Localized outbreak of a willow leaf beetle: plant vigor or natural enemies?

The possible roles of plant quality (vigor) and natural enemies in the development of a localized out-break of the leaf beetle Galerucella lineola (Coleoptera: Chrysomelidae) in a stand of Salix cinerea were investigated. Caged and uncaged larvae on six bushes in the outbreak area were compared with caged and uncaged larvae on six bushes in an adjacent nonoutbreak area in terms of performance. In 1997, when the studies were performed, the natural density of the insect (beetles plus eggs) was six times higher in the outbreak area compared with the nonoutbreak area. Even though the vigor (measured as shoot length) of bushes in the outbreak was 72% higher than that of bushes in the nonoutbreak area, we found no difference between areas in the performance (survival, developmental time, pupal weight) of caged larvae or in the willingness of caged females to lay eggs. Among larvae exposed to natural enemies, the disappearance rate was significantly higher in the nonoutbreak area. The density of generalist predators was significantly higher in the nonoutbreak than in the outbreak area. We conclude that differences in plant quality, despite the observed difference in plant vigor, could not explain the observed difference in beetle density between areas. Lower predation pressure in the outbreak area could, however, not be excluded as a possible reason for the higher density of leaf beetles in this area. Received: October 18, 1999 / Accepted: February 4, 2000     

Impacts of defoliation by tussock moths (Orgyia vetusta) on the growth and reproduction of bush lupine (Lupinus arboreus)

Abstract.

• 1 Outbreaks of tussock moths (Orygia verusta Bdv.) on bush lupines (Lupinus arboreus Sims) may be very intense, although spatially localized, and may last >10 years. To understand better how such outbreaks may persist, we defoliated lupine bushes using variable realistic numbers of tussock moth larvae, and measured the immediate and delayed impacts of defoliation on the growth and seed production of lupine bushes.
• 2 Immediate effects of defoliation included the production of new leaves that were significantly smaller in diameter and biomass, and lower in water content, than the new leaves of undamaged bushes. Also, seed output was reduced by up to 80% on heavily damaged bushes.
• 3 In the longer term, surviving bushes recovered remarkably well, despite having received the full range of possible (survivable) insect densities. After 1 or 2 years’rapid growth, surviving bushes were not affected in terms of height, basal stem diameter or volume by their previous defoliation. However, there was a significant tendency for juvenile bushes to produce more seeds the more heavily they had been attacked the previous year.
• 4 The ability of the host plant to recover, in terms of its biomass, helps to explain the sustained nature of the insect outbreak.

     

Evidence that predator satiation may restrict the spatial spread of a tussock moth (Orgyia vetusta) outbreak

Western tussock moths (Orgyia vetusta Bdv., Lymantriidae) infest one stand of bush lupine (Lupinus arboreus Sims, Fabaceae) heavily and several other stands very lightly at the Bodega Marine Reserve (Sonoma Co., Calif., USA). We found that the disappearance rates of experimentally placed larvae and pupae were consistently lower in the outbreak area than in non-outbreak areas. For pupae but not larvae, this difference was removed by using tanglefoot to repel nonflying predators. However, the major nonflying predator of pupae, the ant Formica lasioides, was no more abundant in non-outbreak areas than in the outbreak area. We found inverse density-dependence in the rate of attack by F. lasioides on experimental pupae, suggesting this generalist predator is satiated within the outbreak area, but preys more heavily on the moth where the moth is sparse.     

Origin of an insect outbreak: escape in space or time from natural enemies?

Initiation of insect outbreaks is poorly understood, and may involve sporadic events that temporarily release insect populations from predation or parasitism. While studying a declining outbreak of the western tussock moth (Orgyia vetusta) on bush lupine (Lupinus arboreus), we witnessed the onset of a new tussock moth outbreak, separated by 1,000 m in space and 2 months in phenological timing from the original population. This new population underwent explosive growth for 2 years and then collapsed because of a massive die-off of lupines. We tested whether during its growth phase, this new outbreak benefited by escaping in either space or time from the natural enemies attacking the original population. In experimental populations on single bushes, we compared predation and parasitism at the sites of the new and the old outbreak. At the site of the old outbreak, we compared predation and parasitism early and late in the season. Parasitism was significantly lower and population growth significantly higher at the new outbreak site than the old one. Neither seasonal timing, predator exclusion, nor their interaction significantly affected survival at either site. Thus the new outbreak appeared to escape in space from parasitism. These results corroborate our previous experimental findings, which suggest that as predicted by theory, the interaction between the tussock moth and its parasitoids can produce large-scale spatial patterning in population densities.     

