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A model of onshore-offshore change in faunal diversity.

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.

Algorithms↗

A molecular timescale for vertebrate evolution.

A timescale is necessary for estimating rates of molecular and morphological change in organisms and for interpreting patterns of macroevolution and biogeography. Traditionally, these times have been obtained from the fossil record, where the earliest representatives of two lineages establish a minimum time of divergence of these lineages. The clock-like accumulation of sequence differences in some genes provides an alternative method by which the mean divergence time can be estimated. Estimates from single genes may have large statistical errors, but multiple genes can be studied to obtain a more reliable estimate of divergence time. However, until recently, the number of genes available for estimation of divergence time has been limited. Here we present divergence-time estimates for mammalian orders and major lineages of vertebrates, from an analysis of 658 nuclear genes. The molecular times agree with most early (Palaeozoic) and late (Cenozoic) fossil-based times, but indicate major gaps in the Mesozoic fossil record. At least five lineages of placental mammals arose more than 100 million years ago, and most of the modern orders seem to have diversified before the Cretaceous/Tertiary extinction of the dinosaurs.

Animals↗

Do island populations have less genetic variation than mainland populations?

Island populations are much more prone to extinction than mainland populations. The reasons for this remain controversial. If inbreeding and loss of genetic variation are involved, then genetic variation must be lower on average in island than mainland populations. Published data on levels of genetic variation for allozymes, nuclear DNA markers, mitochondrial DNA, inversions and quantitative characters in island and mainland populations were analysed. A large and highly significant majority of island populations have less allozyme genetic variation than their mainland counterparts (165 of 202 comparisons), the average reduction being 29 per cent. The magnitude of differences was related to dispersal ability. There were related differences for all the other measures. Island endemic species showed lower genetic variation than related mainland species in 34 of 38 cases. The proportionate reduction in genetic variation was significantly greater in island endemic than in nonendemic island populations in mammals and birds, but not in insects. Genetic factors cannot be discounted as a cause of higher extinction rates of island than mainland populations.

Animals↗

Evidence from molecular systematics for decreased avian diversification in the pleistocene Epoch.

Pleistocene glaciations have been suggested as major events influencing speciation rates in vertebrates. Avian paleontological studies suggest that most extant species evolved in the Pleistocene Epoch and that species' durations decreased through the Pleistocene because of heightened speciation rates. Molecular systematic studies provide another data base for testing these predictions. In particular, rates of diversification can be determined from molecular phylogenetic trees. For example, an increasing rate of speciation (but constant extinction) requires shorter intervals between successive speciation events on a phylogenetic tree. Examination of the cumulative distribution of reconstructed speciation events in mtDNA phylogenies of 11 avian genera, however, reveals longer intervals between successive speciation events as the present time is approached, suggesting a decrease in net diversification rate through the Pleistocene Epoch. Thus, molecular systematic studies do not indicate a pulse of Pleistocene diversification in passerine birds but suggest, instead, that diversification rates were lower in the Pleistocene than for the preceding period. Documented habitat shifts likely led to the decreased rate of diversification, although from molecular evidence we cannot discern whether speciation rates decreased or extinction rates increased.

Animals↗

Blowout bifurcation with non-normal parameters in population dynamics.

Interesting dynamic behaviors have been associated with the transverse stability of an invariant subspace (e.g., riddled basins or on-off intermittency). Recently, these complex behaviors have been generalized to cases of models with non-normal parameters showing that these behaviors might be widespread. In this Brief Report I demonstrate this type of complex dynamics in models of interacting populations for which the invariant subspace is defined by species extinction. In the context of the current biodiversity crisis, implications of these behaviors for population biology are discussed.

Journal Article↗

Spatially synchronous extinction of species under external forcing.

More than 99% of the species that ever existed on the surface of the Earth are now extinct and their extinction on a global scale has been a puzzle. One may think that a species under an external threat may survive in some isolated locations leading to the revival of the species. Using a general model we show that, under a common external forcing, the species with a quadratic saturation term first undergoes spatial synchronization and then extinction. The effect can be observed even when the external forcing acts only on some locations provided the dynamics contains a synchronizing term. Absence of the quadratic saturation term can help the species to avoid extinction.

Animals↗

Dynamics of speciation and diversification in a metapopulation.

