History vs. current demography: explaining the genetic population structure of the common frog (Rana temporaria)

The amount of genetic variability at neutral marker loci is expected to decrease, and the degree of genetic differentiation among populations to increase, as a negative function of effective population size. We assessed the patterns of genetic variability and differentiation at seven microsatellite loci in the common frog (Rana temporaria) in a hierarchical sampling scheme involving three regions (208-885 km apart), three subregions within regions and nine populations (5-20 km apart) within subregions, and related the variability and differentiation estimates to variation in local population size estimates. Genetic variability within local populations decreased significantly with increasing latitude, as well as with decreasing population size and regional site occupancy (proportion of censured localities occupied). The positive relationship between population size and genetic variability estimates was evident also when the effect of latitude (cf. colonization history) was accounted for. Significant genetic differentiation was found at all hierarchical levels, and the degree of population differentiation tended to increase with increasing latitude. Isolation by distance was evident especially at the regional sampling level, and its strength increased significantly towards the north in concordance with decreasing census and marker-based neighbourhood size estimates. These results are in line with the conjecture that the influence of current demographic factors can override the influence of historical factors on species population genetic structure. Further, the observed reductions in genetic variability and increased degree of population differentiation towards the north are in line with theoretical and empirical treatments suggesting that effective population sizes decline towards the periphery of a species' range.     

The influence of periphyton biomass and density on grazing in Physella virgata

We studied how differences in periphyton colonization interval and snail density affected grazing rates in Physella virgata, and whether snails controlled periphyton biomass. Both egestion rates and incorporation rates of ¹⁴C labeled periphyton were estimated in laboratory experiments. Periphyton biomass increased with field colonization interval in all experiments, but did not consistently influence estimates of grazing rate. However, increased periphyton abundance in one of the experiments could still explain higher grazer rates in that year, although larger snail body size is a confounding explanation. Increased snail density also resulted in decreased grazing rates, as observed in earlier studies with this snail species, as well as in studies with other snail grazers. Our results suggest grazing rates and resulting impacts may change seasonally with variation in either periphyton biomass, grazer life-history stage or population density.     

Reid's paradox revisited: the evolution of dispersal kernels during range expansion

Current approaches to modeling range advance assume that the distribution describing dispersal distances in the population (the "dispersal kernel") is a static entity. We argue here that dispersal kernels are in fact highly dynamic during periods of range advance because density effects and spatial assortment by dispersal ability ("spatial selection") drive the evolution of increased dispersal on the expanding front. Using a spatially explicit individual-based model, we demonstrate this effect under a wide variety of population growth rates and dispersal costs. We then test the possibility of an evolved shift in dispersal kernels by measuring dispersal rates in individual cane toads (Bufo marinus) from invasive populations in Australia (historically, toads advanced their range at 10 km/year, but now they achieve >55 km/year in the northern part of their range). Under a common-garden design, we found a steady increase in dispersal tendency with distance from the invasion origin. Dispersal kernels on the invading front were less kurtotic and less skewed than those from origin populations. Thus, toads have increased their rate of range expansion partly through increased dispersal on the expanding front. For accurate long-range forecasts of range advance, we need to take into account the potential for dispersal kernels to be evolutionarily dynamic.     

The effects of genetic drift during range expansion on geographical patterns of variation: a computer simulation of the colonization of Australia by Bufo marinus

A computer simulation is performed of allele frequency changes resulting from genetic drift at eleven loci during the probable course of colonization of Australia by populations of the Giant Toad, Bufo marinus. The history of twelve populations for which allele frequency data are available is modelled. Account is taken of the likely pattern of relationship among the populations, the effective size of the populations (as indicated by the observed variance of allele frequencies) and the probable isolation of the populations from each other following their separation. In all simulations, allele frequencies at some loci show a significant association with latitude while others do not. In six of ten simulations, there is a significant association between degree of variation at a locus and the presence of a latitudinal cline of allele frequencies. There are also indications of this kind of association in simulations of a uni-directional range expansion. These results demonstrate that such associations, which were also observed in the data from the actual populations, can result from genetic drift during a range expansion, and therefore cannot be taken as evidence of the action of natural selection.     

