Segregation of temporal and spatial distribution between kleptoparasites and parasitoids of the eusocial sweat bee, Lasioglossum malachurum (Hymenoptera: Halictidae, Mutillidae)

Cuckoo bees and velvet ants use different resources of their shared host bees, the former laying eggs on the host pollen stores and the latter on immature stages. We studied the activity patterns of the cuckoo bee Sphecodes monilicornis and the velvet ant Myrmilla capitata at two nesting sites of their host, the social digger bee Lasioglossum malachurum , over a 3 year period. Due to the difference in host exploitation, we expected different temporal patterns of the two natural enemies as well as a positive spatial association with host nest density for both species. At a daily level, S. monilicornis was more abundant between 10.00 and 15.00 h, while M. capitata was most active in the early morning and late afternoon; both species activities were independent from host provisioning activity. The activity of cuckoo bees was in general positively correlated with the density of open host nests (but not with the total number of nests), while that of velvet ants was rarely correlated with this factor. Sphecodes monilicornis was seen both attacking the guard bees and directly entering into the host nests or digging close to nest entrances, while M. capitata only gained access to host nests through digging. We conclude that the temporal and spatial segregation between the two species may be, at least partially, explained both by the different resources exploited and by the different dynamics of host interactions.     

Escape from parasitism: spatial and temporal strategies of a sphecid wasp against a specialised cuckoo wasp

Parasites and parasitoids exert an important selection pressure on organisms and, thus, play an important role for both population dynamics and evolutionary responses of host species. We investigated host-parasite interactions in a brood-caring wasp, the European beewolf, Philanthus triangulum (Hymenoptera, Sphecidae), and asked whether females of this species might employ temporal or spatial strategies to reduce the rate of attack by a specialised brood parasitoid, the cuckoo wasp Hedychrum rutilans (Hymenoptera, Chrysididae). Females of the host species might shift their activity to periods of low parasitoid activity both in the course of the season and in the course of the day. On a spatial scale, aggregated or dispersed nesting might be favoured depending on the form of the density dependence of parasitism. The beginning and end of the flight season of host and parasitoid were nearly identical. Activity of chrysidids relative to beewolves did not change significantly during the flight season. However, relative parasitoid activity declined in the course of the day, suggesting the existence of temporal enemy-free space in the evening hours. Shifting the main activity to the evening hours might be a flexible response of beewolves to the presence of chrysidids. Activity of cuckoo wasps per nest was independent of nest density but the actual rate of parasitism as revealed by nest excavations indicated direct density dependence. Total mortality, however, was inversely density dependent. Thus, in the study population aggregated nesting did not reduce parasitism but minimised total mortality.     

Evaluating the impact of a biological control parasitoid on invasive Vespula wasps in a natural forest ecosystem

The overall impact of the parasitoid Sphecophaga vesparum vesparum on invasive Vespula wasps in New Zealand native beech forest was evaluated by assessing the levels of parasitism achieved and the parasitoid’s effect at nest level and population level. The maximum proportion of nests parasitised was 17%, but there was no significant increase with time (r² = 0.139; p = 0.115). However, there was an exponential reduction in the number of parasitoids produced per parasitised nest from a peak of 570 (SE = 143) parasitoids per nest in 1990, declining to only 15 (SE = 6) parasitoids per nest in 2004. Even when parasitoid density was high, the parasitoid had no detectable impact on the number of small cells or the total host nest size, but it halved the number of large (reproductive) cells produced. This may have resulted in fewer queens produced per parasitised nest. Wasp nest density was highly variable from year to year, but there was no evidence that the wasp population density at the parasitised site (Pelorus Bridge) had declined relative to the five sites where the parasitoid had not established. We conclude that the parasitoid is unlikely to have had any significant effect on wasp populations hitherto, nor is it likely to impact host populations in the future. We recommend other biological control programs adopt pre-release assessment of per capita impact as a way of identifying agents that are more likely to be successful and hence minimising economic and potential ecological costs of biocontrol.     

Do digger wasps time their provisioning activity to avoid cuckoo wasps (Hymenoptera: Crabronidae and Chrysididae)?

