Are Namibian “Fairy Circles” the Consequence of Self-Organizing Spatial Vegetation Patterning?

Causes of over-dispersed barren “fairy circles” that are often surrounded by ca. 0.5 m tall peripheral grasses in a matrix of shorter (ca. 0.2 m tall) grasses in Namibian grasslands remain mysterious. It was hypothesized that the fairy circles are the consequence of self-organizing spatial vegetation patterning arising from resource competition and facilitation. We examined the edaphic properties of fairy circles and variation in fairy circle size, density and landscape occupancy (% land surface) with edaphic properties and water availability at a local scale (<50 km) and with climate and vegetation characteristics at a regional scale. Soil moisture in the barren fairy circles declines from the center towards the periphery and is inversely correlated with soil organic carbon, possibly indicating that the peripheral grass roots access soil moisture that persists into the dry season within fairy circles. Fairy circle landscape occupancy is negatively correlated with precipitation and soil [N], consistent with fairy circles being the product of resource-competition. Regional fairy circle presence/absence is highly predictable using an empirical model that includes narrow ranges of vegetation biomass, precipitation and temperature seasonality as predictor variables, indicating that fairy circles are likely a climate-dependent emergent phenomenon. This dependence of fairy circle occurrence on climate explains why fairy circles in some locations may appear and disappear over time. Fairy circles are only over-dispersed at high landscape occupancies, indicating that inter-circle competition may determine their spacing. We conclude that fairy circles are likely to be an emergent arid-grassland phenomenon that forms as a consequence of peripheral grass resource-competition and that the consequent barren circle may provide a resource-reservoir essential for the survival of the larger peripheral grasses and provides a habitat for fossicking fauna.     

Adopting a spatially explicit perspective to study the mysterious fairy circles of Namibia

The mysterious ‘fairy circles’ are vegetation‐free discs that cover vast areas along the pro‐Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25‐ha aerial images of north‐west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self‐organized vegetation pattern. Finally, we test if an existing partial‐differential‐equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key‐processes in arid areas such as plant competition for water and local resource‐biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas‐leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self‐organized vegetation patterns that emerge from positive biomass‐water feedbacks involving water transport by extended root systems and soil‐water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self‐organization to match the birth‐ and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.     

Experiments Testing the Causes of Namibian Fairy Circles

The grasslands on the sandy soils of the eastern edge of the Namib Desert of Namibia are strikingly punctuated by millions of mostly regularly-spaced circular bare spots 2 to 10 m or more in diameter, generally with a margin of taller grasses. The causes of these so called fairy circles are unknown, but several hypotheses have been advanced. In October 2009, we set up experiments that specifically tested four hypothesized causes, and monitored these 5 times between 2009 and 2015. Grass exclusion in circles due to seepage of subterranean vapors or gases was tested by burying an impermeable barrier beneath fairy circles, but seedling density and growth did not differ from barrier-less controls. Plant germination and growth inhibition by allelochemicals or nutrient deficiencies in fairy circle soils were tested by transferring fairy circle soil to artificially cleared circles in the grassy matrix, and matrix soil to fairy circles (along with circle to circle and matrix to matrix controls). None of the transfers changed the seedling density and growth from the control reference conditions. Limitation of plant growth due to micronutrient depletion within fairy circles was tested by supplementing circles with a micronutrient mixture, but did not result in differences in plant seedling density and growth. Short-range vegetation competitive feedbacks were tested by creating artificially-cleared circles of 2 or 4 m diameter located 2 or 6 m from a natural fairy circle. The natural circles remained bare and the artificial circles revegetated. These four experiments provided evidence that fairy circles were not caused by subterranean vapors, that fairy circle soil per se did not inhibit plant growth, and that the circles were not caused by micronutrient deficiency. There was also no evidence that vegetative feedbacks affected fairy circles on a 2 to 10 m scale. Landscape-scale vegetative self-organization is discussed as a more likely cause of fairy circles.     

