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Diversification and extinction in the history of life.

Analysis of the fossil record of microbes, algae, fungi, protists, plants, and animals shows that the diversity of both marine and continental life increased exponentially since the end of the Precambrian. This diversification was interrupted by mass extinctions, the largest of which occurred in the Early Cambrian, Late Ordovician, Late Devonian, Late Permian, Early Triassic, Late Triassic, and end-Cretaceous. Most of these extinctions were experienced by both marine and continental organisms. As for the periodicity of mass extinctions, no support was found: Seven mass extinction peaks in the last 250 million years are spaced 20 to 60 million years apart.

Animals↗

Long-term persistence of species and the SLOSS problem.

The single large or several small (SLOSS) problem has been addressed in a large number of empirical and theoretical studies, but no coherent conclusion has yet been reached. Here I study the SLOSS problem in the context of metapopulation dynamics. I assume that there is a fixed total amount A(0) of habitat available, and I derive formulas for the optimal number n and area A of habitat patches, where n=A(0)/A. I consider optimality in two ways. First, I attempt to maximize the time to metapopulation extinction, which is a relevant measure for metapopulation viability for rare and threatened species. Second, I attempt to maximize the metapopulation capacity of the habitat patch network, which corresponds both with maximizing the distance to the deterministic extinction threshold and with maximizing the fraction of occupied patches. I show that in the typical case, a small number of large patches maximizes the metapopulation capacity, while an intermediate number of habitat patches maximizes the time to extinction. The main conclusion stemming from the analysis is that the optimal number of patches is largely affected by the relationship between habitat patch area and rates of immigration, emigration and local extinction. Here this relationship is summarized by a single factor zeta, termed the patch area scaling factor.

Animals↗

Ancon sheep: a now disproven example of macroevolution.

The Ancon breed of sheep provided, for decades, a critical support for the existence of major evolutionary changes or jumps called "sports." Putative examples of sports have been used as evidence of rapid macroevolution since Darwin first discussed the Ancon sheep mutation in 1859. Ancon sheep had very short legs that were considered an advantage for shepherds because the sheep were less likely to escape over fences. Many textbooks and articles implied that the breed was an example of how a major new trait could evolve in a single generation. The Ancon sheep example has been used both to prove gradual Darwinism and also to argue for rapid evolution as opposed to gradualism. It now is recognized that Ancon sheep resulted from genetic diseases, and that they usually suffered from achondroplasia. The so-called new breed had so many major health problems that the condition caused the breed's extinction decades ago, yet was mentioned in textbooks as evidence for evolutionary jumps as recently as 2005.

Animals↗

Evolution as a self-organized critical phenomenon.

We present a simple mathematical model of biological macroevolution. The model describes an ecology of adapting, interacting species. The environment of any given species is affected by other evolving species; hence, it is not constant in time. The ecology as a whole evolves to a "self-organized critical" state where periods of stasis alternate with avalanches of causally connected evolutionary changes. This characteristic behavior of natural history, known as "punctuated equilibrium," thus finds a theoretical explanation as a self-organized critical phenomenon. The evolutionary behavior of single species is intermittent. Also, large bursts of apparently simultaneous evolutionary activity require no external cause. Extinctions of all sizes, including mass extinctions, may be a simple consequence of ecosystem dynamics. Our results are compared with data from the fossil record.

Adaptation, Biological↗

The phylogeny of a species-level tendency: species heritability and possible deep origins of Bergmann's rule in tetrapods.

