The spatial structure of variation in salamander survival,body condition and morphology in a headwater stream network

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Multi-scale recruitment patterns and effects on local population size of a temperate reef fish

Recruitment of the temperate reef fish Coris julis was studied across the Azores Archipelago (central North Atlantic), over four consecutive recruitment seasons and at three spatial scales: between islands (separated by 100s of km), sites within islands (separated by 10s of km) and transects within sites (separated by 10s of m). At the largest scale ( i.e . between islands) spatial recruitment patterns were highly variable, suggesting the influence of stochastic processes. Recruitment was spatially consistent within islands, even though magnitude was unpredictable between years, indicating that processes at meso-scales are probably more deterministic. Recruits settled randomly at the transect scale, probably reflecting habitat homogeneity. It was proposed that large and island-scale patterns reflect larval availability, driven by physical and biological processes occurring in the plankton. No evidence was found for a density-dependent relationship between newly settled and 2 week settled C. julis nor between cumulative recruitment and young-of-the-year. It appears that adult density is limited by larval supply (pre-settlement regulation) at low recruitment sites, and determined by post-settlement, density-dependent processes at high recruitment sites. This work is one of few to investigate multiple spatial and temporal scales of recruitment for a coastal fish species inhabiting isolated, temperate oceanic islands and hence, provides a novel comparison to the many studies of recruitment on coral reefs and other, more connected systems.     

Influence of habitat heterogeneity on the distribution of larval Pacific lamprey (Lampetra tridentata) at two spatial scales

1. Spatial patterns in channel morphology and substratum composition at small (1–10 metres) and large scales (1–10 kilometres) were analysed to determine the influence of habitat heterogeneity on the distribution and abundance of larval lamprey. 2. We used a nested sampling design and multiple logistic regression to evaluate spatial heterogeneity in the abundance of larval Pacific lamprey, Lampetra tridentata, and habitat in 30 sites (each composed of twelve 1‐m² quadrat samples) distributed throughout a 55‐km section of the Middle Fork John Day River, OR, U.SA. Statistical models predicting the relative abundance of larvae both among sites (large scale) and among samples (small scale) were ranked using Akaike's Information Criterion (AIC) to identify the ‘best approximating’ models from a set of a priori candidate models determined from the literature on larval lamprey habitat associations. 3. Stream habitat variables predicted patterns in larval abundance but played different roles at different spatial scales. The abundance of larvae at large scales was positively associated with water depth and open riparian canopy, whereas patchiness in larval occurrence at small scales was associated with low water velocity, channel‐unit morphology (pool habitats), and the availability of habitat suitable for burrowing. 4. Habitat variables explained variation in larval abundance at large and small scales, but locational factors, such as longitudinal position (river km) and sample location within the channel unit, explained additional variation in the logistic regression model. The results emphasise the need for spatially explicit analysis, both in examining fish habitat relationships and in developing conservation plans for declining fish populations.     

Dispersal patterns of coastal fish: implications for designing networks of marine protected areas

Information about dispersal scales of fish at various life history stages is critical for successful design of networks of marine protected areas, but is lacking for most species and regions. Otolith chemistry provides an opportunity to investigate dispersal patterns at a number of life history stages. Our aim was to assess patterns of larval and post-settlement (i.e. between settlement and recruitment) dispersal at two different spatial scales in a Mediterranean coastal fish (i.e. white sea bream, Diplodus sargus sargus) using otolith chemistry. At a large spatial scale (～200 km) we investigated natal origin of fish and at a smaller scale (～30 km) we assessed "site fidelity" (i.e. post-settlement dispersal until recruitment). Larvae dispersed from three spawning areas, and a single spawning area supplied post-settlers (proxy of larval supply) to sites spread from 100 to 200 km of coastline. Post-settlement dispersal occurred within the scale examined of ～30 km, although about a third of post-settlers were recruits in the same sites where they settled. Connectivity was recorded both from a MPA to unprotected areas and vice versa. The approach adopted in the present study provides some of the first quantitative evidence of dispersal at both larval and post-settlement stages of a key species in Mediterranean rocky reefs. Similar data taken from a number of species are needed to effectively design both single marine protected areas and networks of marine protected areas.     