Edaphic properties enable facilitative and competitive interactions resulting in fairy circle formation

Millions of generally regularly spaced, roughly circular barren patches called fairy circles occur in a narrow band ca 100 km inland of the south‐west African coast. These generally have conspicuously taller peripheral grasses in a shorter grass matrix. The origins of these fairy circles are controversial, but one possibility is that they are self‐organizing emergent vegetation patterns that are the consequence of interplay between positive (facilitative) and negative (competitive) interactions between grasses. We hypothesized that the coarse textured sand on which fairy circles occur creates a hydraulically and nutritionally connected landscape, in which neighbouring fairy circles competitively influence each other over several metres, while providing opportunity for focusing of resources around the peripheral grasses. To test our hypotheses we conducted three main groups of analyses: 1) we measured grass biomass to assess facilitative and competitive effects of the component grasses; 2) across a region with fairy circles we measured the size and density of fairy circles and correlated that with water infiltration rates into soil; 3) we measured the capacity of soil to conduct water pulses and ¹⁵N tracers. We found evidence of facilitative interactions in the periphery of the fairy circles and competitive suppression of the matrix grass proximal to the periphery. Across the region, fairy circle size was positively correlated with soil infiltration rates and negatively with precipitation. This suggests that fairy circles emerge in soils with high capacity for water flux that enables landscape hydraulic connectivity. Water‐ and ¹⁵N‐pulse experiments showed that edaphic resources were highly mobile, moving up to 7.5 m over a period of 1–3 weeks. We concluded that the evidence is consistent with an emergent vegetation pattern explanation for the origins of fairy circles and that the circles are more closely associated with a highly connective edaphic environment, rather than with particular biota.     

Herbivory by subterranean termite colonies and the development of fairy circles in SW Namibia

1. The growth pattern of Namibian fairy circles was examined in relation to environmental, termite, and plant variables to provide support for the sand termite (Psammotermes allocerus Silvestri) hypothesis of circle origin. 2. New and young circles were associated with the highest number of sand termites and their foraging grass sheetings which were both considerably lower in mature and senescent circles. 3. Circles increased in size with age, and apart from the senescent stage had higher soil moisture levels than the matrix. 4. In laboratory trials sand termites browsed seedling roots, negatively impacting root and leaf number. 5. This provides a potential mechanism for circle formation through central‐based foraging by P. allocerus which eliminates Stipagrostis grass clumps around its nest system. 6. The resulting bare disc is postulated to be maintained through a combination of a depleted seed bank, termite herbivory on seedlings, and excavations by pugnacious ants (Anoplolepis steingroeveri Forel).     

Namibian fairy circles and epithelial cells share emergent geometric order

Fairy circles are enigmatic features of the Namib desert landscape. They are large, almost perfectly circular patches of barren soil in sparse grassland. Although a matter of continuing debate, we make no attempt to explain their origin. The focus of our approach is a statistical analysis of the spatial patterns. These are easily accessible via aerial and satellite imagery. Observations over extended periods of time have revealed that they have a life-cycle of birth, growth and death. It has also been known for some time that the fairy circles are not randomly distributed. Our novel finding is that the connectivity patterns of fairy circles and metazoan epithelial cells are statistically indistinguishable, while remaining clearly distinct from other commonly observed polygonal patternings. This result identifies an analogy between the microscopic world of epithelial cells and the macroscopic realm of the Namib, suggesting that approaches developed specifically for the analysis of microscopic structures may extend into ecologically relevant, macroscopic dimensions.     

Strand switching during rolling circle replication of plasmid-like DNA circles in the mitochondria of the higher plant Chenopodium album (L.)

The structure of sigma-like mitochondrial DNA molecules prepared from suspension cultured cells of Chenopodium album (L.) was studied by electron microscopy. These molecules were highly variable in size, ranging from about 1 to 104 kb, and had single- and double-stranded regions typical for rolling circle replicating intermediates. Partial denaturation studies confirmed that these structures constitute rolling circles. Close inspection of the circle-tail junctions of the replication fork at high magnification suggests that in circles with a double-stranded tail, both strands of the tail seem to be covalently attached to the circle in about 27% of the molecules. This observation can be explained by a phenomenon called strand switching or strand splippage during rolling circle replication, similar to a mechanism proposed for bacterial replicons or in vitro replicating constructs harboring bacteriophage T4 replication origins.     