We develop a simple framework for modeling speciation and diversification as a continuous process of accumulation of genetic (or morphological) differences accompanied by species and subpopulation extinction and/or range expansion. This framework can be used to approach a number of questions such as species-area distribution, species-range size distribution, the rate of ecological turnover, asymmetries of range division between sister species, waiting time until speciation and extinction, the relationship between the geographic range size and the probability of speciation, the relationships between subpopulation-level parameters and metapopulation-level parameters, and the effects of taxonomic level on these rates, distributions, and parameters. We illustrate some of these applications using numerical simulations. We develop approximations describing the dependence of the number of different taxonomic units, their average range size, and the rate of their turnover on the system size, the rate of fixation of genetic (or morphological) changes in local demes, and the rate of local extinction and colonization.

Algorithms↗

The imprint of the geographical, evolutionary and ecological context on species-area relationships.

Species-area relationships (SAR) are fundamental in the understanding of biodiversity patterns and of critical importance for predicting species extinction risk worldwide. Despite the enormous attention given to SAR in the form of many individual analyses, little attempt has been made to synthesize these studies. We conducted a quantitative meta-analysis of 794 SAR, comprising a wide span of organisms, habitats and locations. We identified factors reflecting both pattern-based and dynamic approaches to SAR and tested whether these factors leave significant imprints on the slope and strength of SAR. Our analysis revealed that SAR are significantly affected by variables characterizing the sampling scheme, the spatial scale, and the types of organisms or habitats involved. We found that steeper SAR are generated at lower latitudes and by larger organisms. SAR varied significantly between nested and independent sampling schemes and between major ecosystem types, but not generally between the terrestrial and the aquatic realm. Both the fit and the slope of the SAR were scale-dependent. We conclude that factors dynamically regulating species richness at different spatial scales strongly affect the shape of SAR. We highlight important consequences of this systematic variation in SAR for ecological theory, conservation management and extinction risk predictions.

Biological Evolution↗

[Measurement of calcium content in the blood and other biological fluids by Arsenazo III complexone].

A method of photometric measurement of total calcium in the serum and other biological liquids using Arsenazo III color complexone is offered. Physicochemical characteristics of Arsenazo III (mol extinction, affinity for other ions, spectral characteristics, and its complex with calcium) are described. Calcium was measured in 750 normal subjects, its reference value was 2.15-2.75 mmol/liter, the mean value 2.47 mmole/liter. In semiautomated measurement CV% was about 5%. The reagent is convenient, particularly for automated analyzers; no other reagent are needed, it can be stored for a long time at room temperature, and gives stable results.

Adult↗

Quantitative genetics in conservation biology.

Most of the major genetic concerns in conservation biology, including inbreeding depression, loss of evolutionary potential, genetic adaptation to captivity and outbreeding depression, involve quantitative genetics. Small population size leads to inbreeding and loss of genetic diversity and so increases extinction risk. Captive populations of endangered species are managed to maximize the retention of genetic diversity by minimizing kinship, with subsidiary efforts to minimize inbreeding. There is growing evidence that genetic adaptation to captivity is a major issue in the genetic management of captive populations of endangered species as it reduces reproductive fitness when captive populations are reintroduced into the wild. This problem is not currently addressed, but it can be alleviated by deliberately fragmenting captive populations, with occasional exchange of immigrants to avoid excessive inbreeding. The extent and importance of outbreeding depression is a matter of controversy. Currently, an extremely cautious approach is taken to mixing populations. However, this cannot continue if fragmented populations are to be adequately managed to minimize extinctions. Most genetic management recommendations for endangered species arise directly, or indirectly, from quantitative genetic considerations.

Animals↗

[On the quantitative fluorometric determination of cellular DNA by ethidium bromide and the constancy of the quantum yield of the ethidium DNA-complex in the biological environment (author's transl)].