Universal scaling of species‐abundance distributions across multiple scales

The scale‐dependent species abundance distribution (SAD) is fundamental in ecology, but few spatially explicit models of this pattern have thus far been studied. Here we show spatially explicit neutral model predictions for SADs over a wide range of spatial scales, which appear to match empirical patterns qualitatively. We find that the assumption of a log‐series SAD in the metacommunity made by spatially implicit neutral models can be justified with a spatially explicit model in the large area limit. Furthermore, our model predicts that SADs on multiple scales are characterized by a single, compound parameter that represents the ratio of the survey area to the species’ average biogeographic range (which is in turn set by the speciation rate and the dispersal distance). This intriguing prediction is in line with recent empirical evidence for a universal scaling of the species‐area curve. Hence we hypothesize that empirical SAD patterns will show a similar universal scaling for many different taxa and across multiple spatial scales.     

Scale specific determinants of tree diversity in an old growth temperate forest in China

The major processes generating pattern in plant community composition depend upon the spatial scale and resolution of observation, therefore understanding the role played by spatial scale on species patterns is of major concern. In this study, we investigate how well environmental (topography and soil variables) and spatial variables explain variation in species composition in a 25-ha temperate forest in northeastern China. We used new variation partitioning approaches to discover the spatial scale (using multi-scale spatial PCNM variables) at which environmental heterogeneity and other spatially structured processes influence tree community composition. We also test the effect of changing grain of the study (i.e. quadrat size) on the variation partitioning results. Our results indicate that (1) species composition in the Changbai mixed broadleaf-conifer forest was controlled mainly by spatially structured soil variation at broad scales, while at finer scales most of the explained variation was of a spatial nature, pointing to the importance of biotic processes. (2) These results held at all sampling grains. However, reducing quadrat size progressively reduced both spatially and environmentally explained variance. This probably partly reflects increasing stochasticity in species abundances, and the smaller proportion of quadrats occupied by each species, when quadrat size is reduced. The results suggest that environmental heterogeneity (i.e. niche processes) and other biotic processes such as dispersal work together, but at different spatial scales, to build up diversity patterns.     

Large-scale manipulation of mayfly recruitment affects population size

Recruitment establishes the initial size of populations and may influence subsequent population dynamics. Although strong inference can be made from empirical relationships between recruitment and population sizes, a definitive test of recruitment limitation requires manipulating recruitment at relevant spatial and temporal scales. We manipulated oviposition of the mayfly Baetis bicaudatus in multiple streams and measured the abundance of late-stage larvae at the end of the cohort. Based on fundamental knowledge of mayfly behavior, we increased, eliminated, or left unmodified preferred mayfly oviposition sites in 45-m reaches of streams (N = 4) of one high-altitude drainage basin in western Colorado, USA. We compared egg densities before (2001) and after the manipulation (2002) using paired t tests and compared larval densities before and after the manipulation among treatments using repeated measures analysis of variance. This manipulation altered not only egg densities, but also larval abundances 1 year later. Compared to the previous year, we experimentally increased egg densities at the addition sites by approximately fourfold, reduced egg densities to zero in the subtraction sites, and maintained egg densities in the control sites. After the manipulation, larval densities increased significantly by a factor of approximately 2.0 in the addition sites and decreased by a factor of approximately 2.5 in the subtraction sites. This outcome demonstrates that dramatic changes in recruitment can limit larval population size at the scale of a stream reach, potentially masking previously observed post-recruitment processes explaining the patterns of variation in abundance of a stream insect. Furthermore, our results emphasize the importance of preferred oviposition habitats to population sizes of organisms.     