High mortality,fluctuation in numbers,and heavy subterranean insect herbivory in bush lupine,Lupinus arboreus

Sporadic patchy die-off of bush lupine, Lupinus arboreus, has long been known. We describe in detail a series of these incidents on the central California coast, based upon observational and comparative evidence. Stands of thousands of plants die, while nearby mature plants live on. In some sites, repeated die-off followed by regeneration from the seed bank has led to the cover and density of this woody, perennial plant fluctuating widely over the 40 year period for which records exist. Root damage by caterpillars of the ghost moth or swift Hepialus californicus (Lepidoptera, Hepialidae) is a major cause of individual bush death and a probable cause of die-off of stands of lupine. Hidden from view underground, a few of these insects readily kill a juvenile or young mature plant by girdling and reaming-out roots. The mass mortality of L. arboreus that we observed involved heavy root damage by these caterpillars in evenaged stands of plants in their first (1.5-year-old) or second (2.5-year-old) flowering season. The injured plants set seed before dying. Older, larger bush lupines better withstood root damage. In plants aged 3 or more years, damage and mortality were correlated with the intensity of ghost moth caterpillars in the roots. At the highest intensity (mean = 37.5, maximum = 62 caterpillars/root), a stand of large, old L. arboreus suffered 41% mortality; 45% of root cambium (median value) was destroyed by feeding caterpillars. Mass death of mature L. arboreus was not correlated with folivory, and leaf damage ranged from nil to moderate in instances of die-off. The western tussock moth, Orgyia vetusta, accounted for the highest levels of folivory, but this insect was rare when die-offs occurred. The lowest lupine mortality rates in our study occurred where tussock caterpillar intensities were high and where plants were repeatedly defoliated by this insect. However, experimental defoliation by high, but realistic, intensities of tussock moth caterpillars resulted in some mortality of mature bushes, and the combined effects of leaf and root herbivory have yet to be assessed. In its natural range on the California coast, bush lupine has several additional species of insect herbivores that can be locally abundant and injurious to the plant, although none is associated with die-off. Subterranean natural enemies of ghost moth caterpillars may play a role in the patchy waxing and waning of this shrub. Locally, a new species of entomophagous nematode (Heterorhabditis sp.) cause high mortality in the soil, before ghost moth caterpillars have entered the root. This natural enemy may thus afford lupines protection from heavy underground herbivory.     

Lack of strong induced or maternal effects in tussock moths (Orgyia vetusta) on bush lupine (Lupinus arboreus)

Both induced and maternal effects may create delayed negative feedback on the population growth of herbivorous insects. I tested for these effects in a chronically dense population of tussock moths (Orgyia vetusta) feeding on bush lupines (Lupinus arboreus). Experimental bushes received different realistic levels of defoliation by tussock moths in the preceding year, and experimental moth larvae came from mothers that had experienced either high or low levels of crowding as larvae in the previous year. Weight of female moths at pupation was not affected by prior foliar damage, and showed only a weak (12%) effect of maternal crowding. Rates of early larval disappearance, from aerial dispersal (ballooning) or other causes, were unaffected by either foliar damage or maternal crowding, and were very low except when larvae were placed on dead bushes. These results help to explain why the tussock moth population has maintained high densities at the study site for >10 years.     

