Place-based management at different spatial scales

Place-based management is any management action having implications for a specified area. Place-based management is seen as a key component to practical implementation of ecosystem approach to management, with marine spatial planning (MSP) being the currently most promoted approach. In the present paper we address the challenges of place-based management at local, regional and global (oceanic) spatial scales using case studies from the Northeast Atlantic with examples from Norway. Both ecological, governance and management complexity increases with increasing geographic scale, with associated increases in uncertainty and thus increasing need for managing under the precautionary approach. A process where (ecologically) valuable and vulnerable areas are defined early on is essential to successful place-based management under the ecosystem approach. Integrating across sectors and achieving necessary cooperation between involved institutions and stakeholders is also necessary.     

Allometric scaling predicts preferences for burned patches in a guild of East African grazers

The high herbivore diversity in savanna systems has been attributed to the inherent spatial and temporal heterogeneity related to the quantity and quality of food resources. Allometric scaling predicts that smaller-bodied grazers rely on higher quality forage than larger-bodied grazers. We replicated burns at varying scales in an East African savanna and measured visitation by an entire guild of larger grazers ranging in size from hare to elephant. We found a strong negative relationship between burn preference and body mass with foregut fermenters preferring burns to a greater degree than hindgut fermenters. Burns with higher quality forage were preferred more than burns with lower quality forage by small-bodied grazers, while the opposite was true for large-bodied grazers. Our results represent some of the first experimental evidence demonstrating the importance of body size in predicting how large herbivores respond to fire-induced changes in plant quality and quantity.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Aggregation influences coral species richness at multiple spatial scales

The spatial dispersion of individuals across multiple spatial scales can significantly influence biodiversity patterns. Here we characterize the dispersion of corals in reef assemblages distributed across a 10000-km longitudinal biodiversity gradient from Indonesia to the Society Islands, using a multiscale sampling design. Our results indicate that most coral species were aggregated among 10-m transect samples across this vast distance. Using observed and randomized species sampling curves, we show that aggregation reduced the number of species per transect, site, and island sample on average by 13-27%. Across site, island, and regional scales, aggregation also reduced the area under species sampling curves by an average of 2.7-6.5%. The level of aggregation was relatively constant across spatial scales within regions and did not vary among habitats. However, there was significant variation among regions using transect samples across individual sites. Specifically, aggregation reduced the species richness per transect and the area under species sampling curves nearly twice as much in the Indonesian biodiversity hotspot than in the Society Islands. As a significant component of the spatial structure of coral assemblages, aggregation should be integrated into our understanding of coral community dynamics and the development of conservation strategies designed to protect these communities.     

Genetic structure of the common sole Solea vulgaris at different geographic scales

The genetic structure of the flatfish Solea vulgaris was investigated on several spatial scales and at the temporal level through analysis of electrophoretic variation at 8 to 12 polymorphic enzyme loci. No differentiation was apparent at the temporal scale. Some differentiation was detected at and above the regional scale. Isolation by distance was evidenced by the significant correlation between genetic and geographic distances, and by the consistency of the results of multiple-locus correspondence analysis with geographic sampling patterns. The analysis suggested that the geographic unit of population structure (i.e. a geographical area corresponding to a panmictic or nearly panmictic population) lies within a radius of the order of 100 km. The isolation-by-distance pattern in S. vulgaris contrasted with the known genetic structures of other flatfish species of the northeastern Atlantic and Mediterranean in a way that may be related to the range of their respective temperature tolerances for eggs and larvae.     

