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.     

Multi-scale spatial pattern of recruitment in the barnacles Semibalanus cariosus at fishing ports on the Kameda peninsula, southern Hokkaido, Japan

The potential causes of the variable nature of recruitment of marine organisms can be inferred from the scales over which they vary. Sampling for recruits of Semibalanus cariosus on the intertidal concrete tetrapods at 21 fishing ports along the Kameda Peninsula, southern Hokkaido, Japan, was conducted at the end of the recruitment season in 1994 at three spatial scales: at each fishing port, separated by several km; at two sites at each fishing port, separated by several hundred m; and on three blocks at each site, separated by 1–2 m. At all spatial scales, recruitment intensity was independent of adult densities. Recruitment densities significantly varied within all spatial-scales, however, 85.6% of the total variances was estimated to be due to variation among ports. Such km-scale variation of recruitment intensities coincided with the hydrographic pattern of the direction of coastal current.     

Patterns and persistence of larval retention and connectivity in a marine fish metapopulation

Connectivity, the demographic linking of local populations through the dispersal of individuals, is one of the most poorly understood processes in population dynamics, yet has profound implications for conservation and harvest strategies. For marine species with pelagic larvae, direct estimation of connectivity remains logistically challenging and has mostly been limited to single snapshots in time. Here, we document seasonal and interannual patterns of larval dispersal in a metapopulation of the coral reef fish Amphiprion polymnus. A 3‐year record of larval trajectories within and among nine discrete local populations from an area of approximately 35 km was established by determining the natal origin of settled juveniles through DNA parentage analysis. We found that spatial patterns of both self‐recruitment and connectivity were remarkably consistent over time, with a low level of self‐recruitment at the scale of individual sites. Connectivity among sites was common and multidirectional in all years and was not significantly influenced by seasonal variability of predominant surface current directions. However, approximately 75% of the sampled juveniles could not be assigned to parents within the study area, indicating high levels of immigrations from sources outside the study area. The data support predictions that the magnitude and temporal stability of larval connectivity decreases significantly with increasing distance between subpopulations, but increases with the size of subpopulations. Given the considerable effort needed to directly measure larval exchange, the consistent patterns suggest snapshot parentage analyses can provide useful dispersal estimates to inform spatial management decisions.     

Using hierarchical sampling to understand scales of spatial variation in early coral recruitment

Ecological patterns are created by processes acting over multiple spatial and temporal scales. By combining spatially explicit sampling with variance components models, the relative importance of spatial scale to overall variability can be determined. We used a spatially structured experimental design in the Mombasa Marine National Park in Kenya to quantify variation in coral recruitment across four spatial scales (~1–1,000 m) and to generate hypotheses about processes affecting recruitment and potential sources of post-settlement mortality during early life history. For the dominant recruiting corals (Pocillopora spp.), variation in recruitment on surfaces protected from fish grazing was greatest at the largest spatial scale examined (1,000 m). We hypothesize that recruitment on protected surfaces varies mainly with larval delivery due to different lagoonal circulation and water flow between sites. Conversely, variation on surfaces exposed to fishes was greatest at the smallest spatial scale (1 m). We hypothesize that recruitment on exposed surfaces mainly reflects local differences in the scale and intensity of fish grazing, which may obscure larval delivery patterns. Spatial variation in recruitment can affect many ecological processes and factors, including growth, survival to maturity, the distribution of habitat, and variation in species interaction strengths. This study demonstrates how spatially explicit sampling, followed by variance components modeling to partition variance across scales, can help to identify potential drivers of patterns at each relevant scale.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.     

Isolation by resistance across a complex coral reef seascape

A detailed understanding of the genetic structure of populations and an accurate interpretation of processes driving contemporary patterns of gene flow are fundamental to successful spatial conservation management. The field of seascape genetics seeks to incorporate environmental variables and processes into analyses of population genetic data to improve our understanding of forces driving genetic divergence in the marine environment. Information about barriers to gene flow (such as ocean currents) is used to define a resistance surface to predict the spatial genetic structure of populations and explain deviations from the widely applied isolation-by-distance model. The majority of seascape approaches to date have been applied to linear coastal systems or at large spatial scales (more than 250 km), with very few applied to complex systems at regional spatial scales (less than 100 km). Here, we apply a seascape genetics approach to a peripheral population of the broadcast-spawning coral Acropora spicifera across the Houtman Abrolhos Islands, a high-latitude complex coral reef system off the central coast of Western Australia. We coupled population genetic data from a panel of microsatellite DNA markers with a biophysical dispersal model to test whether oceanographic processes could explain patterns of genetic divergence. We identified significant variation in allele frequencies over distances of less than 10 km, with significant differentiation occurring between adjacent sites but not between the most geographically distant ones. Recruitment probabilities between sites based on simulated larval dispersal were projected into a measure of resistance to connectivity that was significantly correlated with patterns of genetic divergence, demonstrating that patterns of spatial genetic structure are a function of restrictions to gene flow imposed by oceanographic currents. This study advances our understanding of the role of larval dispersal on the fine-scale genetic structure of coral populations across a complex island system and applies a methodological framework that can be tailored to suit a variety of marine organisms with a range of life-history characteristics.     