Geographic signatures in species turnover: decoupling colonization and extinction across a latitudinal gradient

High latitude communities have low species richness and are rapidly warming with climate change. Thus, temporal changes in community composition are expected to be greatest at high latitudes. However, at the same time traits such as body size can also change with latitude, potentially offsetting or increasing changes to community composition over time. We tested how zooplankton communities (copepods and cladocerans) have changed over a 25–75 year time span by assessing colonization and extinction rates from lakes across an 1800 km latitudinal gradient, and further tested whether species traits predict rates of community change over time. Lake‐level dissimilarity, measured with Sorenson distance, decreased at higher latitudes. This decrease was due to higher colonization rates of cladocerans in lower latitude lakes and consistent extinction rates across the latitudinal gradient. At the species level, colonization increased with regional occupancy, and tended to be higher for smaller bodied, locally abundant, species. Local extinction rates were negatively correlated with local abundance and regional occupancy, but were not influenced by body size. None of these species‐specific characteristics changed predictably with latitude. Contrary to our expectations, low‐latitude zooplankton communities changed more rapidly than high‐latitude communities by becoming more species rich, not by losing species that were historically present. Moreover, colonization and extinction trends suggest that lakes have become increasingly dominated by species with smaller body sizes and that are already common locally and regionally. Together, these findings indicate that rates of species turnover in freshwater lakes across a latitudinal gradient are not predicted by rates of temperature change, but that turnover is nonetheless resulting in trait‐shifts that favour small, generalist species.     

The extra-limital spread of an invasive species via 'stowaway' dispersal: toad to nowhere?

The mechanisms by which invasive species spread through new areas can influence the spatial scale of their impact. Although previous research has focused on 'natural' dispersal rates following initial introductions, human-aided inadvertent dispersal by 'stowaways' on commercial and domestic transport is thought to be a major contributor to long-distance dispersal. Few data exist to support this assumption. Cane toads Bufo marinus were introduced to north-eastern Australia in 1935, and have since dispersed rapidly through the tropics. Based on information accumulated by community groups in Sydney, 400 km south of the cane toads' current Australian distribution, we document high rates of translocation (at least 50 toads arriving in Sydney per year). Most toads were translocated on commercial truck transport carrying landscaping and building materials from the current range of the cane toads in New South Wales and Queensland, and resulted in highly clumped locations of toad arrival reflecting primary truck transport destinations. Most introductions involved single toads (68 of 102 translocation events), but some introductions involved two to 19 animals. Adults of both sexes were represented equally but juveniles were rarely detected. High rates of translocation of adult toads of both sexes suggest that the eventual distribution of cane toads in Australia may be limited by the animals' bioclimatic tolerances rather than by an inability to reach suitable habitats, even in areas far distant from the toads' current range.     

Exploring tree species colonization potentials using a spatially explicit simulation model: implications for four oaks under climate change

Climate change impacts tree species differentially by exerting unique pressures and altering their suitable habitats. We previously predicted these changes in suitable habitat for current and future climates using a species habitat model (DISTRIB) in the eastern United States. Based on the accuracy of the model, the species assemblages should eventually reflect the new quasi‐equilibrium suitable habitats (~2100) after accounting for the lag in colonization. However, it is an open question if and when these newly suitable habitats will be colonized under current fragmented landscapes and realistic migration rates. To evaluate this, we used a spatially explicit cell‐based model (SHIFT) that estimates colonization potentials under current fragmented habitats and several estimates of historical migration rates at a 1 km resolution. Computation time, which was previously the biggest constraint, was overcome by a novel application of convolution and Fast Fourier Transforms. SHIFT outputs, when intersected with future suitable habitats predicted by DISTRIB, allow assessment of colonization potential under future climates. In this article, we show how our approach can be used to screen multiple tree species for their colonization potentials under climate change. In particular, we use the DISTRIB and SHIFT models in combination to assess if the future dominant forest types in the north will really be dominated by oaks, as modelled via DISTRIB. Even under optimistic scenarios, we conclude that only a small fraction of the suitable habitats of oaks predicted by DISTRIB is likely to be occupied within 100 years, and this will be concentrated in the first 10–20 km from the current boundary. We also show how DISTRIB and SHIFT can be used to evaluate the potential for assisted migration of vulnerable tree species, and discuss the dynamics of colonization at range limits.     

