On the optimal compromise for a dispersing parasitoid

Vector optimization concepts are used to demonstrate that key characteristics of searching and oviposition by the parasitoid Anagrus delicatus (Mymaridae, Hymenoptera) are consistent with the idea that it forages to achieve an optimal compromise, in an uncertain environment, between maximizing ovipositions at suitable sites and dispersing its eggs as widely as possible.     

Patch-leaving rules for parasitoids with imperfect host discrimination

Abstract.

• 1 The solitary parasitoid Anagrus delicatus attacks hosts (plant-hopper eggs) that exist in distinct batches and that are readily detectable to the foraging parasitoid. However, Anagrus delicatus attacks only a small fraction of the available hosts within a batch of eggs and then disperses (Cronin & Strong, 1993a). Here we address the question: Why is A.delicatus abandoning seemingly high quality patches?
• 2 A parasitoid with an imperfect ability to discriminate between unparasitized hosts and hosts that it has attacked earlier within the same patch experiences a risk of self-superparasitism when attacking multiple hosts within a single patch. Self-superparasitism can incur costs in the form of lost time and eggs. Early patch leaving can be favoured as a means of avoiding the costs of self-superparasitism.
• 3 A simple static model demonstrates that patch leaving is favoured by low costs of travelling to a new patch, high error rates in discriminating previously self-parasitized hosts, and high levels of parasitism in the currently occupied patch.
• 4 A more detailed dynamic state variable model, parameterized for A.delicatus, demonstrates that this parasitoid's seemingly enigmatic behaviour can be explained under our hypothesis. In order for this to be the case, we predicted that A.delicatus cannot recognize previously parasitized hosts. Subsequent to our prediction, Cronin & Strong (1993b) demonstrated that experienced A.delicatus do not avoid ovipositing in previously self-parasitized hosts.
• 5 Optimal patch leaving rules can be highly sensitive to even very low host discrimination error rates, which may be widespread among parasitoids.

     

Superparasitism and mutual interference in the egg parasitoid Anagrus delicatus (Hymenoptera: Mymaridae)

Abstract.

• 1 In nature, interference among Anagrus delicatus (Hymenoptera: Mymaridae) parasitoids reduced the per-capita number of hosts parasitized. Interference increased with parasitoid density.
• 2 Anagrus delicatus did not avoid parasitizing hosts that had recently been parasitized by conspecific wasps. Evidence indicated that this superparasitism was largely a random process, increasing with the ratio of parasitized to unparasitized hosts.
• 3 Individual parasitoid efficiency, the number of hosts killed per wasp per unit time, decreased with increasing wasp density. This occurred whether wasps searched the patch together (simultaneously) or one by one (sequentially), and was the result of an increase in time spent superparasitizing hosts at higher wasp density. This is known as indirect mutual interference.
• 4 Increasing numbers of parasitoids together on the same patch caused a significant decline in the rate and per-capita number of hosts parasitized. However, there was not a correspondent decline in searching efficiency with increasing wasp density (i.e. no direct mutual interference).
• 5 These forms of parasitoid density dependence should contribute to the stability of the host—parasitoid interaction.

     

Production of Anagrus epos Girault (Hymenoptera: Mymaridae) on Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae) eggs

Anagrus epos Girault (Hymenoptera: Mymaridae) is a natural enemy candidate for a classical biological control program targeting the glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), in California. Little is known about the biology or ecology of A. epos when it utilizes GWSS eggs as a host. Here, we report the results of laboratory studies that describe the host age preference for oviposition, longevity of A. epos adults provided with different food sources, and developmental rates at six different constant temperature regimes. Anagrus epos is a gregarious parasitoid in GWSS eggs with up to 14 adults emerging from each GWSS egg. In choice and no-choice tests for oviposition, A. epos females successfully parasitized all developmental ages of GWSS eggs (1–8 days old). In choice tests, parasitism rates were significantly higher in 1-, 3-, 4-, and 5-day-old GWSS eggs than in 2-, 6-, 7-, and 8-day-old eggs. If provided with honey and water, honey only, water only, or no food or water, A. epos females lived on average 8.2, 4.7, 2.6, and 1.6 days, respectively. Anagrus epos required 294.1 degree-days above a lower temperature threshold of 12.4 °C to develop from egg to adult (eclosion). Our results provide baseline information useful in the development of an efficient parasitoid mass rearing program for A. epos release and evaluation in California.     

