Patchy Configuration Research Articles

Are Ecosystem Engineering Traits Fixed or Flexible: A Study on Clonal Expansion Strategies in Co-occurring Dune Grasses

Many vegetated coastal ecosystems are formed through ecosystem engineering by clonal vegetation. Recent work highlights that the spatial shoot organization of the vegetation determines local sediment accretion and subsequently emerging landscape morphology. While this key engineering trait has been found to differ between species and prevailing environmental conditions, it remains unknown how the interplay of both factors drive shoot organization and therefore landscape morphology. Here, we compared the spatial shoot organization of young, clonally expanding plants of the two dominant European dune grass species: sand couch (Elytrigia juncea) and marram grass (Ammophila arenaria) across a range of coastal dune environments (from Denmark to France). Our results reveal that, on average, sand couch deployed a more dispersed shoot organization than marram grass, which has a patchy (Lévy-like) organization. Whereas sand couch exhibited the same expansion strategy independent of environmental conditions, marram grass demonstrated a large intraspecific variation which correlated to soil organic matter, temperature and grain size. Shoot patterns ranged from a clumped organization correlating to relatively high soil organic matter contents, temperature and small grain sizes, to a patchy configuration with intermediate conditions, and a dispersed organization with low soil organic matter, temperature and large grain size. We conclude that marram grass is flexible in adjusting its engineering capacity in response to environmental conditions, while sand couch instead follows a fixed expansion strategy, illustrating that shoot organization results from the interaction of both species-specific and environmental-specific trait expression.

Flow-divergence feedbacks control propagule retention by in-stream vegetation: the importance of spatial patterns for facilitation

Facilitation (enhancement of propagule retention in this case) is increasingly recognized as an important driver of biodiversity, but it is still unknown if facilitation during dispersal and colonization is affected by self-organized spatial pattern formation. We investigated the ability of in-stream submerged macrophyte patches to trap the vegetative propagules of three species (Berula erecta, Groenlandia densa, Elodea nuttallii in two size classes: 13–22 and 40–48 cm long), and to potentially benefit the colonization of these three species. We tested the effects of propagule traits, hydrodynamic forcing, and spatial patch configuration on propagule trapping. Propagule buoyancy was negatively correlated with trapping chance, while propagule size did not influence trapping. Species-specific differences in buoyancy were maintained for weeks after fragmentation. Propagule retention was interactive and conditional upon the interplay between incoming flow velocities and vegetation spatial patterning. In the flume experiment at low flows, a patchy configuration (one patch filling 66% of the flume width) retained more surface-drifting propagules (B. erecta, G. densa), than near-homogeneous cover (two patches close together, filling the entire flume width). In contrast, retention of sinking E. nuttallii propagules increased in the two-patch configurations. In flume and field releases where patches did not completely fill the channel width, water flowed around the patches rather than over or through them. This resulted in low-flow velocity areas within patches where canopies were upright and propagules were retained, and higher velocity flows around patches. In contrast, when vegetation filled the channel width, water could not be diverted laterally around the patches and preferentially flowed over them, causing the canopies to bend and reduce their trapping capacity. In flume experiments at high flows, retention of all species decreased, regardless of vegetation configuration, as propagules passed over the reconfigured vegetation canopies. These findings on the interplay of water movement and patch reconfiguration suggest that environmental heterogeneity generated by the self-organizing behavior of aquatic plants might enhance colonization of sessile organisms, calling for landscape-scale processes like dispersal to be better investigated.

General spatial spectral variation in rocky intertidal communities from three biogeographical regions