Population dynamics of coral-reef fishes: Controversial concepts and hypotheses

Abstract Knowledge of processes that drive the local population dynamics of coral-reef fishes is important for managing reef fisheries, and for using these species as models for understanding the ecology of demersal marine fishes in general. However, the reef-fish literature is replete with poorly defined concepts and vague hypotheses regarding the issue of population dynamics. Dichotomous arguments, such as whether or not recruitment drives population dynamics, are misdirected because they fail to incorporate several important concepts. First, changes in local population size are driven by four demographic rates (birth, death, immigration and emigration), all of which must be studied to understand population dynamics. Second, all populations that persist do so because at least one of these demographic rates operates in a density-dependent way that is both sufficiently strong and appropriately time-lagged. Therefore, identifying the source(s) of direct density dependence is critical for understanding the limits to variation in population size (i.e. population regulation). Third, regulation does not imply a simple point equilibrium in population size; density dependence in populations of reef fishes is bound to lie within a field of stochastic variation, and thus be difficult to detect. Since its formal origin in 1981, the ‘recruitment limitation’ hypothesis for explaining local population dynamics in reef fishes has undergone ambiguous changes in definition that threaten its usefulness. ‘Recruitment, ‘originally defined as the appearance of newly settled fish on a reef, more recently is often measured months after settlement, thus confounding pre- and post-settlement processes. ‘Limitation, ‘ which originally referred to recruitment being so low as to preclude local populations from reaching densities where resources were limiting, is more recently defined as an absence of any form of density dependence after settlement. The most effective means of testing whether post-settlement mortality is in fact density-independent is to examine patterns of mortality directly, rather than indirectly by interpreting the shape of the relationship between initial recruit density and subsequent adult density within a cohort (the recruit-adult function). Understanding the population dynamics of coral-reef fishes will require a more equitable focus on all four demographic rates, be they density dependent or not, as well as greater attention to identifying sources of density dependence. Such a pluralistic focus necessitates integrated studies of both pre- and post-settlement processes conducted at multiple spatial and temporal scales. For example, recent evidence suggests that density-dependent pre-dation on new recruits that have settled among reefs at different densities may prove to be an important source of local population regulation, especially via the aggregative response of transient piscivores.     

Recruitment limitation: A theoretical perspective

Abstract A theoretical analysis of the concept of recruitment limitation leads to the conclusion that most populations should he regarded as jointly limited by recruitment and interactions between individuals after recruitment. The open nature of local marine systems does not permit avoidance of density-dependent interactions; it simply may make them more difficult to detect. Local populations consisting of settled organisms may not experience density-dependent interactions under some circumstances, but the entire species population consisting of the collection of local populations and their planktonic larvae must have density-dependent dynamics. Any local population of settled individuals can escape density dependence if sufficient density dependence occurs among planktonic larvae or within other local populations. Common conceptions of density dependence are too narrow, leading too often to the conclusion that it is absent from a system. It is equally wrong to expect that density-dependent interactions after settlement determine local population densities independently of recruitment. Special circumstances allowing density dependence to act strongly and quickly are needed before density dependence can neutralize the effects of recruitment. Recruitment limitation and density-dependent interactions therefore should not be regarded as alternatives but as jointly acting to determine the densities of marine benthic populations. Moreover, the interaction between fluctuating recruitment and density dependence is potentially the most interesting feature of recruitment limitation. For example, this interaction may be an important diversity-maintaining mechanism for marine systems.     

The role of recruitment in structuring patterns of small-scale spatial variability in intertidal and subtidal algal turfs

Recruitment is often important in structuring patterns of distribution and abundance of algal assemblages. Intertidal and subtidal turfing algal assemblages consistently vary on small spatial scales (tens of centimetres), and this variability may be due to patterns of recruitment varying on similar spatial scales. The validity of this model was evaluated by testing the hypothesis that the numbers and types of taxa recruiting to turfs would vary at small spatial scales within intertidal and within subtidal habitats. Abundances of algal recruits were estimated on sandstone plates that were placed at a number of spatial scales within intertidal and within subtidal habitats (centimetres to tens of metres). Significant differences in entire assemblages were found only between habitats. This was explained by abundances of individual taxa, which generally varied between intertidal and subtidal habitats or between sites within habitats. Only small proportions of the overall spatial variation (dissimilarity) could be explained at the scale of replicate recruitment plates that were centimetres apart. Results indicate that while recruitment may contribute to differences between intertidal and subtidal habitats, it cannot explain the small-scale spatial variability in established turfing algal assemblages within these habitats. There was some evidence to suggest that recruitment may contribute to variability in established turfing algal assemblages but only over longer time scales than examined here.     