The fluorescent staining of fixed A9 cells (mouse fibroblasts) with ethidium bromide E has been investigated qualitatively and quantitatively. E is bound to DNA, RNA and protein. After enzymatic RNA digestion and staining at pH = 4.0 the dyestuff is bound to DNA specifically. The formed E-DNA-complex is stoichiometric if DNA will be saturated by highdye concentration, CF = 1.0 x 10(-2) M. The stoichiometric factor n = 0.21 was determined from Scatchard binding isotherms and by spectrophotometric titration. The binding of E to DNA and RNA has been investigated by binding isotherms. E is bound to DNA by intercalation and by ionic interaction. It is bound to RNA by ionic interaction only. The ionically bound dyestuff can be replaced by salts like LiCl from DNA and RNA. Only intercalated E remains. The competitive salt effect can be used to avoid the enzymatic RNA digestion in the specific staining of DNA. Within the cell the E-DNA-complex fluoresces strongly. Using a microspectrophotometer we determined the fluorescence intensity J and the extinction E in cell sections of q = 1 mu 2 area. The fluorescence was excited in the minimum of the absorbance of the E-DNA complex at lambda 1 = 365 nm. The extinction E2 was measured in the short wavelength band lambda 2 = 260 nm. Under these conditions we received a linear relation J = J (E2) up to high extinctions E2. The quantum yield Q of the E-DNA-complex is nearly constant and independent of the biological environment. Q is also unaltered by LiCl. Within the limits of error intercalated and ionic bound dye have the same quantum yeild. Under the conditions of stoichiometric staining the amount of E-DNA-complex and therefore DNA in the cell can be determined by J. The extinction coefficient epsilon 2 = 45 200 M-1 cm-1 of the E-DNA-complex at 260 nm was calculated from the concentration of dependence of the absorbance spectra using the law of mass action. By epsilon 2 the amount of DNA in the cell is accessible from J. According to our measurements with A9 cells the superficial density of DNA has the order of magnitude rho N = mN/q = 10(1) nmol cm-2, the amount mN of DNA per section 1 = 1 mu 2, mN = 10(-1) fmol, and the DNA concentration in the cell nucleus CN = 10(-1) M. The ray pass and diagram of electronics of the based microspectrophotometer are described.

Animals↗

Specific phobia: a disorder of fear conditioning and extinction.

Specific phobia is the most prevalent of the anxiety disorders. Although there have been relatively few studies of its psychobiology and pharmacotherapy, there is a rich laboratory of literature on fear conditioning and extinction and a clear evolutionary perspective. Advances in the cognitive-affective neuroscience of fear processing may ultimately lead to new approaches to the clinical management of phobias.

Adult↗

Effects of temporal variability on rare plant persistence in annual systems.

Traditional conservation biology regards environmental fluctuations as detrimental to persistence, reducing long-term average growth rates and increasing the probability of extinction. By contrast, coexistence models from community ecology suggest that for species with dormancy, environmental fluctuations may be essential for persistence in competitive communities. We used models based on California grasslands to examine the influence of interannual fluctuations in the environment on the persistence of rare forbs competing with exotic grasses. Despite grasses and forbs independently possessing high fecundity in the same types of years, interspecific differences in germination biology and dormancy caused the rare forb to benefit from variation in the environment. Owing to the buildup of grass competitors, consecutive favorable years proved highly detrimental to forb persistence. Consequently, negative temporal autocorrelation, a low probability of a favorable year, and high variation in year quality all benefited the forb. In addition, the litter produced by grasses in a previously favorable year benefited forb persistence by inhibiting its germination into highly competitive grass environments. We conclude that contrary to conventional predictions of conservation and population biology, yearly fluctuations in climate may be essential for the persistence of rare species in invaded habitats.

California↗

Foot-and-mouth disease virus evolution: exploring pathways towards virus extinction.

Foot-and-mouth disease virus (FMDV) is genetically and phenotypically variable. As a typical RNA virus, FMDV follows a quasispecies dynamics, with the many biological implications of such a dynamics. Mutant spectra provide a reservoir of FMDV variants, and minority subpopulations may become dominant in response to environmental demands or as a result of statistical fluctuations in population size. Accumulation of mutations in the FMDV genome occurs upon subjecting viral populations to repeated bottleneck events and upon viral replication in the presence of mutagenic base or nucleoside analogs. During serial bottleneck passages, FMDV survive during extended rounds of replication maintaining low average relative fitness, despite linear accumulation of mutations in the consensus genomic sequence. The critical event is the occurrence of a low frequency of compensatory mutations. In contrast, upon replication in the presence of mutagens, the complexity of mutant spectra increases, apparently no compensatory mutations can express their fitness-enhancing potential, and the virus can cross an error threshold for maintenance of genetic information, resulting in virus extinction. Low relative fitness and low viral load favor FMDV extinction in cell culture. The comparison of the molecular basis of resistance to extinction upon bottleneck passage and extinction by enhanced mutagenesis is providing new insights in the understanding of quasispecies dynamics. Such a comparison is contributing to the development of new antiviral strategies based on the transition of viral replication into error catastrophe.

Biological Evolution↗

Allee effects and the risk of biological invasion.