Aim Understanding general spatial patterns of multiple species is a central aim of biogeography. Spatial analyses of coastal marine assemblages have shown an emergent pattern in that the variability in population abundances increases with increasing spatial scale of observation; i.e. a “red” spectrum of variation in population abundances. However, the generalization of these patterns to other ecological levels and across biogeographical regions remains unexplored. Here, we evaluate the consistency of a reddened relationship between the scale of observation and the variance in ecosystem properties in sessile intertidal rocky-shore assemblages across three biogeographical regions. Location Replicate sites located in temperate (southern-central Chile), sub-Antarctic (Chilean South Patagonia) and Antarctic (West Antarctic Peninsula) regions. Methods A total of 16,225 photo-quadrants were used to analyse local patterns of total community cover, bare substratum, total algal cover and total invertebrate cover. We used spectral analyses in which the spatial variance of each property was decomposed along a gradient of increasing distances between observations. Cross-semivariograms were used to analyse the spatial covariation of dominant taxa. Results The analysed ecosystem properties showed a general reddened spectrum of variation that resembled patchy configurations of local communities. Additionally, cross-semivariance analyses indicated mainly negative relationships of covariation between the abundances dominant sessile species at each site, suggesting a general trend of spatial asynchrony between competing species. Main conclusions To the best of our knowledge, this is the first study showing a general reddened pattern of spatial variation in relevant ecosystem-level properties across the Pacific shore of South America, southern Patagonia and West Antarctic Peninsula. These patterns reflect the result of multiple biotic and abiotic processes that interact across spatial scales. In particular, we suggest that processes that generate spatial asynchrony between competing species, such as compensatory dynamics, are relevant to maintain the spatial structure of rocky-shore communities.

Crustacean decapod assemblages associated with fragmented Posidonia oceanica meadows in the Alboran Sea (Western Mediterranean Sea): composition, temporal dynamics and influence of meadow structure

AbstractThe decapod assemblage associated with a Posidonia oceanica meadow located near its western limit of biogeographic distribution was studied over an annual cycle. Fauna samples were taken seasonally over a year (five replicates per season) in two sites located 7 km apart, using a non‐destructive sampling method (airlift sampler) for the seagrass. The dominant species of the assemblage, Pisidia longimana, Pilumnus hirtellus and Athanas nitescens, were associated with the protective rhizome stratum, which is mainly used as a nursery. The correlations between decapod assemblage structure and some phenological parameters of the seagrass shoots and wave height were negative or null, which reflects that species associated with the rhizome had a higher importance than those associated with the leaf stratum. The abundance and composition of the decapod assemblage as well as the ecological indexes displayed a seasonality trend with maximum values in summer‐autumn and minimum in winter‐spring, which were related to the seawater temperature and the recruitment periods of the dominant species. The spatial differences found in the structure and dynamics of the assemblages may be due to variations in the recruitment of the dominant species, probably as a result of the influence of local factors (e.g. temperature, currents) and the high dispersal ability of decapods, together with the patchy configuration and the surrounding habitats. The studied meadows are fragmented and are integrated within a mosaic of habitats (Cymodocea nodosa patches, algal meadows, rocky and sandy bottoms), which promotes the movement of individuals and species among them, maintaining a high species richness and evenness.

Estimating landscape resistance to dispersal

Dispersal is an inherently spatial process that can be affected by habitat conditions in sites encountered by dispersers. Understanding landscape resistance to dispersal is important in connectivity studies and reserve design, but most existing methods use resistance functions with cost parameters that are subjectively chosen by the investigator. We develop an analytic approach allowing for direct estimation of resistance parameters that folds least cost path methods typically used in simulation approaches into a formal statistical model of dispersal distributions. The core of our model is a frequency distribution of dispersal distances expressed as least cost distance rather than Euclidean distance, and which includes terms for feature-specific costs to dispersal and sex (or other traits) of the disperser. The model requires only origin and settlement locations for multiple individuals, such as might be obtained from mark–recapture studies or parentage analyses, and maps of the relevant habitat features. To evaluate whether the model can estimate parameters correctly, we fit our model to data from simulated dispersers in three kinds of landscapes (in which resistance of environmental variables was categorical, continuous with a patchy configuration, or continuous in a trend pattern). We found maximum likelihood estimators of resistance and individual trait parameters to be approximately unbiased with moderate sample sizes. We applied the model to a small grizzly bear dataset to demonstrate how this approach could be used when the primary interest is in the prediction of costs and found that estimates were consistent with expectations based on bear ecology. Our method has important practical applications for testing hypotheses about dispersal ecology and can be used to inform connectivity planning efforts, via the resistance estimates and confidence intervals, which can be used to create a data-driven resistance surface.