One of the most widely recognized generalizations in biology is Bergmann's rule, the observation that, within species of birds and mammals, body size tends to be inversely related to ambient temperature. Recent studies indicate that turtles and salamanders also tend to follow Bergmann's rule, which hints that this species-level tendency originated early in tetrapod history. Furthermore, exceptions to Bergmann's rule are concentrated within squamate reptiles (lizards and snakes), suggesting that the tendency to express a Bergmann's rule cline may be heritable at the species level. We evaluated species-level heritability and early origination of Bergmann's rule by mapping size-latitude relationships for 352 species onto a tetrapod phylogeny. When the largest available dataset is used, Bergmann's rule shows significant phylogenetic signal, indicating species-level heritability. This represents one of the few demonstrations of heritability for an emergent species-level property and the first for an ecogeographic rule. When species are discretely coded as showing either Bergmann's rule or its converse, parsimony reconstructions suggest that: (1) the tendency to follow Bergmann's rule is ancestral for tetrapods, and (2) most extant species that express the rule have retained this tendency from that ancient ancestor. The first inference also generally holds when the discrete data or size-latitude correlation coefficients are analyzed using maximum likelihood, although the results are only statistically significant for some versions of the discrete analyses. The best estimates of ancestral states suggest that the traditional adaptive explanation for Bergmann's rule-conservation of metabolic heat-was not involved in the origin of the trait since that origin predates the evolution of endothermy. A more general thermoregulatory hypothesis could apply to endotherms and some ectotherms, but fails to explain why salamanders have retained Bergmann's rule. Thus, if thermoregulation underlies the origin of a Bergmann's rule tendency, this trait may have been continuously maintained while its cause changed. Alternatively, thermoregulation may not underlie Bergmann's rule in any tetrapod group. The results also suggest that many extinct groups not included in our analyses followed Bergmann's rule.

Animals↗

The ethanol self-administration context as a reinstatement cue: acute effects of naltrexone.

A major difficulty in treating alcohol addiction is the high rate of relapse even after prolonged abstinence. Relapse can be triggered by several factors, including stress, re-exposure to the drug, conditioned discrete stimuli and exposure to the context in which alcohol consumption occurred. The present study investigated the role played by the environmental context on ethanol relapse using an extinction/reinstatement animal model: rats were trained to self-administer ethanol in a distinctive context, and extinction occurred in a setting that differed by visual, tactile and olfactory properties; reinstatement was tested by placing the animals into the ethanol-associated context in the absence of ethanol. We found that re-exposure to the ethanol-associated context significantly increased responses on the ethanol-paired lever. The increase in responding required the presence of the complete configuration of the multimodal context. The non-selective opioid receptor antagonist naltrexone (0.3 mg/kg) administered 20 min prior to the reinstatement test significantly attenuated context-induced reinstatement of lever press responding, compared with saline-treated subjects. These data indicate that the environmental context associated with ethanol availability influences ethanol-seeking behavior in the rat, and that endogenous opioids are involved in this process. Our findings are in accordance with clinical reports demonstrating naltrexone efficacy in the treatment of alcohol relapse in humans, and indicate that the context-induced reinstatement model described here may be useful to investigate the biological mechanisms underlying alcohol relapse.

Alcohol Drinking↗

[Computer simulation of different types of evolutionary process].

The computer model of two alternative variants of biological evolution is proposed. The first variant supposes random while the second--directed change of individual features, thus corresponding to the Darwinian and non-Darwinian evolution. The evolution of fish communities in fresh waters serves as a particular example. The model is executed using object-oriented method of programming and mathematical apparatus of fuzzy logics. The investigation of the model showed that process of Darwinian evolution is connected with significantly greater species diversity and variability of evolutionary process trajectories than non-Darwinian one. On the other hand, non-Darwinian type of evolution provides fast achievement of high individual fitness, especially under conditions of constant environment. Non-Darwinian type evolution failed in big evolutionary alteration (for example, transition to predation); while the Darwinian evolution under the same conditions can produce such alterations though it took more time and many extinct species. Phylogenetic tree of Darwinian evolution is always more complex than of non-Darwinian one under the same conditions.

Animals↗

[Electrical reactions in the cat brain during differentiation of photic stimuli].

Study of EEG responses to photic conditioned stimuli in cats has shown that the extent of brain activation depends not on the physical strength of the stimulus, but on its biological significance. The longest activation is recorded to a positive signal, even if it is the weakest flash of light. The intensity of EEG responses to negative stimuli is determined by the nearness of their physical parameters to those of the positive signal. Repeated presentation of a non-reinforced flash leads to a gradual extinction of EEG reactions. Brain activation in response to a reinforced positive signal persists unchanged even after a large number of its repetitions. The revealed properties of brain electrical reactions to stimuli of different significance accord with those previously recorded in response to acoustic stimuli. In both cases the data are in agreement with hypothesis of a central system of signal analysis, which performs the estimation of stimuli according to their biological quality.