Strategies adopted by parasitoids and kleptoparasites co-evolve with the defensive adaptations of their hosts, and vice-versa. Hedychrum rutilans and Hedychrum nobile are brood parasites of, respectively, Philanthus triangulum and Cerceris arenaria, two digger wasps that share most aspects of their nesting biology (solitary females dig aggregated nests in the ground and mass-provision the brood with paralyzed insects). We tested the hypothesis that similarity in the hosts’ nesting habits corresponds to similar defensive strategies against these cuckoo wasps. Peak provisioning activity by P. triangulum occurred in late afternoon (and early morning in 1 year) while peak H. rutilans activity was in early afternoon. In contrast, peak provisioning by C. arenaria and peak H. nobile activity occurred in early afternoon. Thus, P. triangulum (as previously found in other populations) appears to have timed its provisioning to avoid its brood parasite whereas C. arenaria did not, rejecting our hypothesis. The daily activity of both chrysidid wasps was positively correlated to air temperature. Host nest density positively affected only H. rutilans activity, in agreement with previous reports for other populations of this species, whereas the daily pattern of activity of the host was the key correlate with the activity of H. nobile. Mortality due to cuckoo wasps was low for both digger wasps, although it was somewhat higher at sites with many nests of P. triangulum. We suggest that perhaps the degree to which digger wasps time their activity to avoid cuckoo wasps is related to the degree of specialization in host choice by their brood parasite. H. rutilans is more highly dependent on P. triangulum, which may have had a greater effect on the timing of provision by its host.     

Presence of an unsuitable host diminishes the competitive superiority of an insect parasitoid: a distraction effect

Competitive interactions between parasitoid species are traditionally evaluated when they compete for a single host species. Yet, the presence of additional host species can alter competitive interactions, even if the host is unsuitable for parasitoid development. In alfalfa of the mid-western USA, a native parasitoid species, Praon pequodorum, was once a dominant natural enemy, but it has become rare since the introduction of another parasitoid, Aphidius ervi. Despite A. ervi’s competitive superiority for their most common host, the pea aphid Acyrthosiphum pisum, P. pequodorum still persists at low densities. We performed a suite of laboratory and field studies to determine if the presence of an alternative host, the spotted alfalfa aphid Therioaphis maculata, may mitigate A. ervi’s competitive superiority and facilitate P. pequodorum’s persistence. We show that spotted alfalfa aphids reduce the foraging efficiency of both parasitoid species for pea aphids, despite spotted alfalfa aphids being an unsuitable host. This decrease in efficiency, however, was not symmetrical; the presence of spotted alfalfa aphids had a greater detrimental effect on A. ervi foraging for pea aphids. This might facilitate the persistence of the competitively inferior P. pequodorum. Our study suggests that indirect effects generated by the presence of alternative hosts are important for understanding parasitoid–host dynamics and overall insect community structure.     

Invertebrate predation of 15N-marked prey in semi-field wheat enclosures

One of the most famous examples of successful, classical biological control in Japan is the introduction of the parasitoids Coccobius fulvus and Aphytis yanonensis against the citrus pest arrowhead scale Unaspis yanonensis. Together, they comprise a host‐parasitoid system that has been demonstrated to be stable. To test the conventional theory that successful biological control of pests occurs through the establishment of a low stable equilibrium, brought about by the density‐dependent responses of natural enemies to the pest species, sampling was carried out at five sites in the field during 2000 and 2001 to examine the relationship between the rate of parasitism by C. fulvus and the density of its host. The data were analysed using three statistical techniques at nine spatial scales. Contrary to conventional theoretical predictions, each method of analysis detected very little density‐dependence at any spatial level in this study. Parasitoid aggregations independent of host density were not sufficient to stabilise host–parasitoid interactions. Our results suggest that neither spatial density‐dependent nor density‐independent parasitism is necessary for successful biological control, or for the stability of the host–parasitoid system. We propose an alternative mechanism: a spatial refuge induced by parasitoid introduction may stabilise a system.     