Production of diaspores at the landscape level regulates local colonization: an experiment with a spore‐dispersed moss

Effective dispersal is crucial to species inhabiting transient substrates in order for them to be able to persist in a landscape. Bryophytes, pteridophytes, lichens and fungi all have wind‐dispersed small diaspores and can be efficiently dispersed if their diaspores reach air masses above canopy height. However, empirical data on dispersal over landscape scales are scarce. We investigated how the colonization of an acrocarpous clay‐inhabiting pioneer moss, Discelium nudum, varied between sites that differed in connectivity to potential dispersal sources at spatial scales from 1 to 20 km in a region in northern Sweden. We recorded the colonization on ?25 introduced clay heaps at each of 14 experimental sites some months after the dispersal period. The colonization rate ranged from 0–82% and had a statistically significant relationship with a proxy for potential habitats (amount of clay‐dominated soil) in a buffer of 20 km radius surrounding the experimental sites (and also weakly with the amount of substrate in a 10 km buffer). There were no significant relationships between colonization rate and connectivity at smaller scales (1 and 5 km). We made a rough estimate of the number of spores available for dispersal in a landscape, given the amount of clay‐dominated soil, by recording the number of Discelium nudum colonies in two 25 × 25 km landscapes. The estimated available spore numbers in the different 20 km buffers were of the same order of magnitude as the deposition densities at the experimental sites calculated from the colonization rates. The results suggest that the spores of species with scattered occurrences and small diaspores (25 μm) in open landscapes can be deposited over extensive areas, at rates high enough to drive colonization patterns. This also implies that regional connectivity may be more important than local connectivity for these kinds of species.     

Post‐glacial colonization routes coincide with a life‐history breakpoint along a latitudinal gradient

Although adaptive divergence along environmental gradients has repeatedly been demonstrated, the role of post‐glacial colonization routes in determining phenotypic variation along gradients has received little attention. Here, we used a hierarchical Q_ST–F_ST approach to separate the roles of adaptive and neutral processes in shaping phenotypic variation in moor frog (Rana arvalis) larval life histories along a 1,700 km latitudinal gradient across northern Europe. This species has colonized Scandinavia via two routes with a contact zone in northern Sweden. By using neutral SNP and common garden phenotypic data from 13 populations at two temperatures, we showed that most of the variation along the gradient occurred between the two colonizing lineages. We found little phenotypic divergence within the lineages; however, all phenotypic traits were strongly diverged between the southern and northern colonization routes, with higher growth and development rates and larger body size in the north. The Q_ST estimates between the colonization routes were four times higher than F_ST, indicating a prominent role for natural selection. Q_ST within the colonization routes did not generally differ from F_ST, but we found temperature‐dependent adaptive divergence close to the contact zone. These results indicate that lineage‐specific variation can account for much of the adaptive divergence along a latitudinal gradient.     

Vegetation Colonization in a Restoring Tidal Marsh: A Remote Sensing Approach

Although remote sensing offers the ability to monitor wetland restoration, few have tested automated methods for quantifying vegetation change. We implemented a semiautomated technique using color infrared aerial photography and a common vegetation index, Normalized Difference Vegetation Index (NDVI), to document vegetation colonization in a restoring salt marsh. Change in vegetation over a period of 10 years was analyzed using a postclassification comparison technique where each image year was classified individually into vegetated and nonvegetated areas using NDVI thresholds and then differenced between years to identify areas of vegetation change. Vegetated and nonvegetated areas were identified using this technique, as were areas and time periods of vegetation change. By comparing classified NDVI imagery, we calculated that 90% of our study site was vegetated 10 years after restoration. This study demonstrated that high-resolution remotely sensed data can be analyzed with common geospatial software to monitor change in a rapidly vegetating wetland and that long time frames with yearly image acquisition are needed to quantify plant colonization rates. This method was effective at detecting change in vegetation over time in a variable tidal marsh environment using imagery that had inconsistent specifications and quality across years. Inconsistencies included interannual climate variation, phenology, and presence of algae, as well as differences in pixel size and image brightness. Our findings indicate that remote sensing is useful for postrestoration monitoring of tidal marsh ecosystems.     