Scaling effects of grazing in a semi-arid grassland

Abstract. Previous studies have demonstrated relationships between spatial scale and spatial pattern and developed general hypotheses of scaling effects. Few studies, however, have examined the interactive relationship between scale and pattern-driving processes such as grazing. The goal of this study is to evaluate scale-dependent patterns across three spatial scales for three grazing intensities over 45 yr and to identify some mechanisms that may be associated with scale related differences. Correlation analysis and analysis of the coefficients of variation indicate that the relationships between units are dependent upon spatial scale and treatment. Across all grazing treatments, the relationship between units of the same scale becomes stronger as the spatial scale is increased. However, the rate of increase in the correlation coefficient is different for each treatment. The coefficient of variation responded inversely across scales with the greatest variation between small-scale units and little difference between the intermediate- and large scales. In addition to different relationships between units at each scale, differences in heterogeneity within treatments over time is illustrated by the relationship between small-scale units within each treatment and their associated larger scale units. The strongest relationship occurred in the heavily grazed treatments where correlation coefficients of small-scale units with intermediate- and large-scale units were ca. 0.60, indicating similar dynamics across scales. For the moderately grazed and ungrazed treatments this relationship varied from 0.40 to 0.47. Results from this study suggest that grazing alters scaling effects. Variability between small-scale units was greatest in the ungrazed treatment which had greater heterogeneity and less predictability than grazed treatments because of the influence of grazing on plant morphology, demography and composition. At the intermediate scale, relationships between units were fairly similar with the least variation occurring in the moderately grazed treatment. Alternatively, variation between large-scale units was greatest in the moderately grazed treatment because of the relationship between rest cycles, weather patterns, and patch grazing. Therefore, grazing can have a positive, a negative, or no influence on heterogeneity between units depending upon the scale of observation. Evaluation of long-term dynamics across these treatments at the same small spatial scale results in different variances within each treatment which may violate assumptions of some statistical and experimental designs. Therefore, evaluations of temporal dynamics should consider scale relative to the relationship between plant size, density and longevity (relative scale).     

Large‐scale,multidirectional larval connectivity among coral reef fish populations in the Great Barrier Reef Marine Park

Larval dispersal is the key process by which populations of most marine fishes and invertebrates are connected and replenished. Advances in larval tagging and genetics have enhanced our capacity to track larval dispersal, assess scales of population connectivity, and quantify larval exchange among no‐take marine reserves and fished areas. Recent studies have found that reserves can be a significant source of recruits for populations up to 40 km away, but the scale and direction of larval connectivity across larger seascapes remain unknown. Here, we apply genetic parentage analysis to investigate larval dispersal patterns for two exploited coral reef groupers (Plectropomus maculatus and Plectropomus leopardus) within and among three clusters of reefs separated by 60–220 km within the Great Barrier Reef Marine Park, Australia. A total of 69 juvenile P. maculatus and 17 juvenile P. leopardus (representing 6% and 9% of the total juveniles sampled, respectively) were genetically assigned to parent individuals on reefs within the study area. We identified both short‐distance larval dispersal within regions (200 m to 50 km) and long‐distance, multidirectional dispersal of up to ~250 km among regions. Dispersal strength declined significantly with distance, with best‐fit dispersal kernels estimating median dispersal distances of ~110 km for P. maculatus and ~190 km for P. leopardus. Larval exchange among reefs demonstrates that established reserves form a highly connected network and contribute larvae for the replenishment of fished reefs at multiple spatial scales. Our findings highlight the potential for long‐distance dispersal in an important group of reef fishes, and provide further evidence that effectively protected reserves can yield recruitment and sustainability benefits for exploited fish populations.     

The scale-dependent importance of habitat factors and dispersal limitation in structuring Great Lakes shoreline plant communities