Size Estimates in Ebbinghaus Illusion in Adults and Children of Different Age

Psychophysical experiments were performed with the participation of 297 subjects (adults and 6- to 17-year-old children and adolescents). Perception of the size of a central stimulus surrounded by other stimuli was studied. Subjects had to estimate the size of a circle with a diameter of 22 angular min, which was surrounded by four similar figures with a diameter of either 11 or 44 angular min at a distance of either 11 or 33 angular min. The misperception of the stimulus size was dependent on the subject's age, the distance between the circles, and the size of the flanking circles. The smaller flanking circles located at any distance produced size underestimation in younger children; these circles led to size overestimation in case of their location at a short distance in most adults and adolescents. The larger flanking circles produced underestimation in all age groups, but, in adults, unlike children, this illusion decreased with increasing distance. The illusion mechanism and its possible connection with selective attention are discussed.     

Ants and the enigmatic Namibian fairy circles – cause and effect?

1. Parts of the Namibian landscape show extensive surface perturbation in the form of long‐lived, yet dynamic ‘fairy circles'. While exerting profound ecological effects on 7.3% of the land surface, the origin and nature of these large bare discs embedded in an arid grassland matrix remains unresolved. 2. We found no evidence to support the current hypothesis of a termite origin for fairy circles but instead observed a strong spatial association between fairy circles and large nests of the ant Black pugnacious ant Anoplolepis steingroeveri Forel, with much higher ant abundances on the circles compared with the matrix. 3. Aggression trials showed that different colonies of A. steingroeveri were located on different circles, and that the species was polydomous. 4. Fairy circles and Pogonomyrmex ant nests both have a bare disc surrounding the nest, are overdispersed (evenly spaced), and are associated with elevated soil moisture. Fairy circle soils exhibited a five‐fold increase in soil moisture when compared with the matrix. 5. Senescent Stipagrostis obtusa (Delile) Nees seedlings were only observed on the circles and not in the matrix, and were found to have a reduction in both root length and number of roots. 6. Anoplolepis steingroeveri excavated the root system of both S. obtusa seedlings on the disc and Stipagrostis ciliata (Desf.) de Winter grasses on the perimeter of the circles, where they tended honeydew‐secreting Meenoplidae bugs that fed on grass roots and culms. The bugs occurred almost exclusively on grasses associated with the circles. This ant–bug interaction is a possible mechanism for the observed reduction in root length and number of senescent grass seedlings on the circles.     

The variability of the circle of Willis: univariate and bivariate analysis

The results are presented of a statistical analysis of the variability of the circle of Willis using univariate and bivariate methods. For this purpose 100 circles of Willis were available. From each circle 19 indexes of arterial size were determined, the basilar artery was measured in two places. Half the circumference was measured. This data yielded no evidence of differences between left- and right-sided vessels in the sample. An important source of variation is the general size of all vessels considered. When the data are cleared from this general size variation, correlation coefficients reveal interesting relations between the vessels. The posterior communicating arteries are strongly related to the ipsilateral carotid artery, whereas a strong inverse relationship exists with the basilar artery and the precommunicating part of the ipsilateral posterior cerebral artery. These relationships can be understood from the expected patterns of the blood flow in these vessels. Similar relationships can be found in the anterior part of the circle of Willis and in the vertebro-basilar junction. In a different manner, based on previous haemodynamic studies, the relation between blood flow and vessel size within the circle of Willis can be demonstrated by relating the ratios of the sizes of afferent and efferent arteries to the sizes of the posterior communicating arteries, an "intuitive" model. The supposed correlations of the outcome of this "intuitive" model with the size of the communicating arteries appeared to by highly significant. It is concluded that the variations of the circle of Willis are related to the individual variations of the blood flow in this arterial network.     