Growth variability of juvenile soles Solea solea and Solea senegalensis, and comparison with RNA : DNA ratios in the Tagus estuary, Portugal

Growth variability and condition of juvenile soles Solea solea and Solea senegalensis , were assessed through RNA : DNA estimates and compared to absolute growth rates. Higher mean cohort RNA : DNA ratios were observed for cohort I at the beginning of estuarine occurrence for both species (4·42 and 4·87, for S. solea and S. senegalensis respectively). Despite different estuarine colonization habits, no significant differences were observed between RNA : DNA monthly variation for both sole species within the same year ( P  > 0·05 for 2003 and 2004). Juvenile S. senegalensis showed significant differences between RNA : DNA ratios obtained for the two nursery areas ( P  < 0·001). The decrease of seasonal growth rates with fish age was similar to seasonal variation of mean RNA : DNA values. Thus the RNA : DNA pattern of juvenile S. solea and S. senegalensis reflected growth and estuarine colonization patterns.     

Systematic evaluation of factors controlling Perkinsus marinus transmission dynamics in lower Chesapeake Bay

The transmission of Perkinsus marinus in eastern oysters Crassostrea virginica in relation to water temperature, host oyster mortality, and water-column abundance of anti-P. marinus antibody-labeled cells was systematically examined for 20 mo at a site in the lower York River, Virginia, USA. Uninfected sentinel oysters were naturally exposed to the parasite at 2 wk intervals throughout the course of the study to determine the periodicity and rates of parasite transmission. The timing and magnitude of disease-associated oyster mortalities in a local P. marinus-infected oyster population were estimated by monitoring a captive subset of the local oyster population. Flow cytometric immunodetection methods were employed to estimate the abundance of P. marinus cells in water samples collected 3 times each week. The acquisition of P. marinus infections by na?ve sentinel oysters occurred sporadically at all times of the year; however, the highest incidence of infection occurred during the months of August and September. This window of maximum parasite transmission coincided with the death of infected hosts within the captive local oyster population. Counts of antibody-labeled cells ranged from 10 to 11900 cells l(-1), with the highest abundances in July and August coincident with maximum summer temperatures. A statistically significant relationship between water-column parasite abundance and infection-acquisition rate was not observed; however, highest parasite-transmission rates in both years occurred during periods of elevated water-column abundance of parasite cells. These results support the prevailing model of P. marinus transmission dynamics by which maximum transmission rates are observed during periods of maximum P. marinus-associated host mortality. However, our results also indicate that transmission can occur when host mortality is low or absent, so alternative mortality-independent dissemination mechanisms are likely. The results also suggest that atypically early-summer oyster mortality from Haplosporidium nelsoni infection, at a time when infections of P. marinus are light, has a significant indirect influence on P. marinus transmission dynamics. Elimination of these hosts prior to late-summer P. marinus infection-intensification effectively reduces the overall number of P. marinus cells disseminated.     

Systematics and evolutionary history of heavy metal tolerant Thlaspi caerulescens in Western Europe: evidence from genetic studies based on isozyme analysis

Thlaspi caerulescens is distributed in Europe on metalliferous and not metalliferous soils. Individuals from populations growing on heavy metal contaminated soils are well known as hyperaccumulators of zinc and cadmium. The taxonomical treatment of subspecies of Thlaspi caerulescens is unsettled. We investigated the degree of genetic variation among 28 populations of Thlaspi caerulescens from Europe with isozyme analysis to compare inter- and intrapopulational diversity. British material from heavy metal contaminated environments recognized as Thlaspi sylvestre and T. occitanicum are quite similar to each other on the level of isozyme polymophisms, but they are more closely related to populations from non-contaminated stands from Scandinavia and Middle Europe than to metallophytes distributed in Continental Europe. Our findings indicate that a taxonomical subdivision of T. caerulescens is not possible and, furthermore, heavy metal tolerance might have evolved twice in populations of Thlaspi caerulescens from different areas. The trait of zinc tolerance and hyperaccumulation is frequently found in numerous relatives of Thlaspi caerulescens, and it is suggested that this trait has been established and manifested in populations from metalliferous sites during postglacial colonization. From Scandinavia only non-metallophytes are known. These populations are very similar to each other on the isozyme level. This fits to the hypothesis that Thlaspi caerulescens was introduced to Scandinavia in recent times by human activity. Despite full self-compatibility we estimated varying outcrossing rates up to 0.88 in the metallophytes and 0.658 in the non-metallophytes depending on population size and structure.     