Lack of sex-biased dispersal promotes fine-scale genetic structure in alpine ungulates

Identifying patterns of fine-scale genetic structure in natural populations can advance understanding of critical ecological processes such as dispersal and gene flow across heterogeneous landscapes. Alpine ungulates generally exhibit high levels of genetic structure due to female philopatry and patchy configuration of mountain habitats. We assessed the spatial scale of genetic structure and the amount of gene flow in 301 Dall’s sheep (Ovis dalli dalli) at the landscape level using 15 nuclear microsatellites and 473 base pairs of the mitochondrial (mtDNA) control region. Dall’s sheep exhibited significant genetic structure within contiguous mountain ranges, but mtDNA structure occurred at a broader geographic scale than nuclear DNA within the study area, and mtDNA structure for other North American mountain sheep populations. No evidence of male-mediated gene flow or greater philopatry of females was observed; there was little difference between markers with different modes of inheritance (pairwise nuclear DNA F ST = 0.004–0.325; mtDNA F ST = 0.009–0.544), and males were no more likely than females to be recent immigrants. Historical patterns based on mtDNA indicate separate northern and southern lineages and a pattern of expansion following regional glacial retreat. Boundaries of genetic clusters aligned geographically with prominent mountain ranges, icefields, and major river valleys based on Bayesian and hierarchical modeling of microsatellite and mtDNA data. Our results suggest that fine-scale genetic structure in Dall’s sheep is influenced by limited dispersal, and structure may be weaker in populations occurring near ancestral levels of density and distribution in continuous habitats compared to other alpine ungulates that have experienced declines and marked habitat fragmentation.

Patchiness and Demographic Noise in Three Ecological Examples

Understanding the causes and effects of spatial aggregation is one of the most fundamental problems in ecology. Aggregation is an emergent phenomenon arising from the interactions between the individuals of the population, able to sense only—at most—local densities of their cohorts. Thus, taking into account the individual-level interactions and fluctuations is essential to reach a correct description of the population. Classic deterministic equations are suitable to describe some aspects of the population, but leave out features related to the stochasticity inherent to the discreteness of the individuals. Stochastic equations for the population do account for these fluctuation-generated effects by means of demographic noise terms but, owing to their complexity, they can be difficult (or, at times, impossible) to deal with. Even when they can be written in a simple form, they are still difficult to numerically integrate due to the presence of the “square-root” intrinsic noise. In this paper, we discuss a simple way to add the effect of demographic stochasticity to three classic, deterministic ecological examples where aggregation plays an important role. We study the resulting equations using a recently-introduced integration scheme especially devised to integrate numerically stochastic equations with demographic noise. Aimed at scrutinizing the ability of these stochastic examples to show aggregation, we find that the three systems not only show patchy configurations, but also undergo a phase transition belonging to the directed percolation universality class.

Tropical metacommunities along elevational gradients: effects of forest type and other environmental factors

Elevational gradients provide a natural experiment for assessing the extent to which the structure of animal metacommunities is molded by biotic and abiotic characteristics that change gradually, or is molded by aspects of plant community composition and physiognomy that change in a more discrete fashion. We used a metacommunity framework to integrate species‐specific responses to environmental gradients as an approach to detect emergent patterns at the mesoscale in the Luquillo Mountains of Puerto Rico. Elements of metacommunity structure (coherence, species turnover and range boundary clumping) formed the basis for distinguishing among random, checkerboard, Gleasonian, Clementsian, evenly spaced and nested patterns. Paired elevational transects (300–1000 m a.s.l.) were sampled at 50 m intervals to decouple underlying environmental mechanisms: a mixed forest transect reflected changes in abiotic and biotic conditions, including forest type (i.e. tabonuco, palo colorado and elfin forests), whereas another transect reflected changes in environmental conditions but not forest type, as its constituent plots were located within palm forest. Based on distributional data (presence versus absence of species), the mixed forest transect exhibited Clementsian structure, whereas the palm forest transect exhibited quasi‐Gleasonian structure. In contrast, the distribution of modes in species abundance was random with respect to the latent environmental gradient in the mixed forest transect and clumped with respect to the latent environmental gradient in the palm forest transect. Such contrasts suggest that the environmental factors affecting abundance differed in form or type from those affecting distributional boundaries. Variation among elevational strata with respect to the first axis of correspondence from reciprocal averaging was highly correlated with elevation along each transect, even though axis scores were not correlated between mixed forest and palm forest transects. This suggests that the identity of the environmental characteristics, or the form of response by the fauna to those characteristics, differed between the two elevational transects. Despite the proximity of the transects, the patchy configuration of palm forest, and the pervasive distribution of the dominant palm species, the relative importance of abiotic variables and habitat in structuring gastropod metacommunities differed between transects, which is remarkable and attests to the sensitivity of metacommunity structure to environmental variation.