Niche-based and neutral models of community structure posit distinct mechanisms underlying patterns in community structure; correlation between species’ distributions and habitat factors points to niche assembly while spatial pattern independent of habitat suggests neutral assembly via dispersal limitation. The challenge is to disentangle the relative contributions when both processes are operating, and to determine the scales at which each is important. We sampled shoreline plant communities on an island in Lake Michigan, varying the extent and the grain of sampling, and used both distance-based correlation methods and variance partitioning to quantify the proportion of the variation in plant species composition that was attributable to habitat factors and to spatial configuration independent of habitat. Our results were highly scale dependent. We found no distance decay of plant community similarity at the island scale (1−33 km). All of the explained variation (32%) in species composition among samples at this scale was attributed to habitat factors. However, at a site intensively sampled at a smaller scale (5−1,200 m), similarity of species composition did decay with distance. Using a coarse sampling grain (transects), habitat factors explained 40% of the variation, but the purely spatial component explained a comparable 22%. Analyzing plots within transects revealed variation in species composition that was still jointly determined by habitat and spatial factors (18 and 11% of the variance, respectively). For both grain sizes, most of the habitat component was spatially structured, reflecting an abrupt alongshore transition from sandy dunes to cobble beach. Space per se explained more variation in species composition at a second site where the habitat transition was more gradual; here, habitat acted as a less selective filter, allowing the signal of dispersal limitation to be detected more readily. We conclude that both adaptation to specific habitat factors and habitat-independent spatial position indicative of dispersal limitation determine plant species composition in this system. Our results support the prediction that dispersal limitation—a potentially, but not necessarily, neutral driver—is relatively more important at smaller scales.     

Small-scale spatial variability in intertidal and subtidal turfing algal assemblages and the temporal generality of these patterns

Spatial and temporal variation in patterns of distribution and abundance of algal assemblages is large and often occurs at extremely small spatial and temporal scales. Despite this, few studies investigate interactions between these scales, that is, how patterns of spatial variation change through time. This study investigated a number of scales of spatial variation (from tens of centimetres to kilometres) in assemblages of intertidal and subtidal turfing algae. Significant differences were found in the composition and abundances of species in assemblages of turf at all spatial scales tested. Much of the variation among assemblages could, however, be explained at the scale of quadrats (tens of centimetres apart) (27±1.4 (SE)% of dissimilarity) with an additional 7±1.2% explained at the scale of sites (tens of metres apart) and 10±1.5% at the scale of locations (kilometres apart). Although the greatest dissimilarity in assemblages occurred at the scale of habitats, this accounted for a relatively small proportion of the overall variation in assemblages. These patterns were consistent through time, that is, at each sampling time the spatial scale explaining the greatest proportion of variation in assemblages was replicate quadrats separated by tens of centimetres. These patterns appear to be due to small-scale variation in patterns of distribution and abundances of the individual species that comprise turfing algal assemblages. The results of this experiment suggest that large scale processes have less effect on patterns of variability of algal assemblages than those occurring on relatively smaller spatial scales and that small-scale spatial variation should not be considered as simply “noise”.     

Dispersal of sea lamprey larvae during early life: relevance for recruitment dynamics

Dispersal is an important early life history process that influences fish population dynamics and recruitment. We studied larval sea lamprey (Petromyzon marinus) dispersal by combining spatially explicit field sampling, genetic methods, and laboratory experiments to investigate how far sea lamprey larvae can disperse away from nests during their first growing season; subsequent dispersal by age 1 of sea lamprey; and the effect of density on larval dispersal. In two study streams sea lamprey larvae were observed to have moved >150 m downstream from the most likely source nest within 2–3 weeks of hatching. Conversely, randomization trials suggested that for both streams age 0 larvae were found closer to full siblings than would be expected if dispersal was not constrained by distance. Restricted dispersal was also observed for age 1 larvae in five streams, although for this age class full siblings were more commonly found to be separated by >1,000 m. Laboratory experiments indicated a significant effect of density on the movement of larval sea lamprey, with more larval movement at higher densities. Temperature also affected movement significantly, with reduced larval movements at cooler temperatures. Our findings suggest that larval sea lamprey dispersal is sufficient to minimize the likelihood of strong density-dependent effects on recruitment, even with large population sizes.     