The influence of host diversity and composition on epidemiological patterns at multiple spatial scales

Spatial patterns of pathogen prevalence are determined by ecological processes acting across multiple spatial scales. Host-pathogen interactions are influenced by community composition, landscape structure, and environmental factors. Explaining prevalence patterns requires an understanding of how local determinants of infection, such as community composition, are mediated by landscape characteristics and regional-scale environmental drivers. Here we investigate the role of local community interactions and the effects of landscape structure on the dynamics of barley and cereal yellow dwarf viruses (B/CYDV) in the open meadows of the Cascade Mountains of Oregon. B/CYDV is an aphid-transmitted, generalist pathogen of over 100 wild and cultivated grass species. We used variance components analysis and model selection techniques to partition the sources of variation in B/CYDV prevalence and to determine which abiotic and biotic factors influence host-pathogen interactions in a Cascades meadowsystem. B/CYDV prevalence in Cascades meadows varied by host species identity, with a significantly higher proportion of infected Festuca idahoensis individuals than Elymus glaucus or Bromus carinatus. Although there was significant variation in prevalence among host species and among meadows in the same meadow complex, there was no evidence of any significant variation in prevalence among different meadow complexes at a larger spatial scale. Variation in prevalence among meadows was primarily associated with the local community context (host identity, the relative abundance of different host species, and host species richness) and the physical landscape attributes of the meadow. These results highlight the importance of local host community composition, mediated by landscape characteristics such as meadow aspect, as a determinant of the spatial pattern of infection of a multi-host pathogen.     

Bet hedging in a guild of desert annuals

Evolutionary bet hedging encapsulates the counterintuitive idea that organisms evolve traits that reduce short-term reproductive success in favor of longer-term risk reduction. It has been widely investigated theoretically, and many putative examples have been cited including practical ones such as the dormancy involved in microbe and weed persistence. However, long-term data on demographic variation from the actual evolutionarily relevant environments have been unavailable to test for its mechanistic relationship to alleged bet hedging traits. I report an association between delayed germination (a bet hedging trait) and risk using a 22-year data set on demographic variation for 10 species of desert annual plants. Species with greater variation in reproductive success (per capita survival from germination to reproduction x per capita fecundity of survivors) were found to have lower average germination fractions. This provides a definitive test using realistic data on demographic variance that confirms the life history prediction for bet hedging. I also showed that the species with greater long-term demographic variation tended to be the ones with greater sensitivity of reproductive success to variation among years in growing-season precipitation.     

Beta diversity at different spatial scales: plant communities in organic and conventional agriculture.

Biodiversity studies that guide agricultural subsidy policy have generally compared farming systems at a single spatial scale: the field. However, diversity patterns vary across spatial scales. Here, we examined the effects of farming system (organic vs. conventional) and position in the field (edge vs. center) on plant species richness in wheat fields at three spatial scales. We quantified alpha-, beta-, and gamma-diversity at the microscale in 800 plots, at the mesoscale in 40 fields, and at the macroscale in three regions using the additive partitioning approach, and evaluated the relative contribution of beta-diversity at each spatial scale to total observed species richness. We found that alpha-, beta-, and gamma-diversity were higher in organic than conventional fields and higher at the field edge than in the field center at all spatial scales. In both farming systems, beta-diversity at the meso- and macroscale explained most of the overall species richness (up to 37% and 25%, respectively), indicating considerable differences in community composition among fields and regions due to environmental heterogeneity. The spatial scale at which beta-diversity contributed the most to overall species richness differed between rare and common species. Total richness of rare species (present in < or = 5% of total samples) was mainly explained by differences in community composition at the meso- and macroscale (up to 27% and 48%, respectively), but only in organic fields. Total richness of common species (present in > or = 25% of total samples) was explained by differences in community composition at the micro- and mesoscale (up to 29% and 47%, respectively), i.e., among plots and fields, independent of farming system. Our results show that organic farming made the greatest contribution to total species richness at the meso (among fields) and macro (among regions) scale due to environmental heterogeneity. Hence, agri-environment schemes should exploit this large-scale contribution of beta-diversity by tailoring schemes at regional scales to maximize dissimilarity between conservation areas using geographic information systems rather than focusing entirely at the classical local-field scale, which is the current practice.     