Interspecific competition and insect herbivory reduce bush lupine (Lupinus arboreus ) seedling survival

Seedlings suffer high mortality in most plant populations, with both competition and herbivory proposed as being important mechanisms causing seedling death. The relative strength of these factors, however, is often unknown. Here I ask how interspecific competition for light and insect herbivory jointly affect seedling survival of bush lupine (Lupinus arboreus), a native shrub common to coastal California. Bush lupine seedlings germinate in grasslands during winter, and throughout spring potentially compete for light with surrounding fast-growing annual grasses. By early summer, after grasses have died, seedlings can be defoliated by a locally abundant caterpillar, the western tussock moth (Orgyia vetusta). I examined the relative importance of competition and insect herbivory on seedling survival in two separate experiments. First, I compared seedling mortality in plots either exposed to or protected from tussock moth larvae. Plants were protected from herbivory by the judicious use of insecticide; control plants were sprayed with water. Tussock moth herbivory resulted in significantly greater (31%) seedling mortality. To determine the effects of interspecific competition for light on seedling survival, I manipulated the density of grass surrounding lupine seedlings. I removed all vegetation surrounding some individuals, and left intact vegetation surrounding others. Reducing competition resulted in a 32% increase in seedling survival from February to June, as well as changes in seedling growth. To determine whether there are interactive effects of competition and herbivory on seedling survival, I enclosed tussock moth larvae on half of all surviving seedlings within each of the two prior competition treatments, comparing growth and survival of defoliated and undefoliated seedlings. Defoliation in June led to an additional 50% mortality for individuals that had grown with competitors through spring, and a 53% additional mortality for seedlings that grew without competitors through spring. Thus, although competition and herbivory both caused substantial seedling mortality, there was no statistical interaction between these factors. Competition-free plants were not less vulnerable to herbivory than plants that previously grew with competitors. Taken together, these experiments indicate that competition and herbivory are both important sources of mortality for bush lupine seedlings. Received: 4 April 1996 / Accepted: 5 November 1996     

Facilitation of tiger moths by outbreaking tussock moths that share the same host plants

1.?Ecologists have argued about the commonness and strength of interspecific competition between insect herbivores, but facilitation between herbivores has received much less consideration. We previously found that when two species of folivorous caterpillars co-occurred on a shared host plant, feeding by early season tiger moth caterpillars reduced the growth and reproduction of later season tussock caterpillars. However, densities of tussock caterpillars in summer were positively correlated with densities of tiger moth caterpillars the following spring. 2.?In this study, we experimentally manipulated numbers of feeding tussock caterpillars and found that they facilitated tiger moth caterpillars. 3.?The depth of the litter layer beneath host lupine bushes was positively correlated with the number of tussock caterpillars feeding on each bush. Experimental additions of litter beneath lupine canopies during summer resulted in increased numbers of tiger moth caterpillars in the following spring, indicating a causal role of litter. Litter potentially provides food, habitat and protection from desiccation and predation. We failed to find evidence that tussock caterpillars facilitated tiger moth caterpillars by mechanisms independent of litter. 4.?Our study demonstrates that facilitation may operate between insect herbivores, across life-stages through indirect interactions that are non-trophic. Facilitation operated by a novel mechanism, the accumulation of litter which was a by-product of feeding by one species was valuable to a second species. Facilitation persisted in time and space far beyond the creation of litter by tussock caterpillars which should be considered important ecosystem engineers from the point of view of tiger moths. Facilitations that involve habitat modification may generally connect species that do not interact directly or trophically, and have not previously been considered to affect one another.     

Density-dependent foraging behaviors in a parasitoid lead to density-dependent parasitism of its host

Empirical studies of spatial heterogeneity in parasitism by insect parasitoids have focused largely on patterns, while the many possible underlying mechanisms have been little studied in the field. We conducted experimental and observational studies on Tachinomyia similis (Diptera: Tachinidae) attacking western tussock moths (Orgyia vetusta; Lepidoptera: Lymantriidae) on lupine bushes at Bodega Bay, Calif., USA. We examined several foraging behaviors that have been hypothesized to create density-dependent variation in parasitism rates, including spatial aggregation of parasitoids to high host density, mutual interference among searching parasitoids and decelerating functional responses of the parasitoid. At the spatial scale of individual bushes, we detected both aggregation to a high density and a decelerating functional response. The resulting spatial pattern of parasitism was best fit by two models; one included an effect of parasitoid aggregation and the other included an effect of aggregation and a decelerating functional response. Most of the variation in parasitism was not correlated with density of O. vetusta.     