Offshore refuges support higher densities and show slower population declines of wintering Ruddy Turnstones Arenaria interpres

ABSTRACT

Capsule: Wintering Ruddy Turnstones Arenaria interpres occur in higher densities and their populations decline less on, or close to, offshore refuges than on mainland sites subject to greater levels of human disturbance.

Aim: To compare wintering densities of Ruddy Turnstones and changes in counts across time from sites with differing levels of human disturbance.

Methods: Long-term counts of Ruddy Turnstones (1998/1999 to 2015/2016) were used from 19 sites (two offshore refuges and 17 mainland sites subject to higher levels of human disturbance) in northeast England. No direct measure of human disturbance was available for our mainland sites; instead we used questionnaires (n?=?690) to understand how far people travelled to visit the coast and then used this distance with human population densities in a buffer around each site as a proxy for human disturbance levels.

Results: After controlling for the extent of their preferred habitat at each site (rocky shore) we found: (i) the closer each of the 19 sites was to the nearest offshore refuge the higher the density of Ruddy Turnstones and (ii) bird counts were stable at the two refuge sites, whereas, on average, counts declined at the 17 mainland sites. However, no relationship was found between Ruddy Turnstone counts from 17 mainland sites and human population densities within differing distances from each site (up to 10?km).

Conclusions: Our work suggests that Ruddy Turnstones made greater use of relatively undisturbed areas (offshore refuges) than those subject to greater disturbance by humans (mainland sites). Although the use of refuges and mainland in our study area was not well known, observations from 11 radio-tagged Ruddy Turnstones suggest that individual birds did use both locations. In a broader context, our work concurs with other studies that highlight the need for refuges with limited or no human access.     

The influence of salt and nitrogen on herbivore abundance: direct and indirect effects

We report on the influence of experimentally increased interstitial salinity and plant nitrogen on the abundance of the delphacid planthopper, Prokelisia marginata (Van Duzee) (Homoptera: Delphacidae), which feeds on salt marsh cordgrass, Spartina alterniflora. We also report the effects of these treatments on parasitism of P. marginata eggs by the fairyfly parasitoid, Anagrus sophiae (Hymenoptera: Mymaridae). Soil salinity was significantly elevated following the addition of salt pellets broadcast over the ground and plant foliar nitrogen was significantly increased after the addition of fertilizer. The addition of fertilizer increased P. marginata densities on Spartina but addition of salt did not. Neither treatment significantly affected levels of egg parasitism by A. sophiae. In this system direct effects of plants on their herbivores via changes in plant chemistry appear more important than indirect effects of plants on herbivores via their natural enemies. Received: 1 August 1997 / Accepted: 29 September 1997     

The responses ofPraon spp. parasitoids to aphid sex pheromone components in the field

In autumn 1991, aphid parasitoids of the genusPraon (Hymenoptera; Braconidae) were caught in water traps with lures containing synthetic aphid sex pheromone components at three sites in England and one in Germany. At two of the English sites and at the German site, the traps were placed in winter cereal fields whilst the third English site was in woodland. Three species were caught,P. volucre, P. dorsale andP. abjectum. Those caught in cereal fields were almost entirelyP. volucre, whilstP. dorsale dominated at the woodland site. Of the known aphid sex pheromone components, the most effective lure was the (+)-(4aS, 7S, 7aR)-nepetalactone. Nepetalactone traps placed at the woodland site in spring and summer caught fewPraon females, and attraction may be confined to the autumn, when sexual female aphids are present in the field. Male parasitoids did not respond to the aphid pheromones at any time, although they were caught in suction traps operated at the woodland site during the autumn. At the cereal sites height had a significant influence on the efficiency of the pheromone traps, those placed just above the crop canopy being most effective. There was no evidence that any other genus of parasitoid responded to aphid sex pheromones at these sites.     

The impact of sulfur on the reproductive success of <Emphasis Type="Italic">Anagrus</Emphasis> spp. parasitoids in the field