Physiological capacity and environmental tolerance in two sandhopper species with contrasting geographical ranges: Talitrus saltator and Talorchestia ugolinii

We investigated the physiological plasticity and environmental tolerance of two phylogenetically closely related, ecologically similar and co-occurring species of supralittoral amphipods differing drastically in the size of their geographical ranges. A series of physiological traits were characterised for the Corsican-endemic Talorchestia ugolinii Bella-Santini and Ruffo and the widespread Talitrus saltator Montagu. The effect of body mass, temperature and salinity on heart rate (used as proxy for metabolic activity and stress), the effect of temperature on oxygen consumption and the tolerance to salinity exposure were investigated in both species, together with the characterisation of haemolymph osmoregulation in T. ugolinii. Our results showed that there is a clear difference in the resting metabolic rates and physiological capacity, as well as environmental tolerance, between T. saltator and T. ugolinii, with T. saltator overall showing a broader physiological niche. Although T. ugolinii showed a relatively good ability to regulate its haemolymph osmotic concentration (similar to that previously described for T. saltator), it demonstrated a lower tolerance to exposure to hypo-osmotic stress. In addition, a consistent picture emerged between the ability to control the cardiac function and the capacity to actively respond to osmotic stress. The physiological findings are discussed in relation to the known ecology and geographical distribution of T. ugolinii.     

Comparing network analysis methodologies for consumer–resource relations at species and ecosystems scales

This research compares two existing methodologies, mixed trophic impact analysis and utility analysis, which use network analysis to evaluate the direct, pair-wise, and indirect, holistic, ecological relations between ecosystem compartments. The two approaches have many similarities, but differ in some key assumptions which affect both the final results and interpretations. Here, we briefly introduce both methodologies through a series of two simple examples; a 3-compartment competition model and a 3-compartment food chain model, and then apply the methodologies to a 15-compartment ecosystem model of the Chesapeake Bay. This example demonstrates how implementing the various conceptual and methodological assumptions lead to differing results. Notably, the overall number of positive relations is greatly affected by the treatment of the self-interactions and the handling of detritus compartments lead to a distinction between ecological or trophic relations. We recommend slight changes to both methodologies, not necessarily in order to bring them completely together, but because each has some points which are stronger and better defensible.     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Genetic structure of the flounders Platichthys flesus and P. stellatus at different geographic scales

The genetic structure of the flounders Platichthys flesus L. and P. stellatus Pallas was investigated on different spatial scales through analysis of allozyme variation at 7 to 24 polymorphic loci in samples collected from different regions (Baltic Sea, North Sea, Brittany, Portugal, western Mediterranean, Adriatic Sea, Aegean Sea and Japan) in 1984 to 1987. No geographic variation was evident within a region. Some pattern of differentiation by distance was inferred within the Atlantic, while the Mediterranean comprised three geographically isolated populations and was itself geographically isolated from the Atlantic (fixed allele differences at up to three loci were found among P. flesus populations from the Atlantic, the western Mediterranean, the Adriatic Sea, the Aegean Sea and also P. stellatus from the coast of Japan). Sea temperature during the reproductive period probably acts as a barrier to gene flow between populations. Genetic distances among European flounder populations (P. flesus) were higher than, or of the same magnitude as, the genetic distance between Pacific (P. stellatus) and European flounder populations, suggesting that P. flesus is paraphyletic and/or there is no phylogenetic basis to recognising P. stellatus as a different species. The divergence between P. flesus and P. stellatus was thus inferred to be more recent than the divergence between the present P. flesus populations from the NE Atlantic and eastern Mediterranean. The eastern Mediterranean populations are thought to originate from the colonisation of the Mediterranean by a proto-P. flesus/P. stellatus ancestor, whereas the present western Mediterranean population has undergone a more recent colonisation event by P. flesus. Patterns of mitochondrial DNA variation, established on a smaller array of P. flesus samples, were in accordance with the geographic patterns inferred from the allozyme survey. In addition, they supported the hypothesis of a two-step colonisation of the western Mediterranean. These results contribute to our understanding of the biogeography of the Mediterranean marine fauna, especially the group of boreal remnants to which P. flesus belongs. Received: 7 February 1997 / Accepted: 26 March 1997     

Stepping-stone gene flow in the solitary coral Balanophyllia elegans: equilibrium and nonequilibrium at different spatial scales