Persistent, localized outbreaks in the western tussock moth Orgyia vetusta: the roles of resource quality, predation and poor dispersal

1. On stands of the abundant perennial lupins Lupinus arboreus and L. chamissonis on the California coast, the western tussock moth Orgyia vetusta Bdv. occurs as only a few localized populations, which are locally dense enough to defoliate the host plants and yet do not appear to spread.
2. To determine what limits the spatial distribution of this insect, and therefore its global abundance, experiments were performed at three sites, examining the roles of host-plant quality, predator abundance, rates of predation and dispersal.
3. At no site did host quality, predator abundance or rates of predation vary spatially in a way that would explain the location of the outbreak. However, adult female tussock moths are flightless, and rates of long-distance aerial dispersal ('ballooning') by first-instar larvae were very low at all sites. Poor dispersal is the only consistent explanation for the limited distribution of the tussock moth.     

The influence of natural enemies on brood production inIps typographus (Col. scolytidae) with special reference to egg-laying and predation byThanasimus formicarius (Col.: Cleridae)

The predatorThanasimus formicarius (L.) (Coleoptera, Cleridae) and its preyIps typographus (L.) (Coleoptera, Scolytidae) were studied in the laboratory and the field. In the laboratory, 11T. formicarius laid 71–132 eggs (mean=162) during 66–123 days. During this time they ate 66–132I. typographus adults per pair (male +female). The number of eggs laid per female was not correlated with life span or the number ofIps eaten. In the field, predation byT. formicarius larvae and other natural enemies onI. typographus brood was studied in the last year of an outbreak. Caged and uncaged spruce bolts attacked byI. typographus were used, and pairs ofT. formicarius were released in the cages. The treatments were: uncaged bolts, caged bolts withoutT. formicarius, caged bolts with 4T. formicarius pairs, and caged bolts with 8T. formicarius pairs. The productivity ofI. typographus was highest in the caged bolts withoutT. formicarius (mean=4.5 offspring/female) and lowest in the uncaged bolts (mean=0.9 offspring/female). The density ofI. typographus galleries was similar in the different treatments. Hence, the variation in productivity between treatments could not have been due to differences in the levels of intraspecific competition. There was no difference in bark beetle productivity or density ofT. formicarius larvae between bolts with 4 pairs ofT. formicarius and bolts with 8 pairs (mean=2.5 offspring/female). This indicates that some kind of interference occurred betweenT. formicarius individuals (e.g. cannibalism) and that a maximum level of predation was reached. Predation by larvae ofMedetera spp. (Diptera, Dolichopodidae),Thanasimus spp. and other beetles, and parasitism by wasps (Hymenoptera, Pteromalidae) probably caused the low productivity in the uncaged bolts.      

Intraspecific competition and subterranean herbivory: individual and interactive effects on bush lupine

High mortality of plants growing in dense monospecific stands (i.e. self-thinning) usually results from intense intraspecific competition. However, inconspicuous below-ground insect herbivory might be a potent but overlooked source of mortality within dense stands of plants, particularly if crowding limits a plant's ability to compensate for herbivore damage. Here I ask how high conspecific density influences a plant's ability to cope with heavy below-ground insect herbivory.
I manipulated conspecific density and exposure to an abundant root-borer, the ghost moth ( Hepialus californicus ), and examined the impacts on the fecundity, growth, and survival of bush lupine ( Lupinus arboreus ), a fast-growing shrub that grows in dense monospecific stands in coastal grasslands. Both herbivory and intraspecific competition affected seed production, size, and mortality of bush lupine over the two years of the experiment. Plants consistently produced fewer seeds when growing at high versus low density and ghost moth herbivory also significantly reduced seed production. The negative effects of herbivory on plant fecundity were similar, regardless of plant density. In contrast, plant survival was affected by both competition, herbivory, and the interaction of these factors. In high density plots, plant survival was uniformly low (averaging 0.45–0.50); plants exposed to herbivores died from heavy herbivory, and plants protected from herbivores died due to intense intraspecific competition that compensated for losses due to herbivory. In low density plots, ghost moth herbivory similarly reduced lupine survival, from an average survival probability of 0.94 in plots protected from these herbivores to 0.55 in plots exposed to herbivory. Thus, results show that regardless of plant density, below-ground herbivory can be a potent source of mortality.     