Pesticides targeted at pest species have often been demonstrated to have strong adverse effects on the survival of biological control agents in short-term laboratory bioassays; however, studies examining the influence of pesticides on the actual reproductive success of biological control agents in the field are rare. Because natural enemy reproduction is often directly tied to biological control success, effects of pesticides on reproduction are of central importance. Here we use a new technique to examine the influence of sulfur, a fungicide widely used in grape production, on the reproductive success of Anagrus erythroneurae (Hymenoptera: Mymaridae) and Anagrus daanei (Hymenoptera: Mymaridae), egg parasitoids of the grape leafhopper, Erythroneura elegantula (Homoptera: Cicadellidae). Sulfur has previously been shown to be highly toxic to Anagrus spp. in short-term laboratory and field bioassays, creating the expectation that sulfur should also reduce Anagrus reproductive success in the field. Surprisingly, in two studies, the first comparing the oviposition success of Anagrus collected live in paired sulfur-treated versus untreated vineyards and the second comparing the lifetime reproductive success of Anagrus collected at the end of their lives in unpaired sulfur-treated versus untreated vineyards, we found no effect of sulfur on parasitoid reproductive success. In this system, traditional short-term assays of laboratory toxicity do not appear to predict effects on parasitoid reproductive success, suggesting that demographic approaches to assessing the disruptive effects of pesticides may have an important role in designing IPM programs.     

The effect of sulfur on biological control of the grape leafhopper, <Emphasis Type="Italic">Erythroneura elegantula</Emphasis>, by the egg parasitoid <Emphasis Type="Italic">Anagrus erythroneurae</Emphasis>

We examined the toxicity of a fungicide, sulfur, to the egg parasitoid Anagrus erythroneurae (Hymenoptera: Mymaridae) Trjapitsyn and Chiapini and the vineyard leafhopper pest Erythroneura elegantula Osborn (Homoptera: Cicadellidae) and tested whether or not the use of sulfur in the field affects biological control of E. elegantula. Using field cage bioassays, we demonstrated that sulfur is toxic to adult A. erythroneurae parasitoids, but not toxic to adult E. elegantula leafhoppers. We nonetheless found in a field experiment that sulfur produced no changes in rates of parasitism or E. elegantula egg density, and generated only a very small increase in the density of E. elegantula nymphs. These results suggest that sulfur, although toxic to A. erythroneurae, is not highly disruptive of E. elegantula biological control in vineyards. Our results suggest that simple bioassays of acute toxicity may not accurately predict the impact of agricultural chemicals on biological control.     

The use of energy stores in the puerulus of the spiny lobster Jasus edwardsii across the continental shelf of New Zealand

Nektonic pueruli of the spiny lobster, Jasus edwardsii, were caught from two locations about 20 km apart across the continental shelf on the south east of the North Island, New Zealand. The pueruli were assayed for total protein, glucose, glycogen, and lipid content. Only the lipid content differed between pueruli caught onshore and offshore (mean difference=3.1 mg or 3.4% of dry mass). The average difference in lipid content measured over this distance was used to calculate the rate of energy consumption and timing for pueruli to actively swim from the continental shelf to shore. These results confirmed previous theoretical estimates and indirect measures. Furthermore, the rate of energy consumption would allow all of the pueruli caught offshore to swim to shore based on their total measured lipids. However, some individuals with low energy stores may be energetically compromised at arrival which may affect their subsequent moulting and survival. The results of this study indicate that lipid is the primary format for energy storage of the nektonic puerulus of the spiny lobster and that these lipid reserves have sufficient energetic capacity to allow the puerulus to actively swim the distance across the shelf to settle on the coast.     

Consequences of nitrogen and phosphorus limitation for the performance of two planthoppers with divergent life-history strategies

Phytophagous insects have a much higher nitrogen and phosphorus content than their host plants, an elemental mismatch that places inherent constraints on meeting nutritional requirements. Although nitrogen limitation is well documented in insect herbivores, phosphorus limitation is poorly studied. Using factorial experiments in the laboratory and field, in which levels of soil nitrogen and phosphorus were manipulated, we studied the relative consequences of macronutrient limitation for two herbivores, namely the phloem-feeding planthoppers Prokelisia dolus and P. marginata. These planthoppers inhabit the salt marshes of North America where large stands of their Spartina host plant are found. Notably, these congeners differ in their dispersal abilities; P. marginata is dispersive whereas P. dolus is sedentary. Both nitrogen and phosphorus subsidies enhanced the nitrogen and phosphorus content of Spartina. When P. dolus and P. marginata were raised on plants with an enriched nitrogen signature, they exhibited greater survival, grew to a larger size, developed more rapidly, and achieved higher densities than on nitrogen-deficient plants. However, P. marginata experienced greater fitness penalties than P. dolus on nitrogen-deficient plants. Phosphorus limitation and associated fitness penalties were not as severe as nitrogen limitation for P. marginata, and were not detected in P. dolus. The tempered response of P. dolus to N- and P-deficient Spartina is probably due to its greater investment in feeding musculature and hence ability to compensate for nutrient deficiencies with increased ingestion. To cope with deteriorating plant quality, P. dolus employs compensatory feeding, whereas P. marginata disperses to higher quality Spartina. When its option of dispersal is eliminated and P. marginata is confined on nutrient-deficient plants, its performance is drastically reduced compared with P. dolus. This research highlights the importance of interfacing herbivore life-history strategies with ecological stoichiometry in order to interpret the consequences of macronutrient limitation on herbivore performance and population dynamics.     