High nymphal host density and mortality negatively impact parasitoid complex during an insect herbivore outbreak

Insect herbivore outbreaks frequently occur and this may be due to factors that restrict top-down control by parasitoids, for example, host-parasitoid asynchrony, hyperparasitization, resource limitation and climate. Few studies have examined hostparasitoid density relationships during an in sect herbivore outbreak in a n atural ecosystem with diverse parasitoids. We studied parasitization patterns of Cardiaspina psyllids during an outbreak in a Eucalyptus woodland. First, we established the trophic roles of the parasitoids through a species-specific multiplex PCR approach on mummies from which parasitoids emerged. Then, we assessed host-parasitoid density relationships across three spatial scales (leaf, tree and site) over one yeas We detected four endoparasitoid species of the family Encyrtidae (Hymenoptera);two primary parasitoid and one heteronomous hyperparasitoid Psyllaephagus species (the latter with female development as a primary parasitoid and male development as a hyperparasitoid), and the hyperparasitoid Coccidoctonuspsyllae. Parasitoid development was host-synchronized, although synchrony between sites appeared constrained during winter (due to temperature differences). Parasitization was predominantly driven by one primary parasitoid species and was mostly inversely host-density dependent across the spatial scales. Hyperparasitization by C. psyllae was psyllid-density dependent at the site scale, however, this only impacted the rarer primary parasitoid. High larval parasitoid mortality due to density-dependent nymphal psyllid mortality (a consequence of resource limitation) compounded by a summer heat wave was incorporated in the assessment and resulted in density independence of host-parasitoid relationships. As such, high larval parasitoid mortality during insect herbivore outbreaks may contribute to the absence of host density-dependent parasitization during outbreak events.     

Evaluation of Diachasmimorpha longicaudata (Hymenoptera: Braconidae) as a mortality factor of Ceratitis capitata (Diptera: Tephritidae) infesting Citrus species under laboratory and field-cage conditions

Ceratitis capitata (or medfly) is one of the major pests currently affecting fruit crops in northwestern Argentinian Citrus-producing areas. Medfly populations are sustained in large exotic fruits, such as Citrus paradisi, Citrus aurantium and Citrus sinensis, which are known to hinder the activity of almost all native parasitoid species. Therefore, a feasible approach to controlling medfly involves the use of exotic parasitoids such as Diachasmimorpha longicaudata. In this study, the prediction that parasitoid females would be proficient at finding medfly larvae infesting the Citrus species mentioned earlier was tested. Particularly, the variation in fruit species preference by parasitoid females, the efficacy of the parasitoid to kill medfly and the effect of host density on parasitoid performance were determined. Parasitoids were allowed to forage for 8 h on grapefruits and oranges artificially infested with medfly larvae under controlled (laboratory) and uncontrolled (field cage) environmental conditions. Fruit choice and no-choice tests were performed. Results were assessed by comparing the number of female visits to and ovipositor insertions into the fruit, and parasitoid emergence, parasitism and additional host mortality percentages. Parasitoid preference for visiting larger fruits (sour orange and grapefruit) may be related to the increased fruit surface area. Ovipositional activity on fruit was influenced by the variation of the larval host density per unit of fruit surface. The higher parasitism rates recorded from sweet orange would be mainly a result of both increased host density and fruit physical features. Nevertheless, D. longicaudata showed the capacity to parasitise hosts in all Citrus species tested.     

Parasitoids of the goldenrod gall moth: effects of scale on spatial density dependence

Summary A parasitoid assemblage consisting of four hymenopteran species caused larval and pupal mortality of the host Gnorimoschema gallaeosolidaginis. In the absence of discrete host patches, the relationship between densities of hosts and parasitized hosts was examined on scales of 0.25, 1, 4, 16 and 25 m² within a 400 m² plot and between ten 16 m² plots within a 1·3 km² area. Within the 400 m² plot, positive density dependence was observed on scales of 1, 4 and 16 m². Rates of parasitism were found to be density independent across the ten 16 m² plots. The nature of parasitoid and host spatial patterns were examined at the 400 m² plot by using Goodall's (1974) paired-quadrat variance function, and by fitting the observed distribution of galls and parasitized galls to the Poisson and negative binomial on three scales. Postive density dependence at the 400 m² plot occurred in the context of a host density gradient and may indicate aggregation of search time by parasitoids. However, significant responses on scales of 4 and 16 m² may only reflect the sum of parasitoid responses on lower scales because of this gradient. Predominance of significant responses on a 1 m² scale may be related to the high degree of variation in host density, and limits to active searching range of parasitoids.     

Do distances among host patches and host density affect the distribution of a specialist parasitoid?