Lifelong leukocyte telomere dynamics and survival in a free‐living mammal

Telomeres play a fundamental role in the maintenance of genomic integrity at a cellular level, and average leukocyte telomere length (LTL) has been proposed as a biomarker of organismal aging. However, studies tracking LTL across the entire life course of individuals are lacking. Here, we examined lifelong patterns of variation in LTL among four birth cohorts of female Soay sheep (Ovis aries) that were longitudinally monitored and sampled from birth to death. Over the first 4 months of life, there was within‐individual loss of LTL, consistent with findings in the human and primate literature, but there was little evidence of consistent LTL loss associated with age after this point. Overall, we observed only weak evidence of individual consistency in LTL across years and over the entire lifespan: Within‐individual variation was considerable, and birth cohorts differed markedly in their telomere dynamics. Despite the high levels of LTL variation within the lifetimes of individuals, there remained significant associations between LTL and longevity. Detailed analysis of the longitudinal data set showed that this association was driven by improved survival of individuals with longer LTL over the first 2 years of life. There was no evidence that LTL predicted survival in later adulthood. Our data provide the first evidence from a mammal that LTL can predict mortality and lifespan under natural conditions, and also highlight the potentially dynamic nature of LTL within the lifetimes of individuals experiencing a complex and highly variable environment.     

Genetic variation in the murine lifespan response to dietary restriction: from life extension to life shortening

Chronic dietary restriction (DR) is considered among the most robust life-extending interventions, but several reports indicate that DR does not always extend and may even shorten lifespan in some genotypes. An unbiased genetic screen of the lifespan response to DR has been lacking. Here, we measured the effect of one commonly used level of DR (40% reduction in food intake) on mean lifespan of virgin males and females in 41 recombinant inbred strains of mice. Mean strain-specific lifespan varied two to threefold under ad libitum (AL) feeding and 6- to 10-fold under DR, in males and females respectively. Notably, DR shortened lifespan in more strains than those in which it lengthened life. Food intake and female fertility varied markedly among strains under AL feeding, but neither predicted DR survival: therefore, strains in which DR shortened lifespan did not have low food intake or poor reproductive potential. Finally, strain-specific lifespans under DR and AL feeding were not correlated, indicating that the genetic determinants of lifespan under these two conditions differ. These results demonstrate that the lifespan response to a single level of DR exhibits wide variation amenable to genetic analysis. They also show that DR can shorten lifespan in inbred mice. Although strains with shortened lifespan under 40% DR may not respond negatively under less stringent DR, the results raise the possibility that life extension by DR may not be universal.     

Reconstruction of individual life histories of rodents from their teeth and bone

Age changes of bone tissue and molar cementum, and formation of daily layers in dentin of incisors enable us to estimate age and maximum lifespan, season of birth, season of death, sexual maturation, sex in adults, and in combination with tetracycline prebaiting, movements and space distribution of rodents. In hibernating rodents, the hibernation zone formed in incisor dentin can be used to assess inter- and intrapopulation variations in seasonal activity and growth. This approach is illustrated by examples from the study of rodents of the generaApodemus andSicista.     

Figural After-Effects or the Contrast Illusion

The contrast illusion was studied systematically for a long time by D. N. Uznadze. Experimental findings characteristic of this illusion were published by him first in the Russian (8) and then in the German (18) languages. To produce this illusion, two circles of different size were tachistoscopically presented to the subject 10 or 15 times. After each presentation the subject was required to state which of the circles was the larger: the right or the left. This was the experiment to establish a set [ustanovka] in the course of which the subject developed the readiness, or set, to perceive unequal objects, as is established in critical (control) tests. In the critical test, the subjects, to whom circles of equal size are shown, continue to perceive them as unequal, because they act against a pre-developed set. The effect based on an established set lasts for several displays and, what is of particular interest to us in the given instance, the circle which seems larger is not the one on the side where the larger circle had been presented in the experiments to produce persistent images, but that on the opposite side (where the smaller circle had been seen). This was the manner in which the contrast illusion was discovered, subsequently named the "Uznadze effect" by J. Piaget, who in turn proposed a technique for measuring the strength of this illusion. (17)     