Limited dispersal should result in genetic differences between populations proportional to geographic distances of separation. This association between gene flow and distance can be disrupted by (1) continuing genetic exchange among distant populations, (2) historical changes in gene flow, and (3) physical barriers or corridors to dispersal. The movements of larvae are thought to determine dispersal capability in benthic marine invertebrates. The solitary scleractinian Balanophyllia elegans Verrill possesses crawling larvae capable of only limited dispersal. Paradoxically, however, inferred levels of gene flow between pairs of localities spread over much of the 4000 km range of B. elegans exhibited a weaker relationship with geographical separation than that expected for a linear array of populations in which all genetic exchange takes place between adjacent populations. In this paper, I examined the pattern of gene flow (inferred from the frequencies of eight polymorphic allozyme loci) in B. elegans at a smaller (1 to 50 km) spatial scale to determine (1) whether gene flow at this spatial scale conformed to the expectations of the stepping-stone model, and (2) whether continuing long-distance gene flow or historical changes in gene flow were responsible for the weak relationship between gene flow and distance observed previously at the rangewide spatial scale. Between May and August 1992, I collected 75 adults from each of 18 localities along the coast of Sonoma County, California, USA. These populations of B. elegans were significantly subdivided both among localities separated by 1 to 50 km (F _LT =0.053, Se=0.0075) and among patches separated by 4 to 8 m (F PL=0.026, SE=0.0023). The observed slope and correlation (r ²=0.54) between inferred levels of gene flow and the geographic distance at the 1 to 50 km spatial scale conformed to equilibrium expectations (obtained by simulation) for a linear stepping-stone model, although those from the rangewide spatial scale did not. This implies that the mechanisms conferring patterns of inferred genetic differentiation between localities in B. elegans differ fundamentally with spatial scale. At a scale of 1 to 50 km, continuing gene flow and drift have equilibrated and the process of isolation-bydistance may facilitate local adaptive change. At a broader spatial scale, historical changes in gene flow, perhaps affected by late Pleistocene climatic fluctuations, disrupt the equilibration of gene flow and genetic drift, so that genetic differentiation may not increase continuously with separation between populations.     

Population dynamics of the barnacle <Emphasis Type="Italic">Chthamalus montagui</Emphasis> at two spatial and temporal scales in northern Spain

This study describes spatial and temporal patterns of variability in population parameters in the barnacle Chthamalus montagui Southward in three localities of northern Spain and evaluates whether density-dependent settlement may regulate population dynamics. The sampling design considered two spatial scales, localities and sites within localities, and two temporal scales, years and six month intervals. Density, amount of free space, mortality, growth rate and magnitude of settlement (both absolute and per unit of free space) were obtained from photographs of permanent quadrats and from direct counts in the field. The number of settlers in scraped and untouched quadrats was used to estimate the importance of the presence of conspecifics in settlement. Significant variation at the two spatial and temporal scales was found for most parameters. Large spatial and temporal variations in adult mortality rate, density, and settlement were observed. Patterns of mortality were not consistent with differences in density among localities. Differences in settlement among localities were maintained through time. We suggest that magnitude of settlement is regulated by persistent features such as topography or local water circulation. We assume that early post-settlement mortality does not differ among localities. In the absence of differential mortality, settlement determines average population density within localities. Within localities, settlement was independent of density and free space. No consistent evidence was found on preferential settlement at the vicinity of conspecifics. The main conclusion is that density-dependent settlement is not relevant for the regulation of the populations of C. montagui in the northern Spain. Regulation might occur by density-dependent processes within the adult fraction of the population and/or the larval phase before settlement.     