The effect of spatial habitat structure and patchiness may differ among species within a multi-trophic system. Theoretical models predict that species at higher trophic levels are more negatively affected by fragmentation than are their hosts or preys. The absence or presence of the higher trophic level, in turn, can affect the population dynamics of lower levels and even the stability of the trophic system as a whole. The present study examines different effects of spatial habitat structure with two field experiments, using as model system the parasitoid Cotesia popularis which is a specialist larval parasitoid of the herbivore Tyria jacobaeae. One experiment examines the colonisation rate of the parasitoid and the percentage parasitism at distances occurring on a natural scale; the other experiment examines the dispersal rate and the percentage parasitism in relation to the density of the herbivore and its host plant. C. popularis was able to reach artificial host populations at distances up to the largest distance created (at least 80 m from the nearest source population). Also, the percentage parasitism did not differ among the distances. The density experiment showed that the total number of herbivores parasitised was higher in patches with a high density of hosts, regardless of the density of the host plant. The percentage parasitism, however, was not related to the density of the host. The density of the host plant did have a (marginally) significant effect on the percentage parasitism, probably indicating that the parasitoid uses the host plant of the herbivore as a cue to find the herbivore itself. In conclusion, the parasitoid was not affected by the spatial habitat structure on spatial scales that are typical of local patches.     

Zombie fire ant workers: behavior controlled by decapitating fly parasitoids

Laboratory observations were conducted on four separate red imported fire ant, Solenopsis invicta, colonies that contained workers parasitized by the decapitating fly, Pseudacteon tricuspis. Parasitized S. invicta workers remained inside the nest during parasitoid larval development and left the nest approximately 8 – 10 hours before decapitation by the parasitoid. When parasitized ants left the nest, they were highly mobile, were responsive to tactile stimuli, and showed minimal defensive behavior. Ants ultimately entered into a grass thatch layer, where they were decapitated and the fly maggots pupariated. This study reveals that parasitized ants exhibit behaviors that are consistent with host manipulation to benefit survival of the parasitoid. Received 9 November 2006; revised 26 January 2007; accepted 7 February.     

Density-dependent dispersal in host-parasitoid assemblages

Most spatial population models assume constant rates of dispersal. However, in a given community, dispersal may not only depend on the density of conspecifics, i.e. density‐dependent dispersal, but also on the density of other species, a phenomenon we term ‘community‐dependent dispersal’. We co‐vary the densities of both the beetle host Callosobruchus chinensis and its parasitoid wasp, Anisopteromalus calandrae, in a laboratory study and record the proportions of each species that disperse within a two‐hour period. The parasitoid in these systems exhibits community‐dependent dispersal – dispersing more frequently when parasitoid density is high and larval host density is low. This supported our prediction that individuals should disperse according to competition for available resources. However, in this study the host's dispersal was independent of density. We suggest that this may be due to less intense selection acting on host dispersal strategies than on the parasitoid. We consider some possible consequences of community‐dependent dispersal for a number of spatial population processes. A well‐known host‐parasitoid metapopulation model is expanded so that it includes a greater range of dispersal functions. When the model is parameterised with the parasitoid community‐dependent dispersal function observed in the empirical study, similar population dynamics are obtained as when fixed‐rate dispersal functions are applied. The importance of dispersal functions for invasions of both competitive and host‐parasitoid systems is also considered. The model results demonstrate that understanding how individuals disperse in response to different species’ population densities is important in determining the rate of spread of an invasion. We suggest that more empirical studies are needed to establish what determines dispersal rate and distance in a range of species, combined with theoretical studies investigating the role of the dispersal function in determining spatial population processes.     

Parental Care Syndromes in House Sparrows: Positive Covariance Between Provisioning and Defense Linked to Parent Identity

Parents of many species provide multiple forms of care to their offspring. In many birds, parents often provision offspring with food and defend them from predators and/or nest‐site competitors. We tested how these two forms of parental care covary in a wild population of house sparrows (Passer domesticus). Using a behavioral reaction norm approach, we found that nestling provisioning exhibited between individual differences and positively covaried with a measure of nest defense: the propensity to attack a heterospecific nest box competitor, the European starling (Sturnis vulgaris). This result would seem to support parental investment theory and suggests that high‐provisioning parents have high‐value offspring, which they will defend more vigorously than low‐provisioning parents. In addition, we found that parents with nestlings that hatched earlier in the season and grew faster approached a model starling more frequently and tended to be more likely to strike the starling. However, we also found that although brood value explained significant variation in both nestling provisioning and nest defense, it did not eliminate the positive, between‐individual relationship between provisioning and defense. This suggests that some of the correlation between provisioning and defense is tied to individual identity and hence may be a behavioral syndrome in which differences between individuals in underlying attributes produce correlated behaviors.     