Early‐life telomere length predicts lifespan and lifetime reproductive success in a wild bird

Poor conditions during early development can initiate trade‐offs that favour current survival at the expense of somatic maintenance and subsequently, future reproduction. However, the mechanisms that link early and late life‐history are largely unknown. Recently it has been suggested that telomeres, the nucleoprotein structures at the terminal end of chromosomes, could link early‐life conditions to lifespan and fitness. In wild purple‐crowned fairy‐wrens, we combined measurements of nestling telomere length (TL) with detailed life‐history data to investigate whether early‐life TL predicts fitness prospects. Our study differs from previous studies in the completeness of our fitness estimates in a highly philopatric population. The association between TL and survival was age‐dependent with early‐life TL having a positive effect on lifespan only among individuals that survived their first year. Early‐life TL was not associated with the probability or age of gaining a breeding position. Interestingly, early‐life TL was positively related to breeding duration, contribution to population growth and lifetime reproductive success because of their association with lifespan. Thus, early‐life TL, which reflects growth, accumulated early‐life stress and inherited TL, predicted fitness in birds that reached adulthood but not noticeably among fledglings. These findings suggest that a lack of investment in somatic maintenance during development particularly affects late life performance. This study demonstrates that factors in early‐life are related to fitness prospects through lifespan, and suggests that the study of telomeres may provide insight into the underlying physiological mechanisms linking early‐ and late‐life performance and trade‐offs across a lifetime.     

Demography,female life history,and reproductive profiles among the chimpanzees of Mahale

Demography provides critical data to increase our understanding of the evolution, ecology, and conservation of primate populations. The chimpanzees of the Mahale Mountains National Park, Tanzania, have been studied for more than 34 yr on the basis of individual identification and standardized attendance records. From this long-term study, we derived the following demographic data: The major cause of death was disease (48%), followed by senescence (24%) and within-species aggression (16%). Fifty percent of Mahale chimpanzees died before weaning. The median ages of female life history variables were: first maximal swelling, 10.0 yr (n = 5); emigration, 11.0 yr (n = 11); and first birth, 13.1 yr (n = 5). The median period of adolescent infertility was 2.8 yr (n = 4) when calculated from the age at immigration to that at first birth. Female fecundity was highest between 20 and 35 yr of age, with an annual birth rate of 0.2. Twenty-six females that were observed from a young age (10-13 yr) to death at various ages (15-40 yr) gave birth to an average of 3.9 and weaned an average of 1.4 offspring. Twenty-five females that were observed from middle age (18-33 yr) to death in older age (31-48) gave birth to an average of 2.7 and weaned an average of 2.0 offspring. The post-reproductive lifespan for female chimpanzees was defined as the number of years that passed from the year when the last offspring was born to the year when the female died, minus 5. Twenty-five percent of old females had a post-reproductive lifespan. The interbirth interval after the birth of a son (x = 72 mo) tended to be longer than that after the birth of a daughter (x = 66 mo). The extent of female transfer, which is a rule in chimpanzees, is influenced by the size and composition of the unit group and size of the overall local community.     

Prioritizing child health interventions in ethiopia: modeling impact on child mortality, life expectancy and inequality in age at death

Background

The fourth Millennium Development Goal calls for a two-thirds reduction in under-5 mortality between 1990 and 2015. Under-5 mortality rate is declining, but many countries are still far from achieving the goal. Effective child health interventions that could reduce child mortality exist, but national decision-makers lack contextual information for priority setting in their respective resource-constrained settings. We estimate the potential health impact of increasing coverage of 14 selected health interventions on child mortality in Ethiopia (2011–2015). We also explore the impact on life expectancy and inequality in the age of death (Gini_health).