Rapid Effects of Marine Reserves via Larval Dispersal

Marine reserves have been advocated worldwide as conservation and fishery management tools. It is argued that they can protect ecosystems and also benefit fisheries via density-dependent spillover of adults and enhanced larval dispersal into fishing areas. However, while evidence has shown that marine reserves can meet conservation targets, their effects on fisheries are less understood. In particular, the basic question of if and over what temporal and spatial scales reserves can benefit fished populations via larval dispersal remains unanswered. We tested predictions of a larval transport model for a marine reserve network in the Gulf of California, Mexico, via field oceanography and repeated density counts of recently settled juvenile commercial mollusks before and after reserve establishment. We show that local retention of larvae within a reserve network can take place with enhanced, but spatially-explicit, recruitment to local fisheries. Enhancement occurred rapidly (2 yrs), with up to a three-fold increase in density of juveniles found in fished areas at the downstream edge of the reserve network, but other fishing areas within the network were unaffected. These findings were consistent with our model predictions. Our findings underscore the potential benefits of protecting larval sources and show that enhancement in recruitment can be manifested rapidly. However, benefits can be markedly variable within a local seascape. Hence, effects of marine reserve networks, positive or negative, may be overlooked when only focusing on overall responses and not considering finer spatially-explicit responses within a reserve network and its adjacent fishing grounds. Our results therefore call for future research on marine reserves that addresses this variability in order to help frame appropriate scenarios for the spatial management scales of interest.     

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.     

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.     

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.     

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 surf zone: a semi-permeable barrier to onshore recruitment of invertebrate larvae?

The supply of larvae to the shore is important for population replenishment and intertidal community dynamics but its variability at most scales is not well understood. We tested the relationship between nearshore mussel larval abundance and intertidal settlement rates over several years at multiple spatiotemporal scales in Oregon and New Zealand. Abundance of competent larvae nearshore and intertidal recruitment rates were simultaneously quantified using collectors mounted at different depths on moorings 50-1100 m from shore, and at adjacent rocky intertidal sites. Total mussel larval abundance and oceanographic conditions were also measured in some locations. At all scales, abundance of nearshore mussel larvae was unrelated to intertidal recruitment rates. In the intertidal, patterns of mussel recruitment were persistent in space, with sites of consistently high or low recruitment. In contrast, nearshore competent larval abundance showed generally similar abundances among sites except for a high, and spatially-inconsistent, variability in Oregon during 1998 only. On moorings, recruitment tended to be greater on midwater collectors than shallower or deeper. Finally, on moorings larval abundance in traps and recruitment on collectors was unrelated. These results suggest that (1) among sites, the size of the nearshore larval pool is relatively uniform while onshore recruitment varies and is unrelated to larval abundance, (2) temporal variability in nearshore larval availability is not strongly expressed onshore, (3) vertical stratification of competent larvae nearshore is strong and may influence transport and recruitment, and (4) within-coast variability in onshore recruitment is strongly driven by processes occurring locally within the surf zone that need to be studied to understand coastal recruitment dynamics.     

Predicting reef fish connectivity from biogeographic patterns and larval dispersal modelling to inform the development of marine reserve networks

Developing networks of no-take marine reserves is often hindered by uncertainty in the extent to which local marine populations are connected to one another through larval dispersal and recruitment (connectivity). While patterns of connectivity can be predicted by larval dispersal models and validated by empirical methods, biogeographic approaches have rarely been used to investigate connectivity at spatial scales relevant to reserve networks (10's–100's of km). Here, species assemblage patterns in coral reef fish were used together with an individual-based model of dispersal of reef fish larvae to infer patterns of connectivity in a ∼300 km wide region in the Philippines that included the Bohol Sea and adjacent bodies of water. A dominant current flows through the study region, which may facilitate connectivity among >100 no-take reserves. Connectivity was first investigated by analysing data on the presence/absence of 216 species of reef fish and habitat variables across 61 sites. Hierarchical clustering of sites reflecting species assemblage patterns distinguished a major group of sites in the Bohol Sea (Bray–Curtis similarity >70%) from sites situated in adjacent bodies of water (bays, channels between islands and a local sea). The grouping of sites could be partly explained by a combination of degree of embayment, % cover of sand and % cover of rubble (Spearman rank correlation, ρ_w = 0.42). The individual-based model simulated dispersal of reef fish larvae monthly for three consecutive years in the region. The results of simulations, using a range of pelagic larval durations (15–45 days), were consistent with the species assemblage patterns. Sites in the model that showed strongest potential connectivity corresponded to the majority of sites that comprised the Bohol Sea group suggested by hierarchical clustering. Most sites in the model that exhibited weak connectivity were groups of sites which had fish assemblages that were least similar to those in the Bohol Sea group. Concurrent findings from the two approaches suggest a strong influence of local oceanography and geography on broad spatial patterns of connectivity. The predictions can be used as an initial basis to organise existing reserves to form ecologically meaningful networks. This study showed that species assemblage patterns could be a viable supplementary indicator of connectivity if used together with predictions from a larval dispersal model and if the potential effect of habitat on the structuring of species assemblages is taken into consideration.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.