Entomopathogenic nematodes: natural enemies of root-feeding caterpillars on bush lupine

A new species of soil-dwelling entomopathogenic nematode Heterorhabditis hepialus killed up to 100% (mean=72%) of root-boring caterpillars of a ghost moth Hepialus californicus in coastal shrub lands. When unchecked, ghost moth caterpillars killed bush lupine, Lupinus arboreus. Here we describe this strange food chain. Although unappreciated by ecologists, entomopathogenic nematodes are widespread and probably one of the most important groups of natural enemies for underground insects. The free-living infective juvenile (IJ) of entomopathogenic nematodes searches for host insects in the soil. A single IJ can kill a host, although several often invade together. After entering the host through a spiracle or other orifice, the IJ regurgitates its symbiotic bacterium, Photorhabdus luminescens, which kills the host within 48 h. The bacteria digest the cadaver and provide food for the exponentially growing nematode population inside. The bacteria produce antibiotics and other noxious substances that protect the host cadaver from other microbes in the soil. When the cadaver is exhausted of resources, IJs break the host integument and can disperse. As many as 420,000 IJs can be produced within a large ghost moth caterpillar. Surface soil of the lupine rhizosphere is the primary habitat of IJs of H. hepialus. Attracted to waste gases emitted by insects, the 0.5-mm-long IJs can move 6 cm/day through moist soil. Prevalences of H. hepialus ranged from as high as 78% of rhizospheres in some lupine stands to almost zero in others, but it was absent from no stand at our study site. Field intensities ranged from 0.003 IJs/cm³ of soil to 7.5 IJs/cm³, and correlated roughly with prevalences among sites. Few ghost moth caterpillars (mean=6.7) succeeded in entering lupine roots where prevalence of H. hepialus was highest, and this stand had lowest mortality (0.02) of mature bush lupine. In the three stands with lowest prevalence (mean = 2%) of this nematode, many caterpillars (mean = 38.5) entered roots, and lupine mortality was high (range = 0.41–1.0). Old aerial photographs indicate that the stands with highest recent nematode prevalence have had little or no mass die-off of lupine over the past 40 years. The photos depict repeated die-offs of lupine during the past four decades in stands with lowest recent prevalence of the nematode. This pattern leads us to entertain the hypothesis that the nematode affects vegetation dynamics indirectly through a trophic cascade. Dispersal of entomopathogenic nematodes is little understood. We found that air drying of soil extirpates H. hepialus and speculate that this nematode is dispersed during the wet season in moist soil bits on the exterior of fossorial insects and mammals. H. hepialus colonized some previously unoccupied lupine rhizospheres during the wet winter-spring season and, obversely, became extinct from some rhizosperes as soil dried in summer. Root-feeding insects have only recently been recognized as a force in communities, and the regulation of these important herbivores is still largely an ecological terra incognita. All evidence indicates that entomopathogenic nematodes are found throughout terrestril ecosystems, and we propose that trophic chains similar to those described in this report should not be uncommon.     

Wood Decomposition Following a Perennial Lupine Die-Off: A 3-Year Litterbag Study

Woody debris is a conspicuous feature of many ecosystems and can be a large pool of stored carbon and nutrients. In the California coastal prairie, yellow bush lupines (Lupinus arboreus) experience mass die-offs, producing large quantities of woody detritus. Live lupines are fed upon by the stem-boring caterpillars of the ghost moth, Hepialus californicus, and outbreaks of ghost moths are one factor contributing to lupine die-offs. A common detritivore, the terrestrial isopod Porcellio scaber, frequently inhabits ghost moth tunnels in lupine wood. We used a litterbag experiment to test the hypothesis that H. californicus increases decomposition of woody lupine detritus by facilitating its use by P. scaber. Isopod access to wood was crossed with simulated ghost moth boring to measure the independent and interactive effects of these two arthropods on total mass loss, as well as on carbon, nitrogen, and lignin dynamics. Isopods initially colonized litterbags but were not more abundant on L. arboreus logs that had simulated ghost moth boring than on logs without boring. They were rare in litterbags collected at 12 months or later and had no effect on wood decomposition. Simulated ghost moth boring increased wood decomposition (P = 0.0021), from 50.5 to 55.1% mass loss after 3 years. This effect was likely due to increased surface area for microbial utilization of the wood. Lupine wood had an initial lignin content of 14.70 ± 0.67%, but lignin did not appear to decompose during the 3 years of this study, and by the end of the experiment accounted for 32.6 ± 1.12% of the remaining wood. Neither ghost moth boring nor isopod access affected lignin loss. Lupine wood from a die-off in 2002 was estimated to have contained three times more nitrogen per unit area than the yearly input of annual grass litter. The slow decomposition of lupine wood, however, restricts the rate at which nitrogen is released into the soil and results in the storage of carbon and nutrients in lupine wood for several years following such die-offs.     