Methods and Findings

We used the Lives Saved Tool to estimate potential impact of scaling-up 14 health interventions in Ethiopia (2011–2015). Interventions are scaled-up to 1) government target levels, 2) 90% coverage and 3) 90% coverage of the five interventions with the highest impact. Under-5 mortality rate, neonatal mortality rate and deaths averted are primary outcome measures. We used modified life tables to estimate impact on life expectancy at birth and inequality in the age of death (Gini_health). Under-5 mortality rate declines from 101.0 in 2011 to 68.8, 42.1 and 56.7 per 1000 live births under these three scenarios. Prioritizing child health would also increase life expectancy at birth from expected 60.5 years in 2015 to 62.5, 64.2 and 63.4 years and reduce inequality in age of death (Gini_health) substantially from 0.24 to 0.21, 0.18 and 0.19.

Conclusions

The Millennium Development Goal for child health is reachable in Ethiopia. Prioritizing child health would also increase total life expectancy at birth and reduce inequality in age of death substantially (Gini_health).     

Life-history strategies associated with local population variability confer regional stability

A widely held ecological tenet is that, at the local scale, populations of K-selected species (i.e. low fecundity, long lifespan and large body size) will be less variable than populations of r-selected species (i.e. high fecundity, short lifespan and small body size). We examined the relationship between long-term population trends and life-history attributes for 185 bird species in the Czech Republic and found that, at regional spatial scales and over moderate temporal scales (100-120 years), K-selected bird species were more likely to show both large increases and decreases in population size than r-selected species. We conclude that life-history attributes commonly associated with variable populations at the local scale, confer stability at the regional scale.     

The redistribution of non-breeding dunlins in response to the post-DDT recovery of falcons

Large-scale changes in predator populations are occurring worldwide due to (re-)introductions, over-exploitation, or recovery after decimation by pesticides and persecution. These widespread changes may affect the distribution of their prey. We studied the continental-scale distributions of non-breeding Calidris alpina pacifica and C. a. hudsonia (Pacific and Atlantic dunlins, respectively), as numbers of their major predators—peregrines (Falco peregrinus) and merlins (F. columbarius; together ‘falcons’)—increased after DDT was banned in 1973. For the period 1975–2010 we compiled the number of dunlins and falcons in each of 244 Christmas Bird Count circles, which cover most of the dunlins' non-breeding ranges. Over the study period, falcons increased by 6.5- (Pacific) and 3.1- (Atlantic) fold, spread to more count circles, and the number of dunlins per falcon fell. The annual total count of the two dunlin sub-species fluctuated strongly and independently. We measured annual aggregation as the expected proportion of a subspecies total found on the same count circle as a randomly selected dunlin. The average aggregation of Pacific dunlins (0.117) was about double that of Atlantic dunlins (0.059), with annual variation largely attributable to changes in a few large count circles. The slope of the aggregative response to year-on-year changes in dunlin numbers was identical on the two coasts. The response to the ongoing falcon increase was positive and significant on the Pacific and slightly negative on the Atlantic. We interpret these results using a version of the ideal free distribution that includes predation danger.     

Maintenance of the 2 microns circle plasmid in populations of Saccharomyces cerevisiae.

The 2 microns circle plasmid is maintained at high frequencies in populations of yeast cells. To find out how the plasmid is maintained, three forces were measured: the selective advantage or disadvantage conferred by 2 microns circles, the rate of generation of [Cir0] cells, and the rate of illegitimate transfer of 2 microns circles from cell to cell. It was found that under the conditions used, 2 microns circles confer a selective disadvantage of about 1%, that [Cir0] cells are generated at the rate of 7.6 x 10(-5) per [Cir+] cell per generation, and that illegitimate transfer of 2 microns circles occurs at a rate less than 10(-7) per recipient cell per generation. The most likely explanation of 2 microns circle maintenance is that the plasmid is sexually transmitted at such a rate that it spreads through populations despite selection against it.