Interactions among bacterial-feeding nematode species at different levels of food availability

Accurate prediction of the biodiversity–ecosystem functioning relationship requires adequate understanding of the interactions among species in a community. Effects of species diversity on ecosystem functioning are usually considered more pronounced with increasing functional dissimilarity, although species within functional groups may also perform non-identical functions and interact with each other. Here we present results of a laboratory experimental study aimed at elucidating whether interspecific interactions among species within a single nematode trophic group, bacterivores, (1) affect population development and community structure, and (2) depend on food availability. We studied the population growth of Rhabditis (Pellioditis) marina, a rhabditid nematode known to favour very high food densities when in monoculture, and of Diplolaimelloides meyli and D. oschei, congeneric Monhysteridae known to perform better in monocultures at intermediate food availability. Both Diplolaimelloides species showed significantly different patterns of food-density dependence in combination culture compared to monoculture. At very high food availability, the rhabditid nematode facilitated growth of both monhysterid species, probably as a result of down-regulation of bacterial density. At the lowest food availabilities, the presence of even low numbers of monhysterid nematodes lead to exclusion of the rhabditid, which at such low food availability has a very inefficient food uptake. At intermediate food availabilities, abundances of both Diplolaimelloides species were strongly depressed in the combination culture, as a result of food depletion by the rhabditid, indirect inhibitory interactions between the two congeneric species, or both. The complexity of the species interactions render predictions on the outcome and functional consequences of changes in within-trophic-group diversity highly problematic.     

Neural network modelling for the analysis of forcings/temperatures relationships at different scales in the climate system

A fully non-linear analysis of forcing influences on temperatures is performed in the climate system by means of neural network modelling. Two case studies are investigated, in order to establish the main factors that drove the temperature behaviour at both global and regional scales in the last 140 years. In particular, our neural network model shows the ability to catch non-linear relationships among these variables and to reconstruct temperature records with a high degree of accuracy. In this framework, we clearly show the need of including anthropogenic inputs for explaining the temperature behaviour at global scale and recognise the role of El Niño southern oscillation for catching the inter-annual variability of temperature data. Furthermore, we analyse the relative influence of global forcing and a regional circulation pattern in determining the winter temperatures in Central England, showing that the North Atlantic oscillation represents the driven element in this case study. Our modelling activity and results can be very useful for simple assessments of relationships in the complex climate system and for identifying the fundamental elements leading to a successful downscaling of atmosphere–ocean general circulation models.     

Patterns of use of space in a guild of territorial reef fishes

There exists on Heron Reef, at the southern end of the Great Barrier Reef, an 8-species guild of ecologically very similar, territorial, herbivorous reef fishes. All individuals of these species maintain territories on rubble substrata throughout juvenile and adult life. Territories are defended from all other guild members. Three rubble patches, each containing residents of 3 guild species: Pomacentrus apicalis, P. wardi, and Abudafduf lachrymatus, have been monitored for 12 to 18 months. This paper examines the patterns of use of space as shown by the antecedent and subsequent histories of sites chosen by 43 new colonists, and sites vacated through the loss of 34 residents on the rubble patches. In addition, variation through time is examined in the total amount of space held in territories on each rubble patch. The 3 species show similar preferences for space as colonists, although adult and juvenile colonists behave differently. Numbers of colonists detected on rubble patches are not proportional to the resident populations of the 3 species. Residents of the 3 species are equal in their abilities to enter spaces vacated through mortality, although they differ slightly in methods used to enter sites. They do not respond preferentially to sites previously occupied by any particular species. P. wardi shows a higher rate of mortality than the other species, and residents of this species are more often dislodged by new colonists. The total amount of space held on any rubble patch did not vary during the year from September, 1972 to October, 1973. The data are interpreted as supporting the hypothesis that the 3 species do not differ in their requirements for space on rubble patches, and that they are in competition for a short supply of such space. Differences exist in their strategies for obtaining and holding such space. These differences are important for explaining the continued presence of P. wardi on rubble patches. P. apicalis and a. lachrymatus are both specialists in holding territories on the upper reef slope. No differences have been detected in their requirements or competitive abilities. P. wardi is a fugitive species on the upper reef slope, coexisting because it maintains a refuge from competition by occupying some un-preferred sites off rubble patches.     