Modeling spatial expansion of invasive alien species: relative contributions of environmental and anthropogenic factors to the spreading of the harlequin ladybird in France

Species distribution models (SDM) have often been used to predict the potential ranges of introduced species and prioritize management strategies. However, this approach assumes equilibrium between occurrences and environmental gradients, an assumption which is violated during the invasion process, where many suitable sites are empty because the species has not yet reached them. Here we considered the invasive ladybird Harmonia axyridis as a case study to show the benefits of using a dynamic colonization–extinction model that does not assume equilibrium. We used a multi‐year occupancy model incorporating environmental, anthropogenic and neighborhood effects, to identify factors that explained spreading variation of this species in France from 2004, when only a few occupied sites were detected, to 2011. We found that anthropogenic factors (urbanization, agriculture, vineyards, and presence/absence of highways) explained more variation in the diffusion process than environmental factors (winter and summer temperatures, wind‐speed, and rainfall). The surface of urbanization was the major anthropogenic factor increasing the probability of colonization. The average summer temperature was the main environmental factor affecting colonization, with a negative effect when high or low. The neighborhood effect revealed that colonization was mostly influenced by contributions coming from a radius of 24 km around the focal cell. The contribution of neighborhood decreases over time, suggesting that H. axyridis is reaching its equilibrium in France. This is confirmed by the small discrepancy observed between the performance of our approach and a SDM approach when predicting a single year occupancy pattern at the end of the study period. Our approach has the advantage of explicitly modelling the state of the biological system during the spatial expansion and identifying colonization constraints. This allows managers to explore the effect of different actions on the system at key moments of the invasion process, hence providing a powerful approach to prioritize management strategies.     

A theoretical study of the invasion of cleared areas by tsetse flies (Diptera: Glossinidae)

Large-scale eradication campaigns against tsetse flies Glossina spp. are giving way to smaller operations aimed at disease and vector containment. There has been little discussion of the effects of these changes in policy. This study estimates the rate at which tsetse re-infect treated areas after the termination of control efforts. Movement is modelled as a diffusion process with a daily root mean square displacement (lambda) of 0.2-1 km-1/2 and population growth as logistic with a growth rate (r) < or =1.5% day-1. Invasion fronts move as the product of lambda and radicalr. For r = 0.75% day-1 a front advances at 2.5 km year-1 for each 100 m increment in lambda. If there are 0.001% survivors in 10% of the treated area, the population recovers to within 1% of the carrying capacity (K) within three years. If the control area is subject to invasion from all sides, a treated block of 10,000 km2 is effectively lost within two years - except at the lowest values of lambda and r. Cleared areas of 100 km2 are lost in a year, as observed in a community-based suppression programme in Kenya. If the treated area is closed to re-invasion, but if there is a block where tsetse survive at 0.0001-0.1% of K, the population recovers within 3-4 years for up to 20 km outside the surviving block. If the surviving flies are more widely spread, re-infection is even more rapid. The deterministic approach used here over-estimates re-invasion rates at low density, but comparisons between control scenarios are still valid. Stochastic modelling would estimate more exactly rates of re-infection at near-zero population densities.     

The historv of introductions of Bufo mariuk (Amphibia: Anura); a natural experiment in evolution

The Central American toad, Bufo marinus , has been extensively introduced throughout the Caribbean and Pacific regions and is now one of the most widespread of terrestrial vertebrates. Details, such as the sources and dates of introductions, the number of individuals introduced and the fates of the introduced populations have been documented and are described. The availability of this historical information makes the introductions of great potential value as a series of evolutionary experiments. They can be used to study the way in which allopatric populations diverge genetically and the effects of population bottlenecks of known size on the genetic characteristics of populations, and to examine theories of rapid speciation caused by genetic revolutions associated with founder events.     