Spatial heterogeneity in the herbaceous layer of a semi-arid savanna ecosystem

Despite increasing recognition of the role spatial pattern can play in ecosystem function, few studies have quantified spatial heterogeneity in savanna ecosystems. The spatial distribution of herbaceous biomass and species composition was measured across three scales in a semi-arid savanna in central Kenya, and patterns were related to environmental variables at different scales. Herbaceous biomass declined across a rainfall gradient and from upper to lower topographic positions, but variation within a site (across 5–50 m) was similar in magnitude to among-site variation associated with rainfall and topography. Geostatistical analyses showed that patchiness at scales of 5–25 m explained 20% of total variation in herbaceous biomass. This pattern arose from the presence of both 5–10-m diameter patches containing high herbaceous biomass (> 170 g m^–2) and 5–10-m diameter patches characterized by nearly bare soil surfaces (< 40 g m^–2). Patch structure was contingent on topography, with larger bare patches at ridgeline and upper hillslope positions. Grass species distributions showed the greatest degree of patch structure and species turnover across distances of 5–45 m. Additional community variation was associated with topography, with minimal variation in species composition across the rainfall gradient. Pattern diversity significantly exceeded levels reported for four other grassland ecosystems, suggesting fundamental differences in local processes generating spatial pattern. It is hypothesized that heterogeneously distributed grazing pressure, interacting with the distribution of shrub canopies, is an important factor generating such high levels of small-scale patch structure in this savanna.     

Scales of mussel bed complexity: structure, associated biota and recruitment

Hierarchically scaled surveys were carried out on beds of the brown mussel Perna perna (Linnaeus) on the South coast of South Africa. The object was to assess spatial and temporal variations in the complexity of mussel beds and to investigate relationships between mussel bed complexity and mussel recruitment. Complexity was divided into three components: physical complexity; demographic complexity; associated biota. A series of variables within each component were recorded at two different scales (10 and 50 cm) within nested quadrats on three separate occasions. The nested ANOVA design explicitly incorporated spatial scale as levels of the ANOVA. These scales were: shores (areas 1 km in length separated by 25 km); transects (areas 20 m in length separated by 100s of meters); 50x50-cm quadrats separated by meters and 10x10-cm quadrats separated by cm) This approach was intended to generate hypotheses concerning direct associations between recruitment and complexity versus co-variation due external processes. Three main questions were addressed: (1) At what scale does each variable of complexity exhibit greatest significant variation? (2) At these scales is there similar ranking of variables of complexity and recruitment? (3) Within this/these scales, is there any significant relationship between the variables measured and mussel recruitment? On two occasions (Nov. 97 and Mar. 98) the majority of variables showed greatest significant variation at the transect-scale. On a third occasion (Oct. 97) most variables showed greatest significant variation at the quadrat-scale and the site-scale. On all occasions a markedly high percentage of the variation encountered also occurred at the smallest scale of the study, i.e., the residual scale of the ANOVA analyses. Some similarity in the ranking of variables occurred at the transect scale. Within the transect-scale, there was little indication of any relationship between variables of complexity and recruitment. Relationships were inconsistent either among transects or among sampling occasions. Overall, the results suggest that a high degree of variation in mussel bed complexity consistently occurs at very small scales. High components of variance generally also occur at one or more larger scales; however, these scales vary with season. Mussel recruitment does not appear to be directly affected by complexity of mussel beds. Instead it appears external factors may influence both complexity and recruitment independently. In addition recruitment may influence complexity rather than vice versa.     

Long‐term consistency in spatial patterns of primate seed dispersal

Seed dispersal is a key ecological process in tropical forests, with effects on various levels ranging from plant reproductive success to the carbon storage potential of tropical rainforests. On a local and landscape scale, spatial patterns of seed dispersal create the template for the recruitment process and thus influence the population dynamics of plant species. The strength of this influence will depend on the long‐term consistency of spatial patterns of seed dispersal. We examined the long‐term consistency of spatial patterns of seed dispersal with spatially explicit data on seed dispersal by two neotropical primate species, Leontocebus nigrifrons and Saguinus mystax (Callitrichidae), collected during four independent studies between 1994 and 2013. Using distributions of dispersal probability over distances independent of plant species, cumulative dispersal distances, and kernel density estimates, we show that spatial patterns of seed dispersal are highly consistent over time. For a specific plant species, the legume Parkia panurensis, the convergence of cumulative distributions at a distance of 300 m, and the high probability of dispersal within 100 m from source trees coincide with the dimension of the spatial–genetic structure on the embryo/juvenile (300 m) and adult stage (100 m), respectively, of this plant species. Our results are the first demonstration of long‐term consistency of spatial patterns of seed dispersal created by tropical frugivores. Such consistency may translate into idiosyncratic patterns of regeneration.     