Effect of legume understorey on decomposition and nutrient content of eucalypt forest litter

Summary In Jarrah (Eucalyptus marginata Donn ex Sm.) forest of south-western Australia dense germination and regeneration of the native legumeAcacia Pulchella R. Br. can occur following moderate to high intensity fire. The effect of this legume understorey on rate of decomposition and change in nutrient content ofE. marginata litter was investigated using the mesh bag techniques and by examining four components of forest floor litter representing increasing stages of decomposition. E. marginata leaf litter confined in mesh bags lost 37% of its initial dry weight in the first 8 months on the forest floor and 44% of its initial dry weight after 20 months. During this period weight loss was similar for leaf litter located in forest without legume understorey and for leaf litter placed under dense stands ofA. pulchella. MixingA. pulchella litter withE. marginata litter had no significant effect on rate ofE. marginata litter breakdown. The presence of understorey vegetation had a marked effect on chemical composition of decomposingE. marginata leaves. After 8 and 20 months exposure on the forest floor, leaf litter in mesh bags placed underA. pulchella understorey had significantly (P<0.001) higher concentration and contained significantly (P<0.001) greater amounts of N, P, K, S, Ca and Mg than leaf litter placed in areas without legume understorey. This effect was particularly marked for N and P. In forest without legume understorey the amounts of these two nutrients inE. marginata leaf litter changed little during the first 20 months of decomposition, but forE. marginata leaf litter in mesh bags underA. pulchella there were absolute gains of up to 68% in the amount of N and 109% in the amount of P during this period. This represents accumulation of N and P from sources outside the litter bags. The concentration of N, P, S, Ca and Mg were higher at each of the four stages of decomposition in eucalypt leaf litter collected from the forest floor beneathA. pulchella compared to eucalypt leaf litter collected in forest without understorey. Concentrations of N, P and S increased with stage of decomposition. Levels of these three nutrients in eucalypt litter from under the legume were 1.5 to 2.9 fold higher than in the same component of litter from forest without understorey. The effect of legume understorey on nutrient concentrations in the forest floor and on Cielement ratios in decomposing litter is discussed in relation to long term rates of litter breakdown and net mineralisation of litter nutrients.     

Egg predation in forest bird communities on islands and mainland

Summary To examine if differences in egg predation rates could explain differences in bird community composition, egg predation was studied in two years on small islands in a South Swedish lake and on the nearby mainland using both natural and artificial nests.In plots with similar vegetation, the combined density of ground- and tree-nesting bird species did not differ between the islands and the mainland. Egg predation rates were similar on islands and the mainland for natural Turdus nests in two years, and for artificial Turdus and Phylloscopus nests. Unmarked and unvisited experimental nests suffered similar rate of egg predation as marked and visited nests. Egg predation rates were higher on natural nests when artificial nests were also put out, increasing the total nest density. Initial egg predation rates in artificial nests were also higher than later when nest density had decreased by 75%.The egg predators involved differed for artificial Phylloscopus nests between the islands and the mainland. Small mammals were apparently responsible for 29% of the predation on the mainland, but none on the islands. Artificial Turdus nests near crow nests suffered from a higher egg predation rate than nests further away from crow nests. Daily survival rates of Turdus nests increased from the laying to the incubation and further to the fledging state.Egg predation can not explain differences in bird community composition between islands and mainland in the present case.     

Measuring the host preference of parasites: An aid in evaluating biotypes ofAnagrus epos [Hym.: Mymaridae]

Populations ofAnagrus epos Girault are being imported and evaluated for the biological control of variegated grape leafhopper,Erythroneura variabilis Beamer, a newly introduced pest of grapes in central California. To distinguish different biotypes, a field test was developed that quantifies host preference.E. epos collected from geographically isolated regions of western United States and northern Mexico showed different levels of preference towards the variegated leafhopper and its close relative the grape leafhopper,Erythroneura elegantula Osborn, a native to central California. Host preference values may aid in the identification of relased biotypes using leafhopper eggs on leaves sampled from release sites.      

