification occurred where upwelled carbonate-rich anoxic waters mixed with warm aerated surface waters, forming regional dysoxia, so that extreme carbonate supersaturation and dysoxic conditions were both required for their growth. Long-term oceanic and atmospheric changes may have contributed to a trigger for ETM formation. In equatorial western Pangea, the earliest microbialites are late Early Triassic, but it is possible that ETMs could exist in western Pangea, if well-preserved earliest Triassic facies are discovered in future work.     

Evolution of C isotopes in the Cambrian of China: implications for Cambrian subdivision and trilobite mass extinctions

Carbon and oxygen isotopes were studied in fossiliferous Cambrian carbonates in northwestern Hunan Province (South China) and in northern Anhui and southern Shandong provinces (North China). Two major C isotope excursions related to biological events occur in the Wangcun section (Yongshun County, northwestern Hunan), which consists of a slope carbonate sequence (510 m thick) containing abundant trilobites. The first C isotope excursion (δ¹³C value shifts from -2.3‰ to 2‰) occurs near the boundary between the Qingxudong and Aoxi formations, close to the traditional Lower-Middle Cambrian boundary. The second excursion (δ¹³C value shifts from 0‰ to 3‰) occurs in the interval between the Linguagnostus reconditus Zone and the Glyptagnostus reticulatus Zone. The base of the G. reticulatus Zone define the base of the Paibi Stage and Furongian Series. Similar C isotope excursions also occur in shallow - water carbonate sections in North China. In Jiagou section near Huainan (Anhui Province), recently considered an important interval for defining the lower-middle Cambrian boundary because of dramatic changes in the trilobite fauna (extinction of redlichiids and appearances of ptychopariids), a negative C isotope excursion (δ¹³C value shifts from +1.21‰ to -1.93‰) occurs at the top of the lower member of the Mantou Formation. In the Gushan section (Changqing County, Shandong Province), a C isotope excursion (δ¹³C value shifts from -0.04‰ to 2.23‰) occurs at the base of the Changshan Formation and is coincident with the base of the Chuangia Zone. This excursion can be correlated with the excursion in the lower part of Glyptagnostus reticulatus Zone in the Wangcun section. The above two distinct C isotope excursions, which occur both in slope carbonates in South China and in shallow - water carbonates in North China, have also been recognized in Cambrian sections on other continents, and they coincide with global mass extinctions of trilobites. The two excursions evidently reflect global changes of Cambrian sea level, and they have utility for Cambrian subdivisions and for both regional and global stratigraphic correlation. In addition, a negative carbon excursion below the base of the Ptychagnostus atavus Zone in the Wangcun section supports previous suggestions that the FAD of P. atavus can be considered as a global correlatable horizon within the middle Cambrian.     

Macroinvertebrate distribution in streams: a comparison of CA ordination with biotic indices

Macroinvertebrates were collected in running waters in Italy, analyzed with correspondence analysis (CA) and with the calculation of 8 biotic indices. Then the CA ordination axes were correlated with 19 environmental variables and with biotic indices.The first CA axis is easily interpreted as an upstream-downstream gradient and is correlated with physical factors (particle size, slope etc.), whereas the second axis separated permanent waters from temporary ones.The first CA axis correlated with many biotic indices suggesting that biotic indices are strongly influenced by physical factors. Multiple regressions with 2 biotic indices as criterion and the 19 environmental factors as predictor variables confirm the importance of physical factors in determining the values of the biotic indices.The advantages and drawbacks of the use of CA instead of biotic indices is discussed.     

Abiotic and biotic factors influencing the winter distribution of predatory insects

Summary Various environmental factors were investigated to analyse those involved in successful overwintering and possibly overwintering site selection for Tachyporus hypnorum and Demetrias atricapillus, both important coleopteran predators of cereal aphids. The results of the study indicated food supply to be important for both predator species during the winter period, although the role of biotic factors in site selection in the autumn could not be clearly demonstrated. The winter distribution of the two species could, however, be explained well in terms of abiotic factors. It is suggested that these and other similar predator species have well-defined overwintering requirements and that these can be exploited in the management of field boundary habitats.     

Uncertainty in identifying local extinctions: the distribution of missing data and its effects on biodiversity measures

Identifying local extinctions is integral to estimating species richness and geographic range changes and informing extinction risk assessments. However, the species occurrence records underpinning these estimates are frequently compromised by a lack of recorded species absences making it impossible to distinguish between local extinction and lack of survey effort—for a rigorously compiled database of European and Asian Galliformes, approximately 40% of half-degree cells contain records from before but not after 1980. We investigate the distribution of these cells, finding differences between the Palaearctic (forests, low mean human influence index (HII), outside protected areas (PAs)) and Indo-Malaya (grassland, high mean HII, outside PAs). Such cells also occur more in less peaceful countries. We show that different interpretations of these cells can lead to large over/under-estimations of species richness and extent of occurrences, potentially misleading prioritization and extinction risk assessment schemes. To avoid mistakes, local extinctions inferred from sightings records need to account for the history of survey effort in a locality.