Animals↗

Effects of competition, colonization, and extinction on rodent species diversity.

Analyses of long-term experimental data from the Chihuahuan desert revealed that species diversity of other rodents was higher on plots from which kangaroo rats (Dipodomys spp.) had been removed. The difference was due to consistently higher colonization and lower extinction probabilities of small granivorous rodents in the absence of competitively dominant kangaroo rats. The results of this ecosystem experiment demonstrate the importance of both competitive exclusion and metapopulation dynamics for biological diversity in a natural community.

Animals↗

Population dynamics of clock-controlled biological species: models and why circadian rhythms are circadian.

Conventional models of population dynamics of biological species do not take into account the fact that activity of most living organisms is under control of biological clocks, which are oscillators with periods near 1 day, 1 year, and so on. Here proposed are new categories of population dynamics models which are combinations of Lotka-Volterra type of equations (as a simplest example; in fact, an arbitrary type may be used) and equations of the biological oscillators. Such models may be particularly useful in addressing to those problems which are inaccessible with conventional models. A simple example of such models with distributed clock periods is proposed and studied numerically and analytically to consider why periods of biological rhythms are not precisely one day or one year, etc., as is often questioned. For this purpose, a notion of survival index is introduced to measure the degree of success in survival. It turns out that clock periods equal or close to that of the milieu are not necessarily advantageous for survival and can even lead to extinction; survival is then most successful at the edge of entrainment. This may provide a clue to answer the question. It is also found that survival is difficult for those species which cannot be entrained by the environmental cycle.

Animals↗

Molar microwear of subfossil lemurs: improving the resolution of dietary inferences.

In this study we use molar microwear analyses to examine the trophic distinctions among various taxa of Malagasy subfossil lemurs. High resolution casts of the teeth of Megaladapis, Archaeolemur, Palaeopropithecus, Babakotia, and Hadropithecus were examined under a scanning electron microscope. Megaladapis was undoubtedly a browsing folivore, but there are significant differences between species of this genus. However, dietary specialists appear to be the exception; for example, Palaeopropithecus and Babakotia probably supplemented their leaf-eating with substantial amounts of seed-predation, much like modern indrids. Hadropithecus was decidedly not like the modern gelada baboon, but probably did feed on hard objects. Evidence from microwear and coprolites suggests that Archaeolemur probably had an eclectic diet that differed regionally and perhaps seasonally. Substantial trophic diversity within Madgascar's primate community was diminished by the late Quaternary extinctions of the large-bodied species (>9 kg).

Adaptation, Physiological↗

Are certain life histories particularly prone to local extinction?

Using stochastic simulations and elasticity analysis, we show that there are inherent differences in the risk of extinction between life histories with different demographies. Which life history is the most vulnerable depends on which vital rate varies. When juvenile survival varies semelparous organisms with delayed reproduction are the most vulnerable ones, while a varying developmental rate puts a greater threat to semelparous organisms with rapid development. Iteroparous organisms are the most vulnerable ones when adult survival varies. Generally, we do not expect to observe organisms in nature having variation in vital rates that produce a high risk of extinction. Given the results here we therefore predict that iteroparous organisms should show low variation in adult survival. Moreover, we predict that semelparous organisms should show low variation in juvenile survival and low variation in developmental rate. The effect of temporal correlation on extinction risk is most pronounced in organisms with semelparous life histories.

Animals↗

Computational method for determination of bone and joint loads using bone density distributions.

Because bone structure is influenced by mechanical loading during ontogeny, the geometry and density distribution of bones contain information about their loading histories. Based on a mathematical theory relating stress history to bone remodeling, we have developed a method to determine dominant bone loading conditions using an optimization procedure. We applied this load determination method using a simplified two-dimensional bone-end finite element model, for which a standard density distribution had been calculated under a given set of loading conditions. With this density distribution, the optimization procedure was used to determine the original loads from a broad set of many plausible basic load distributions and locations. The optimization procedure adjusted the magnitude of each basic load to achieve the desired tissue level attractor stress stimulus throughout the model. The results show that the density-based bone load determination method yields accurate results for basic test cases and, thus, may have potential for estimating in vivo bone loads for both extant and extinct animals.

Animals↗

Food chain dynamics in the chemostat.