Habitat alteration and the conservation of African primates: case study of Kibale National Park, Uganda

Tropical forests and the animals they support are being threatened by accelerating rates of forest conversion and degradation. In a continually fluctuating sociopolitical world, it is often impossible to protect areas from such conversion until the political environment is suitable to pursue conservation goals, by which time, the forests have often been converted to other uses. This reality suggests a need for inquiry into which primate species can persist after different types of disturbances and how quickly primate communities can recover from disturbance. Here we examine the persistence of primate populations in disturbed habitats by providing a case study of patterns of primate abundance in areas of Kibale National Park (766 km2), Uganda, that have been modified by different types and intensities of human activities, primarily commercial logging and agricultural clearing. Distributional surveys at 24 sites and detailed line-transect censuses at six sites demonstrate that primate populations in Kibale are often high and suggest that patterns of population change associated with disturbance are complex. Analysis of the land use coverage of Kibale reveals that abandoned farms (10.3%) and degraded forest (8.7%) now cover 146 km2. Unfortunately, we do not know what proportion of the farms were established on areas that were forest versus grassland. However, if the areas that are now abandoned farms were all once forested, this means that 79 km2 of forest has been lost. Based on density estimates from nearby sites, this would represent a loss of 52,612 monkeys and 200 chimpanzees. Populations would also have been affected by the degradation of the 66 km2 (8.7%) of forest. These estimates of the potential reductions in the primate populations that could have resulted from forest clearing and degradation illustrate the importance of protecting land. A review of the literature illustrates that the biomass of primates found within Kibale is very high in comparison to other locations and thus illustrates the importance of Kibale to regional conservation.     

A model for range expansion of an introduced species: the common waxbill Estrilda astrild in Portugal

Abstract. The common waxbill Estrilda astrild was first introduced to Portugal from Africa in 1964, and has spread across much of the country and into Spain. We modelled the expansion of the common waxbill on a 20 × 20 km UTM grid in 4‐year periods from 1964 to 1999. The time variation of the square root of the occupied area shows that this expansion process is stabilizing in Portugal, and reasons for this are discussed. Several methods used to model biological expansions are not appropriate for the present case, because little quantitative data are available on the species ecology and because this expansion has been spatially heterogeneous. Instead, colonization on a grid was modelled as a function of several biophysical and spatio‐temporal variables through the fitting of a multivariate autologistic equation. This approach allows examination of the underlying factors affecting the colonization process. In the case of the common waxbill it was associated positively with its occurrence in adjacent cells, and affected negatively by altitude and higher levels of solar radiation.     

RANGE CHARACTERISTICS OF PACIFIC COAST BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE SOUTHERN CALIFORNIA BIGHT

Boat-based photoidentification surveys of bottlenose dolphins (Tursiops truncatus) were conducted from 1982 to 1989 in three discrete coastal study areas within the Southern California Bight: (1) Santa Barbara, California; (2) Orange County, California; (3) Ensenada, Baja California, Mexico. A total of 207 recognizable dolphins were identified in these three “secondary” study areas. These individuals were compared to 404 dolphins identified from 1981 to 1989 in our “primary” study area, San Diego, California, to examine the coastal movement patterns of bottlenose dolphins within the Southern California Bight. A high proportion of dolphins photographed in Santa Barbara (88%), Orange County (92%), and Ensenada (88%) were also photographed in San Diego. Fifty-eight percent (n= 120) of these 207 dolphins exhibited back-and-forth movements between study areas, with no evidence of site fidelity to any particular region. Minimum range estimates were 50 and 470 km. Minimum travel-speed estimates were 11-47 km/d, and all dolphin schools sighted during the study were within 1 km of the shore. These data suggest that bottlenose dolphins within the Southern California Bight are highly mobile within a relatively narrow coastal zone. Home-range dimensions and movement patterns for many vettebrate species are influenced, in part, by variation in food resources. The unique range characteristics documented during this study may reflect the highly dynamic nature of this coastal ecosystem and the associated patchy distribution of food resources available to these bottlenose dolphins.     

Socio-geography of human mobility: a study using longitudinal mobile phone data

A relationship between people's mobility and their social networks is presented based on an analysis of calling and mobility traces for one year of anonymized call detail records of over one million mobile phone users in Portugal. We find that about 80% of places visited are within just 20 km of their nearest (geographical) social ties' locations. This figure rises to 90% at a 'geo-social radius' of 45 km. In terms of their travel scope, people are geographically closer to their weak ties than strong ties. Specifically, they are 15% more likely to be at some distance away from their weak ties than strong ties. The likelihood of being at some distance from social ties increases with the population density, and the rates of increase are higher for shorter geo-social radii. In addition, we find that area population density is indicative of geo-social radius where denser areas imply shorter radii. For example, in urban areas such as Lisbon and Porto, the geo-social radius is approximately 7 km and this increases to approximately 15 km for less densely populated areas such as Parades and Santa Maria da Feira.