The relationship between variable host grouping and functional responses among parasitoids of Antispila nysaefoliella (Lepidoptera: Heliozelidae)

Our study investigated the importance of variability in the parasitoid community as a source of selection on host group size using a field population of the tupelo leafminer, Antispila nysaefoliella Clemens, which specializes on tupelo, Nyssa sylvatica Marsh. Larvae were collected from leaves with variable numbers of larvae and screened for parasitism using polymerase chain reaction of mitochondrial cytochrome oxidase I using markers designed specifically for amplifying parasitoid DNA while excluding host DNA. This method of selective PCR was effective for detecting the presence and identifying species of immature stages of three hymenopteran superfamilies: Chalcidoidea, Ichneumonoidea and Platygastroidea, which represented 83.4%, 16.0% and 0.6% of the total detectable parasitism, respectively. Our resulting sequences were then calibrated with sequences from identified adult parasitoids that had been either reared or field‐captured. A cluster analysis revealed 10 distinct clades that showed differences in attack patterns with respect to host traits and season. Total parasitism followed an inverse density‐dependent or density‐independent pattern with respect to host density (number per leaf). However, when parasitoid taxa were considered separately, one clade, which could be a cryptic species of Pnigalio maculipes Crawford (Chalcidoidea: Eulophidae), was found to increase its per leaf attack rate with host density. Our results suggest that parasitoid community composition and differences among species in their attack strategies can play a large role in determining the adaptive advantage of host grouping.     

Density dependence of Anaphes sordidatus (Hymenoptera: Mymaridae) parasitism on eggs of Listronotus oregonensis (Coleoptera: Curculionidae)

Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) oviposits in carrot leaves and the larvae feed in carrot roots. Its eggs are parasitized by Anaphes sordidatus (Girault) (Hymenoptera: Mymaridae) throughout its oviposition period. This parasitoid is the major biotic mortality factor for L. oregonensis. Parasitism by A. sordidatus was studied for three years in sequentially sown plots of carrots, Daucus carota var. sativa. Over the scason, significantly fewer L. oregonensis eggs were oviposited in later sown carrots than in earlier sown carrots because oviposition started later in late sown plots of carrots. A positive temporal density-dependent relationship was observed each year between parasitism rates and host densities. This positive density dependence occurred in early and mid-summer for earlysown carrots where host density reached 1–2 host eggs per plant but disappeared in late summer when host density decreased while parasitism remained high. Latesown carrots had low host egg density (0.2 host egg per plant) and contributed little to the total number of eggs. In these late sown plots, parasitism increased rapidly to over 80% but no density dependence was observed. Spatially, few statistically significant regressions were found but all indicated a positive spatial density-dependence. Most non-significant regressions occurred because the range of egg density was too small between plots for a given date.     

Spatial density‐dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae)

The density‐dependence in parasitism by the robber fly Mallophora ruficauda (Diptera: Asilidae) on scarab beetle larvae (Coleoptera: Scarabaeidae) populations was studied in the present research. Mallophora ruficauda is a pestiferous species common in the open grasslands of the Pampas region of South America. Adults are predators of insects and larvae are solitary parasitoids of third instar larvae of several species of scarab beetle (Coleoptera: Scarabaeidae). In contrast with most studied host‐parasitoid interactions, host searching by M. ruficauda is carried out by both larvae and adults. Typically, robber fly females lay eggs on tall grasses from where larvae drop to the ground, and attack hosts which are buried in the soil. We carried out our study at two spatial scales close to 14 apiaries located in the provinces of Buenos Aires and Entre Ríos (Argentina). We found that parasitism is density‐independent at the larger spatial scale and inversely density‐dependent at the smaller one. We also found that M. ruficauda selects Cyclocephala signaticollis among several scarab beetle species. Specificity is observed both at large and small spatial scales. We discuss the implications of both host specificity and host searching behaviour on the observed parasitism patterns.     