The asymptotic behavior of a tri-trophic food chain model in the chemostat is studied. The Monod-Herbert growth model is used for all trophic levels. The analysis is carried out numerically, by finding both local and global bifurcations of equilibria and of limit cycles with respect to two chemostat control parameters: the dilution rate of the chemostat and the concentration of input substrate. It is shown that the bifurcation structure of the food chain model has much in common with the bifurcation structure of a one-dimensional map with two turning points. This map is used to explain how attractors are created and destroyed under variation of the bifurcation parameters. It is shown that low as well as high concentration of input substrate can lead to extinction of the highest trophic level.

Animals↗

Large-body impact and extinction in the Phanerozoic.

The kill curve for Phanerozoic marine species is used to investigate large-body impact as a cause of species extinction. Current estimates of Phanerozoic impact rates are combined with the kill curve to produce an impact-kill curve, which predicts extinction levels from crater diameter, on the working assumption that impacts are responsible for all "pulsed" extinctions. By definition, pulsed extinction includes the approximately 60% of Phanerozoic extinctions that occurred in short-lived events having extinction rates greater than 5%. The resulting impact-kill curve is credible, thus justifying more thorough testing of the impact-extinction hypothesis. Such testing is possible but requires an exhaustive analysis of radiometric dating of Phanerozoic impact events.

Biological Evolution↗

Dynamic response of Permian brachiopod communities to long-term environmental change.

The fossil record preserves numerous natural experiments that can shed light on the response of ecological communities to environmental change. However, directly observing the community dynamics of extinct organisms is not possible. As an alternative, neutral ecological models suggest that species abundance distributions reflect dynamical processes like migration, competition, recruitment, and extinction. Live-dead comparisons suggest that such distributions can be faithfully preserved in the rock record. Here we use a maximum-likelihood approach to show that brachiopod (lamp shell) abundance distributions from four temporally distinct ecological landscapes from the Glass Mountains, Texas (of the Permian period), exhibit significant differences. Further, all four are better fitted by zero-sum multinomial distributions, characteristic of Hubbell's neutral model, than by log-normal distributions, as predicted by the traditional ecological null hypothesis. Using the neutral model as a guide, we suggest that sea level fluctuations spanning about 10 Myr altered the degrees of isolation and exchange among local communities within these ecological landscapes. Neither these long-term environmental changes nor higher-frequency sea level fluctuations resulted in wholesale extinction or major innovation within evolutionary lineages.

Animals↗

Positive Darwinian evolution in human influenza A viruses.

We earlier suggested that type A human influenza virus genes undergo positive Darwinian selection through immune surveillance. This requires more favorable amino acid replacements fixed in antigenic sites among the surviving lineages than among the extinct lineages. We now show that viral hemagglutinins fix proportionately more amino acid replacements in antigenic sites in the trunk of the evolutionary tree (survivors) than in the branches (nonsurvivors), demonstrating that type A human influenza virus is undergoing positive Darwinian evolution. The hemagglutinin gene is evolving 3 times faster than the nonstructural gene and the average age of the sampled nonsurvivors is only 1.6 years, so that extinction is not only common but rapid.

Amino Acid Sequence↗

Climate change, species-area curves and the extinction crisis.

An article published in the journal Nature in January 2004-in which an international team of biologists predicted that climate change would, by 2050, doom 15-37% of the earth's species to extinction-attracted unprecedented, worldwide media attention. The predictions conflict with the conventional wisdom that habitat change and modification are the most important causes of current and future extinctions. The new extinction projections come from applying a well-known ecological pattern, the species-area relationship (SAR), to data on the current distributions and climatic requirements of 1103 species. Here, I examine the scientific basis to the claims made in the Nature article. I first highlight the potential and pitfalls of using the SAR to predict extinctions in general. I then consider the additional complications that arise when applying SAR methods specifically to climate change. I assess the extent to which these issues call into question predictions of extinctions from climate change relative to other human impacts, and highlight a danger that conservation resources will be directed away from attempts to slow and mitigate the continuing effects of habitat destruction and degradation, particularly in the tropics. I suggest that the most useful contributions of ecologists over the coming decades will be in partitioning likely extinctions among interacting causes and identifying the practical means to slow the rate of species loss.

Animals↗