The biology of pygmy whitefish, Prosopium coulterii, in a closed sub-boreal lake: spatial distribution and diel movements

Using gillnets and trap nets, we examined the spatial distribution, diel movements, and environmental tolerances of pygmy whitefish, Prosopium coulterii, in a small boreal lake in north-central British Columbia. Most gillnets were set below the thermocline but we also fished a shore net in the littoral zone. During the ice-free season (May to November) there was a strong diel onshore–offshore movement: during the day pygmy whitefish were offshore and below the thermocline (water temperatures of 4–6°C) but at night they were inshore and above the thermocline (water temperatures of 12–18°C). This onshore–offshore movement occurred close to the bottom and, regardless of where they were caught, most fish were <4 m off the bottom. Oxygen concentrations in most of the hypolimnion dropped to <5.0 mg l⁻¹ in June and by late August to <1.0 mg l⁻¹; indicating pygmy whitefish can tolerate low oxygen conditions. The catch of pygmy whitefish in gillnets set below the thermocline was highly skewed: 53% of the nets were empty, 37% caught 18 or less fish, and 10% caught 70% of the total catch (742 fish). Trap nets produced similarly skewed results: most trap net sets caught no pygmy whitefish but one set caught over 2,000 individuals. Our catch data suggest that in Dina Lake #1 some pygmy whitefish aggregate.     

Life-table study of Anagrus atomus, an egg parasitoid of the green leafhopper Empoasca decipiens, at four different temperatures

The objective of our study was to assess thepotential of the egg parasitoid Anagrusatomus L. (Hymenoptera: Mymaridae) for controlof the greenhouse leafhopper Empoascadecipiens Paoli (Homoptera: Cicadellidae). Theegg-adult development time, survivorship andreproduction of A. atomus were evaluatedat four constant temperatures (16, 20, 24 and28°C). Developmental time ranged from33.6 days at 16°C to 13.3 days at 28°C. Based on a linear regression ofdevelopment rate on temperature the lowerthreshold was estimated at 8.39°C. Anagrus atomus required 263.2 degree-days tocomplete its development from egg to adult. Theegg-adult survival rate and the sex ratio weresignificantly lower at 28°C than at theother three temperatures tested. The intrinsicrate of increase (r _m) variedsignificantly between all four temperatures.The potential of A. atomus to attackdifferent host ages was additionallyinvestigated. Host eggs were parasitizedthroughout their development but rate ofparasitism was reduced in host eggs older thansix days. The number of eggs parasitized waspositively density dependent but the rate ofparasitism decreased with increasing hostdensity. A maximum rate of parasitism of 62.5%was recorded. The potential impact of the eggparasitoid on the population dynamics of E. decipiens is discussed.     

Within‐year movements and site fidelity of Purple Sandpipers during the nonbreeding season

ABSTRACT Although within‐year site fidelity to specific wintering sites allows shorebirds to use prior knowledge of resources and microhabitats, such fidelity may also make populations more vulnerable to extirpation in the event of increased predation pressure, habitat loss, or disturbance. In the eastern Atlantic, Purple Sandpipers (Calidris maritima) have been found to be highly faithful to specific sites in wintering areas. However, little is known about the use of wintering areas by these sandpipers along the coast of Maine. We quantified movements of 60 radio‐marked Purple Sandpipers in a bay near the mainland and on an offshore cluster of islands along the mid‐coast of Maine during two winters (2005–2006 and 2006–2007). Birds marked in early‐ and mid‐December remained until spring migration, with no evidence of onward migration. Mean maximum distances moved did not differ significantly between either males (8.6 ± 1.0 [SE] km; N= 30) and females (7.4 ± 0.8 km; N= 30) or juveniles (9.9 ± 1.6 km; N= 9) and adults (7.8 ± 1.1 km; N= 26). We also detected no monthly (January–May) differences in maximum distances moved. Sixty percent of marked individuals moved ≤5 km between the two most distant relocations and no birds moved >25 km during the 2‐ to 4‐month tracking period. We attribute the high site fidelity primarily to the plentiful prey base in the study area. During a 2‐d period with severe cold, feeding areas at locations protected from wave action became encased in ice and birds at these locations moved up to 10 km offshore to sites with less ice. Species with strong site fidelity, like wintering Purple Sandpipers, may be at higher risk in the event of large‐scale changes in their food base, increased predation pressure, habitat loss, or disturbance. However, the short‐distance movements made when intertidal feeding areas became encased in ice suggest that Purple Sandpipers could potentially move greater distances in response to changing conditions in their wintering areas.     