An integrated study of the temporal and spatial variation in the supply of propagules, recruitment and assemblages of intertidal macroalgae on a wave-exposed rocky coast, Victoria, Australia

Recruitment is known to influence distributions and abundances of benthic marine organisms. It is therefore important to document patterns of variability in recruitment and how these relate to patterns in established assemblages. This study provides an integrated assessment of the temporal and spatial variation in supply and recruitment of propagules and established populations of several macroalgae. Propagules in water samples from two stages of the incoming tide, recruitment to artificial substrata and percentage cover of species established on the shore were recorded every 2 months from December 1994 to October 1995, in two zones of an intertidal, wave-exposed rocky shore. Variability in recruitment was measured at three spatial scales: 10s cm, 100s cm and 100s m. Availability and recruitment of most taxa were greatest between April and August, although many species had available propagules and recruited throughout the year. Temporal variation in the established assemblages was, however, more species-specific. Differences in established assemblages between zones were reflected in differences in availability and recruitment of propagules between zones. Recruitment could not be predicted directly from supply of propagules, but the two processes were linked. For most species, the greatest variation in recruitment occurred at the smallest spatial scale of 10s cm, although there was also considerable large-scale (between site) variation in recruitment of several species. Results indicate that while pre-and post-settlement mortality are likely to influence macroalgal distribution and abundance, the temporal and spatial variability in supply and recruitment of propagules can explain much of the patchiness in macroalgal assemblages.     

Coexistence patterns of two invasive thistle species, Carduus nutans and C. acanthoides, at three spatial scales

To better understand the competitive processes involved in invasion by congeners, we examine coexistence patterns of two invasive species, Carduus nutans and C. acanthoides, at three spatial scales. A roadside survey of 5 × 5 km blocks in a previously identified overlap zone provided information about the regional scale. At smaller scales, we surveyed four fields of natural co-occurrence, quantifying the spatial patterns at the field scale by randomly placed 1 × 1 m quadrats and at the smallest scale by detailing plant position within the quadrats. The patterns observed are strikingly different at the different scales. At the regional scale, there is positive local autocorrelation in both species but negative cross-correlation between them, consistent with previous surveys. However, at the field scale, there is positive local autocorrelation in both species, and we generally see a positive association between the two species. At the plot scale, when excluding areas of joint absence, there is again a negative association between the two species. This pattern can also be seen at the field scale when excluding plots with joint absence. These results suggest that, at the scale of a field, the strongest factor determining location is aggregation in favorable habitats, which is a stronger force than the competition-induced segregation evidenced at small scales. Lottery competition for spatially aggregated safe sites thus appears to drive the patterns observed at the field scale, while the regional scale pattern may be a result of restricted natural dispersal and invasion history.     

Spatial patterns of species distributions in grazed subalpine grasslands

Spatial patterns of species diversity have important influences on the functioning of ecosystems, and the effect of livestock grazing on spatial heterogeneity can differ depending on the scale of the analysis. This study examined the effects of grazing on the spatial patterns of species distributions and whether the effects of grazing on the spatial distributions of a species vary with the scale of the analysis. Data were collected at three locations in the subalpine grasslands of Ordesa-Monte Perdido National Park and Aísa Valley, Central Pyrenees, Spain, which differed in mean stocking rates. Aspect explained about one-third of the environmental variation in species distributions. In flat areas, spatial variation in species composition varied with grazing intensity at two scales. At a coarse scale (among vegetation patches), grazing promoted patchiness, and among-transect variation in species diversity and grazing intensity were positively correlated. At a fine scale (within vegetation patches), the disruption of the self-organizing processes of the species spatial distributions resulted in a reduction in the long-range spatial autocorrelations of some of the characteristic species and in the homogenization of species spatial distributions. The presence of encroaching Echinospartum horridum had a significant influence on the effect of grazing on south-facing grassland slopes.     