Host stage preference and parasitism behaviour of Aenasius bambawalei an encyrtid parasitoid of Phenacoccus solenopsis

In Pakistan, the cotton mealybug, Phenacoccus solenopsis Tinsley (Sternorrhyncha (Homoptera): Pseudococcidae), is a serious pest of many cultivated plants. A parasitoid, Aenasius bambawalei Hayat (Hymenoptera: Encyrtidae), is associated with P. solenopsis. In order to mass rear A. bambawalei for a biological control programme, it is important to investigate the parasitoid’s host stage preference and its parasitism behaviour for P. solenopsis in order to optimise production. The present study showed that under both choice and no-choice conditions, the parasitoid preferred third instar and pre-reproductive host stage mealybugs for parasitism. Parasitoid larva developing inside the host exhibited a greater longevity, shorter developmental period and longer body size in these preferred host stages. Our study also confirmed that A. bambawalei showed no attraction to male mealybugs and no host feeding on any host stage was recorded. The ability of the parasitoid to effectively discriminate between suitable and non-suitable stages means that it is feasible to rear it on a mixed population.     

The effect of multi‐species host density on superparasitism and sex ratio in a gregarious parasitoid

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Reproduction now or later: optimal host-handling strategies in the whitefly parasitoid Encarsia formosa

We developed a dynamic state variable model for studying optimal host‐handling strategies in the whitefly parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae). We assumed that (a) the function of host feeding is to gain nutrients that can be matured into eggs, (b) oögenesis is continuous and egg load dependent, (c) parasitoid survival is exponentially distributed and (d) parasitoids encounter hosts randomly, are autogenous and have unlimited access to non‐host food sources to obtain energy for maintenance and activity. The most important prediction of the model is that host feeding is maladaptive under field conditions of low host density (0.015 cm^?2) and short parasitoid life expectancy (maximum reproductive period of 7 d). Nutrients from the immature stage that can be matured into eggs are sufficient to prevent egg limitation. Both host density and parasitoid life expectancy have a positive effect on the optimal host‐feeding ratio. Parasitoids that make random decisions gain on average only 35% (0.015 hosts cm^?2) to 60% (1.5 hosts cm^?2) of the lifetime reproductive success of parasitoids that make optimal decisions, independent of their life expectancy. Parameters that have a large impact on lifetime reproductive success and therefore drive natural selection are parasitoid life expectancy and the survival probability of deposited eggs (independent of host density), the number of host encounters per day (when host density is low) and the egg maturation rate and number of host types (when host density is high). Explaining the evolution of host‐feeding behaviour under field conditions requires field data showing that life expectancy in the field is not as short as we assumed, or may require incorporation of variation in host density. Incorporating variation in walking speed, parasitised host types or egg resorption is not expected to provide an explanation for the evolution of host‐feeding behaviour under field conditions.     

Nest site choice by the intertidal spider Desis formidabilis (Araneae: Desidae) and nest utilisation by its hymenopteran egg parasitoid

1. Echthrodesis lamorali Masner, 1968 is the only known parasitoid of the eggs of the intertidal rocky shore spider Desis formidabilis O.P. Cambridge 1890 and is endemic to a small area of South Africa. 2. The abundance of spider nests and parasitoid presence were assessed in relation to their in‐ and between‐shore location at multiple sites within the distribution of E. lamorali along the Cape Peninsula (Western Cape, South Africa). 3. Desis formidabilis nests were more abundant in the mid‐shore zone than higher up or lower down the shore. Spider population sizes also differed between collection sites, with higher numbers recorded on the cooler western coast of the peninsula. 4. Evidence of parasitoid activity was recorded in 43.31% of the 127 nests and 13.85% of the 592 egg sacs they contained. 5. Where parasitoids gained entry to a spider egg sac, oviposition took place into all of the eggs present. 6. Incidence of wasp activity was positively correlated with spider nest concentration, not with height up the shore, suggesting that both the host and parasitoid are tolerant of salt‐water inundation. 7. These results should assist managers of the Table Mountain National Park, in which the full distribution of E. lamorali falls, to better understand this component of rocky shore community dynamics.