Prevalence of Nosema sp. (Microsporidia: Nosematidae) during an outbreak of the jack pine budworm in Ontario

Microsporidia are believed to play little or no role in outbreaks of the jack pine budworm, Choristoneura pinuspinus Freeman (Lepidoptrera: Tortricidae), because the short duration (2–4 years) of those outbreaks may not permit significant build-up of the pathogen. We conducted the first survey of Nosema sp. (Microsporidia: Nosematidae) over the course of a recent jack pine budworm outbreak in Ontario. Between 2004 and 2010 the outbreak defoliated a cumulative total of 1.78 million ha. Microscopic examination of ∼15,000 overwintering larvae collected over 6 years in sites with densities of 3 larvae per branch or more revealed widespread occurrence of Nosema at generally high infection intensities. The pathogen was present in 69.5% of the 518 plots that were monitored. Prevalence of infection was generally low (below 40% in 84% of plots with infected larvae) but reached high levels (80–95%) locally and increased rapidly in most infestations within 1–2 years of onset. We hypothesize that the habit of early-instar larvae to feed on developing male flowers (pollen cones) after spring emergence is critical in allowing rapid build-up of Nosema by increasing efficiency of horizontal transmission (higher density of both infected larvae and egested spores). Nosema infection may contribute to the complexity of jack pine budworm outbreak patterns by affecting egg recruitment and early-instar survival at the stand level in concert with known effects of budworm-induced reductions in pollen cone production on those processes.     

Releases of Trichogramma platneri (Hymenoptera: Trichogrammatidae) in Apple Orchards under a Sterile Codling Moth Release Program

Based on the premise that augmented host numbers may help multiply and support parasitoid populations, the egg parasitoid Trichogramma platneri Nargarkatti was released in apple orchards which were participating in a sterile codling moth, Cydia pomonella (L.), release program. Nonviable eggs resulting from matings involving at least one sterile codling moth partner can be successfully parasitized by T. platneri. Grain moth-reared, as well as codling moth-reared, T. platneri were released either in the spring or in the spring and summer oviposition period(s) of wild codling moth. Sentinel codling moth eggs were hung weekly, for 3-day periods, from May until September to determine fluctuations in T. platneri populations both during and between releases. Low numbers of wild or nonviable codling moth eggs or other susceptible host eggs resulted in sufficient eggs to maintain low spring- or summer-introduced T. platneri populations. T. platneri reduced codling moth damage in trees in which the Trichogramma were released.     

Recolonization and possible recovery of Burrowing Mayflies (Ephemeroptera: Ephemeridae: Hexagenia spp.) in Lake Erie of the Laurentian Great Lakes

Burrowing mayflies of the genus Hexagenia spp. were widely distributed (ca. 80% of sites) and abundant (ca. 160 nymphs/m²) in the western basin of Lake Erie of the Laurentian Great Lakes in 1929–1930, prior to a period of anoxia in the mid 1950s. Nymphs were absent or rare in the basin between 1961 and 1973–1975. In 1979–1991, nymphs were infrequently found (13–46% of sites) in low abundance (3–40 nymphs/m²) near shore (<7.5 km from shore), but were absent or rare offshore (0–7% of sites at 0–1 nymphs/m²). Increased abundance occurred offshore between 1991 (0% of sites) and 1993 (52% of sites at 7/m²). Annual sampling, beginning in 1995, indicates that nymphs increased in both nearshore and offshore waters. By 1997, nymphs were found throughout the lake (88% of sites) at a mean density 40-fold greater (392/m²) than that observed in 1993 (11/m²). In 1998, the distribution of nymphs remained the same as 1997 (88% of sites) but density declined 3-fold (392 to 134/m²). These data indicate that mayflies have recolonized sediments of western Lake Erie and that their abundance may be similar to levels observed before their disappearance in the mid 1950s. However, prior to the mid 1950s, densities were greater in offshore than nearshore waters, but between 1979 and 1998 greater densities occurred near shore than offshore. In addition, there were two areas in the 1990s where low densities consistently occurred. Therefore, recovery of nymphs in western Lake Erie may not have been complete in 1998. At present we do not know the cause for the sudden recolonization of nymphs in large portions of western Lake Erie. Undoubtedly, pollution-abatement programs contributed to improved conditions that would have ultimately led to mayfly recovery in the future. However, the explosive growth of the exotic zebra mussel, Dreissena polymorpha, undoubtedly diverted plankton foods to bottom substrates which could have increased the speed at which Hexagenia spp. nymphs recolonized sediments in western Lake Erie in the 1990s.