Extrinsically and intrinsically generated spatial patterns of algal abundance in an experimental stream

Spatial heterogeneity in species abundance arises from both extrinsic (largely abiotic) and intrinsic (largely biotic) processes. The relative importance of these two types of processes can vary across ecological systems and across temporal and spatial scales. Numerous empirical studies have explored spatial patterns resulting from extrinsic and intrinsic processes, however the interaction of these two types of processes can result in complex patterns that are difficult to test. We used a unique model system consisting of periphytic algae grown on agar in an experimental stream to manipulate an extrinsic and an intrinsic process. We manipulated an extrinsic process by varying the spatial arrangement of nutrients creating both heterogeneous and homogeneous environments for the algae. We manipulated an intrinsic process by introducing a snail herbivore to the system. The resulting spatial algal patterns showed that both types of processes were important in producing spatial abundance patterns and that the patterns occurred at two distinct spatial scales in our system. At the scale of the imposed nutrient heterogeneity, algae “tracked” the differences in nutrient supply rates. The snail herbivores both reduced and promoted spatial patterns in algal abundance at different spatial scales reflecting their species-specific foraging behavior. An ability to detect differences in algal abundance allowed the snails to reduce the power of patterns at the scale of the imposed nutrient heterogeneity; however below a spatial scale of approximately 30 mm the snails could no longer detect differences in algal abundance and so foraged randomly. At this spatial scale the spatial heterogeneity in algal abundance increased and the resulting algal patterns were relatively spatially fixed through time. We suggest that this relative constancy may arise in part from a detected weak Allee effect in algal growth rates.     

Spatio-temporal heterogeneity in coral recruitment around Moorea, French Polynesia: Implications for population maintenance

The spatio-temporal variability of scleractinian coral recruitment was investigated from December 2000 to December 2003 around Moorea Island (French Polynesia). Nine stations, each with 20 terracotta tiles, were placed on the outer reef slope, at 3 sites (Vaipahu, Tiahura, Haapiti), and at 3 depths (6, 12 and 18 m). The relative contribution of the different families of recruits (Pocilloporidae: 60.4%, Poritidae: 18.8%, Acroporidae: 11.2%) and the very low recruitment rates (maximum: 35 recruits per tile, higher mean densities of 10.8 recruits per tile, average 40.77 recruits m^− 2 year^− 1) recorded at Moorea are similar to recruitment patterns recorded on sub-tropical reefs. Over the duration of the study period, we detected a marked seasonal variability in recruitment rates, with the peak recruitment for all families recorded in December-March periods, corresponding to periods of warmest SSTs. However, recruits of Acroporidae were also relatively abundant in the September-December period in some years, which coincides with the known spawning periods of some Acropora species. Total recruitment rate decreased after the first year of the survey, and was probably the result of the bleaching event that occurred in early 2002, which may have reduced fecundity of some coral populations. A lower proportion of recruits were found on the upper surface of the tiles (14.5%), compared to the lower surfaces (57.1%) and sides (28.4%), which is likely the result of intense grazing by herbivorous fish and urchins. We detected a patchy distribution at the station scale, and a significant variation in recruitment patterns among depths and sites. Pocilloporidae recruited more abundantly at 6 and 18 m, whereas Poritidae were generally more abundant at 12 m depth. In contrast, Acroporidae showed no clear depth pattern during the study period. Recruitment was lowest at the most exposed site (Haapiti), especially for Acroporidae, and probably reflects lower settlement rates and/or higher early post-settlement mortality caused by frequent high swells and their associated strong currents. These distinctive characteristics in recruitment patterns of the 3 dominant families of recruits underline the important role of life history strategies in understanding the spatial patterns, dynamics and maintenance processes of coral populations. The relatively low coral recruitment rates recorded from this study indicate that recovery from severe or frequent perturbations will be slow.