Resource distribution influences positive edge effects in a seagrass fish

According to conceptual models, the distribution of resources plays a critical role in determining how organisms distribute themselves near habitat edges. These models are frequently used to achieve a mechanistic understanding of edge effects, but because they are based predominantly on correlative studies, there is need for a demonstration of causality, which is best done through experimentation. Using artificial seagrass habitat as an experimental system, we determined a likely mechanism underpinning edge effects in a seagrass fish. To test for edge effects, we measured fish abundance at edges (0-0.5 m) and interiors (0.5-1 m) of two patch configurations: continuous (single, continuous 9-m2 patches) and patchy (four discrete 1-m2 patches within a 9-m2 area). In continuous configurations, pipefish (Stigmatopora argus) were three times more abundant at edges than interiors (positive edge effect), but in patchy configurations there was no difference. The lack of edge effect in patchy configurations might be because patchy seagrass consisted entirely of edge habitat. We then used two approaches to test whether observed edge effects in continuous configurations were caused by increased availability of food at edges. First, we estimated the abundance of the major prey of pipefish, small crustaceans, across continuous seagrass configurations. Crustacean abundances were highest at seagrass edges, where they were 16% greater than in patch interiors. Second, we supplemented interiors of continuous treatment patches with live crustaceans, while control patches were supplemented with seawater. After five hours of supplementation, numbers of pipefish were similar between edges and interiors of treatment patches, while the strong edge effects were maintained in controls. This indicated that fish were moving from patch edges to interiors in response to food supplementation. These approaches strongly suggest that a numerically dominant fish species is more abundant at seagrass edges due to greater food availability, and provide experimental support for the resource distribution model as an explanation for edge effects.

RESOURCE DISTRIBUTION INFLUENCES POSITIVE EDGE EFFECTS IN A SEAGRASS FISH

According to conceptual models, the distribution of resources plays a critical role in determining how organisms distribute themselves near habitat edges. These models are frequently used to achieve a mechanistic understanding of edge effects, but because they are based predominantly on correlative studies, there is need for a demonstration of causality, which is best done through experimentation. Using artificial seagrass habitat as an experimental system, we determined a likely mechanism underpinning edge effects in a seagrass fish. To test for edge effects, we measured fish abundance at edges (0–0.5 m) and interiors (0.5–1 m) of two patch configurations: continuous (single, continuous 9-m2 patches) and patchy (four discrete 1-m2 patches within a 9-m2 area). In continuous configurations, pipefish (Stigmatopora argus) were three times more abundant at edges than interiors (positive edge effect), but in patchy configurations there was no difference. The lack of edge effect in patchy configurations might be because patchy seagrass consisted entirely of edge habitat. We then used two approaches to test whether observed edge effects in continuous configurations were caused by increased availability of food at edges. First, we estimated the abundance of the major prey of pipefish, small crustaceans, across continuous seagrass configurations. Crustacean abundances were highest at seagrass edges, where they were 16% greater than in patch interiors. Second, we supplemented interiors of continuous treatment patches with live crustaceans, while control patches were supplemented with seawater. After five hours of supplementation, numbers of pipefish were similar between edges and interiors of treatment patches, while the strong edge effects were maintained in controls. This indicated that fish were moving from patch edges to interiors in response to food supplementation. These approaches strongly suggest that a numerically dominant fish species is more abundant at seagrass edges due to greater food availability, and provide experimental support for the resource distribution model as an explanation for edge effects.

Genetic survey of two endangered grasshopper subspecies, Prionotropis hystrix rhodanica and Prionotropis hystrix azami (Orthoptera, Pamphagidae): within- and between-population dynamics at the regional scale