Analyzing the spatial structure of a Sri Lankan tree species with multiple scales of clustering

Clustering at multiple critical scales may be common for plants since many different factors and processes may cause clustering. This is especially true for tropical rain forests for which theories explaining species coexistence and community structure rest heavily on spatial patterns. We used point pattern analysis to analyze the spatial structure of Shorea congestiflora, a dominant species in a 25-ha forest dynamics plot in a rain forest at Sinharaja World Heritage Site (Sri Lanka), which apparently shows clustering at several scales. We developed cluster processes incorporating two critical scales of clustering for exploring the spatial structure of S. congestiflora and interpret it in relation to factors such as competition, dispersal limitation, recruitment limitation, and Janzen-Connell effects. All size classes showed consistent large-scale clustering with a cluster radius of approximately 25 m. Inside the larger clusters, small-scale clusters with a radius of 8 m were evident for recruits and saplings, weak for intermediates, and disappeared for adults. The pattern of all trees could be divided into two independent patterns: a random pattern (nearest neighbor distance > 8 m) comprising approximately 12% of the trees and a nested double-cluster pattern. This finding suggests two independent recruitment and/or seed dispersal mechanisms. Saplings were several times as abundant as recruits and may accumulate several recruit generations. Recruits were only weakly associated with adults and occupied about half of the large-scale clusters, but saplings almost all. This is consistent with recruitment limitation. For approximately 70% (95%) of all juveniles the nearest adult was less than 26 m away (53 m), suggesting a dispersal limitation that may also be related to the critical large-scale clustering. Our example illustrates the manner in which the use of a specific and complex null hypothesis of spatial structure in point pattern analysis can help us better understand the biology of a species and generate specific hypotheses to be further investigated in the field.     

Importance of physical and biological settlement cues used at different spatial scales by the larvae of Semibalanus balanoides

The settling cues used by larvae of Semibalanus balanoides (L.) were examined at large (1 m), medium (1 mm), and small (<300 m) spatial scales, corresponding roughly to choices made during broad and close exploration and final inspection within the settling zone. The experiments were carried out at two locations in Canada in May and June 1984. Samples of substratum surfaces from above (n=87 sites unselected by cyprids) and below (n=84 sites selected by cyprids) the upper limit of the barnacle zone were characterized according to 13 quantitative and semi-quantitative variables (diatom cover, macroalgae, detritus) using scanning electron microscopy (SEM). The data were analysed using a stepwise logistic regression. At the large scale, the presence of an alga (Urospora wormskjoldii Mertens) in the upper uncolonized zone and a diatom (Achnantes parvula Kützing) within the settling zone are potential settlement cues. At the medium scale in the settling zone, larvae select clean sites, devoid of detritus and diatoms or on which detrital matter, when present, is of finer texture than on the unchosen sites within the settling zone. At the smallest scale (<300 m, equal to the average maximum width of the larvae), we tested the hypothesis that the larva seeks optimal microheterogeneity (presumably providing good adhesion). Microheterogeneity of the surface was measured immediately under the larva, around the settling organs, the antennules. Our results show that such surface microheterogeneity is significantly greater and the vertical center of gravity of the surface significantly lower on the sites selected by the cyprids than on the unselected contiguous sites. Fractal measurements carried out on selected and contiguous unselected surfaces indicate that larvae discriminate microheterogeneity levels below a step length of 35 m, a value approximately the size of the diameter of the antennular discs (40 m), which are used to explore the substratum and attach.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Behavioral aspects of competition in a three-species rodent guild of coastal southern California

Summary The influence of vole cycles on the demography, spatial organization, and abundance of potential rodent competitors was studied in three California rodents in the laboratory and in the field. Population densities ofMicrotus californicus were inversely correlated with reproductive success in two potential competitors (Reithrodontomys megalotis andMus musculus). Additionally, high-density vole populations forced these species into suboptimal habitats. During low-density vole populations,Reithrodontomys andMus left their refugia, expanded their usage of habitat, were more active, and their reproductive success was greater. Thus, this work suggests that during vole population peaks, competition with sympatric species of mice is severe. Conversely, when vole populations decline, competition is relaxed. The temporal and spatial organization of the three-species complex supports models of nonequilibrium and fugitive species coexistence.