The Allee effect is a nonlinear phenomenon exhibited in the population dynamics of sparse populations in which the per capita population growth rate increases with increasing population density. In sufficiently sparse populations, the Allee effect may lead to extinction and is known to generate a threshold in the probability of establishment when presented as a function of introduced population size or density. As introduced populations are generally small, Allee effects are probably common in biological invasions and their consideration is necessary for accurately assessing the risk of invasion by many species, including all sexually reproducing species. Bythotrephes longimanus, an invasive, freshwater, cladoceran zooplankter from Europe, is one such species. Here, I review a previously published model of the Allee effect for continuously sexually reproducing species. Then, I develop a new model for seasonally parthenogenetic species such as Bythotrephes, and thereby demonstrate the potential consequences of Allee effects. This result underscores the importance of considering nonlinear phenomena, including thresholds, when conducting risk analysis for biological introductions.

Animals↗

Decreased sexual signalling reveals reduced viability in small populations of the drumming wolf spider Hygrolycosa rubrofasciata.

One of the important goals in conservation biology is to determine reliable indicators of population viability. Sexual traits have been suggested to indicate population extinction risk, because they may be related to viability through condition dependence. Moreover, condition-dependent sexual traits may be more sensitive indicators of population viability than early life-history traits, because deleterious fitness effects of inbreeding tend to be expressed mainly at the end of the species' life history. However, empirical evidence of the significance of sexual behaviour for population viability is missing. In this study, we examined two male sexual traits and survival in 39 different-sized and isolated natural populations of the wolf spider, Hygrolycosa rubrofasciata. We also used several traits to estimate female reproductive success in 25 populations of H. rubrofasciata. According to previous studies, H. rubrofasciata males have a costly and condition-dependent acoustic signal, courtship drumming, which is the target of female choice. Males with a high drumming rate have considerably higher viability than males with a low drumming rate, and females that mate with the more actively drumming males gain genetic benefits in terms of increased offspring viability. Our results show that males in small populations had both lower survival and lower drumming rate than males in larger populations. However, we did not find any evidence for a decline in important early life-history traits (offspring number, hatching success or offspring body mass) or female body mass in small populations. Our results have two important messages for conservation biology. First, they show that sexual traits can be used as sensitive indicators of population viability. Second, the indirect benefits of female choice in terms of good genes might partially compensate for the reduction of viability in declining populations. Also, our results support the view that deleterious effects of small population size are expressed at the end of the species' life history.

Animal Communication↗

Patterns of endangerment in the hawaiian flora.

The Hawaiian flora, because of its great isolation, high levels of endemism, known lineages, and high rates of endangerment, offers unique opportunities to explore patterns of endangerment related to phylogeny, ecological and life history traits, and geographic patterns. Nine percent of the native flora of 1159 taxa are already extinct, and 52.5% are at risk (extinct, endangered, vulnerable, or rare). Risk is strongly associated with limited geographic distribution at several scales: endemic taxa (native only to the Hawaiian Islands) are at far greater risk than indigenous taxa (with both Hawaiian and extra-Hawaiian ranges); single-island endemics are more at risk than multi-island endemics; small islands have the highest proportion of endemic taxa at risk; and endemics with more limited habitat distributions (elevation, community type) are more at risk. Historic population density is a strong predictor of risk, and taxa with low historic population densities are at greatest risk with rapid anthropogenic changes. Among the major islands, Maui Nui has the highest percent of taxa that are extinct. Kaua'i has the lowest percent of extinct taxa and the highest proportion of single-island endemic taxa that are rare. Endemic taxa at risk are associated with distributions in shrublands, forests, bogs, and cliff habitats. Endemic taxa with distributions in low elevation dry habitats have the highest proportion of taxa at risk, but the greatest absolute numbers of taxa at risk have distributions in mesic lowland and montane forests, and in wet montane forests. The life history patterns associated with risk are complicated, and inclusion of the effects of evolutionary relationships (lineages) changes some of these patterns. Species level analyses without respect to lineage shows risk associated with monomorphic (hermaphroditic) breeding systems and bird pollination because of the large number of hermaphroditic, bird-pollinated species in the Campanulaceae. Analyses incorporating the effect of lineage greatly reduce the impact of large lineages and result in an association of risk with insect pollination, and no effect of breeding system. There is no association of lineage size and the percent of taxa at risk within the lineage; endemic taxa from lineages with large radiations are at no greater risk than endemic single-taxon lineages. The percentages of taxa at risk at the family level in the Hawaiian Islands and worldwide (excluding Hawaiian taxa) are positively correlated, although flowering plant families in the Hawaiian Islands have a much greater proportion of taxa at risk. Some of the approaches described here may be useful to predict geographical and biological patterns of endangerment in island and island-like ecosystems under increasing pressures of endangerment and extinction.

Cell Lineage↗