The fragmentation and destruction of natural habitats is one of the major causes of extinction of wild flora and fauna due to human activities. The area around the Mediterranean Basin is particularly affected: this region is one of the world’s richest in terms of biodiversity and is undergoing substantial modification due to extensive changes in land use. We investigated the demographic state (through a genetic survey) of an endangered grasshopper, the geographical distribution of which is restricted to arid open land or glades in south-eastern France. We compared two subspecies: one is severely threatened by the fragmentation of its specific and restricted habitat (Prionotropis hystrix rhodanica in the ‘Crau’ steppe) whereas the other, P.␣h.␣azami, occurs more widely in glades or clearings in the evergreen Mediterranean forest. Genetic diversity at eight microsatellite loci was highly structured, indicating substantial isolation of populations. Within-population diversity was high at all but one site. Genetic drift was the major force involved in this genetic structure, with very little gene flow at the regional scale of observation, consistent with both the limited dispersal of this flightless species and the patchy configuration of its habitat. No significant differences in the extent of genetic diversity or population dynamics were observed between the two subspecies. The observed genetic pattern calls into question the validity of the subspecies status, and shows no strong impact of recent anthropogenic modification of the Crau steppe. The evolution of the Mediterranean xeric and open habitats under both natural and anthropogenic processes is discussed.

Spatial Distribution of Net Diatoms Along Adjacent Water Masses of Different Origin

Larger diatoms (>55 µm) were sampled using closing bottles from on board the R/V Bilim along adjacent water masses of different origin in order to observe patchy assemblages of net diatoms. MDS (Multi-Dimensional Scaling) plots of diatoms in adjacent water masses inhabiting the south-western Black Sea, Bosphorus, Golden Horn, the north-eastern Sea of Marmara and Izmit Bay have shown that patchy configurations of diatom were orientated along particular dominating environmental gradients, mainly temperature and salinity. A two-way evolution takes place in the Bosphorus as the Black Sea water flows towards the Sea of Marmara near the surface and under-neath the Mediterranean inflow, below the pycnocline towards the Black Sea. Considerable changes in salinity and temperature with depth in the region affected the drifting flora considerably, especially its diatom composition. Of all the regions, the south-western Black Sea was the most species diverse and abundant.

The patchy configuration of scaphoid avascular necrosis

The proximal poles of five patients with presumed avascular necrosis of the scaphoid, mostly the result of nonunion, were analyzed histologically for true, pathologically confirmed avascular necrosis of bone. Each specimen was reviewed both qualitatively by an experienced bone pathologist and quantitatively with the use of a computerized image analysis system. Avascular necrosis was found to be variable from one specimen to another as well as within each specimen. Midcentral levels showed a statistically significant greater degree of avascular necrosis than either adjacent palmar or dorsal zones. Both viable and nonviable areas of avascular bone were found in an unpredictable, irregular (patchy) distribution, often in direct apposition to one another. It is apparent from this study that random biopsy alone cannot accurately predict the histologic status of the entire specimen; the clinician must rely on other methods to determine scaphoid viability.

Variance of sightings in the survey of patchily distributed objects

In the field, animals, birds and fishes are often distributed inhomogeneously. In such situations, the variance of sightings of strip or line transect survey would be larger than when they are independently distributed one another. We formulate in this paper the structure of the patchy configuration, and deduce the variance of sightings of strip or line transect survey. From this, we see that it is larger when the patch size (the expected number of objects in each patch) is larger and the patch radius is smaller.

Ocular findings in hemodialysis and following kidney transplantation in childhood and adolescence

Between August 1980 and January 1987, 23 patients undergoing treatment for chronic renal failure underwent eye examinations. Hemodialysis and subsequent kidney transplants were performed in 18 patients; in two patients a kidney transplant was performed alone, and in three others hemodialysis without transplant. The interval between dialysis and transplantation averaged 23.1 months, the mean follow-up after transplantation 20 months. Patients who underwent hemodialysis alone were followed up for periods of two, three and 85 months. The patients' ages when hemodialysis treatment was first instituted ranged from six to 17 years (average 11.8 years). The mean age at the time kidney transplants were performed was 13.6 years (ranging from one to 17 years). Seventeen patients had conjunctival and corneal infiltrations in the area of the palpebral fissure. In two cases infiltrations were confined to the conjunctiva. Four patients had no pathologic changes, in either the cornea or the conjunctiva. Slitlamp examination revealed subcapsular losses of lens transparency in eight patients; these losses were manifested by delicate punctiform and patchy configurations. In nine cases fundus ophthalmoscopy revealed constricted retinal arteries. Within the period of observation all but one of the patients had unchanged vision. The one exception (cystinosis) had reduced visual acuity due to an accumulation of crystalline inclusions in the cornea.