Effects of an early-season folivorous moth on the success of a later-season species,mediated by a change in the quality of the shared host,Lupinus arboreus Sims

Summary Larvae of Platyprepia virginalis (Lep.: Arctiidae) and Orgyia vetusta (Lep.: Lymantriidae) feed on the foliage of bush lupine (Lupinus arboreus Sims) at Bodega Bay, California, USA, in February–April and May–July, respectively. Female O. vetusta attained lower pupal weights and produced fewer eggs on branches of L. arboreus which had experimentally received P. virginalis damage earlier in the same year, compared to those reared on control branches.Growth rates of first-instar O. vetusta were lower on undamaged leaves of previously damaged L. arboreus than on leaves of undamaged controls. This was true for lupine bushes damaged by P. virginalis in the field; and also for greenhouse-grown lupine plants damaged by Estigmene acrea (Lep.: Arctiidae) when excised foliage was used in the assay. However, fifth (last) instar female O. vetusta larvae showed no preference for damaged versus undamaged greenhouse-grown L. arboreus in their feeding or choice of a site for pupation.These experiments demonstrate the potential for competition-like interactions between herbivores mediated by changes in quality of the shared host.     

Unexpected spatial patterns in an insect outbreak match a predator diffusion model

An outbreak of the western tussock moth ( Orgyia vetusta) with a sharp spatial boundary has been hypothesized to be the result of an interaction with mobile parasitoids. We report here a striking correspondence between the qualitative prediction of a general model for a mobile predator and a sessile prey and initial observations of the outbreak of western tussock moth. The model makes the counterintuitive prediction that the prey species will have its highest population density at the edge of an outbreak, which corresponds to observations made of the western tussock moth. This is a striking pattern which should be looked for in other population systems.     

Escape in space from enemies: a comparison between stands with and without enhanced densities of the spruce bark beetle

1 Populations of the spruce bark beetle, Ips typographus (L.), are known to grow rapidly in storm‐disturbed stands as a result of relaxation from intraspecific competition. In the present study, it was tested whether a second mechanism, escape in space from natural enemies, also contributes to the rapid population increase. 2 The experiment was conducted during the initiation phase of five local outbreaks of I. typographus triggered by a storm‐disturbance in November 1995 in southern Sweden. 3 The impact of natural enemies on the ratio of increase (number of daughters per mother) of I. typographus was compared pairwise between disturbed stands with high numbers of storm‐felled trees and undisturbed stands without wind‐felled trees. 4 Enemy impact was assessed by comparing the ratio of increase in uncaged (exposed to enemies) and caged (protected from enemies) bolts colonized by I. typographus prior to being placed in the stands. The experiment was conducted in the second and third summers after the storm‐felling. 5 Enemy impact was about twice as high in stands without wind‐felled trees compared with in stands with wind‐felled trees in the second summer whereas there was no significant difference between the stand types in the third summer. 6 The result demonstrates that spatial escape from enemies contributes to the rapid population growth of I. typographus after storm‐disturbances.     

Growth and survival of larvae of Thaumetopoea pinivora inside and outside a local outbreak area

1 Outbreaks of herbivorous insects tend to be spatially restricted, possibly because of demographic differences between inside and outside the outbreak area. In some cases, the margin of the outbreak area is distinct, allowing comparisons of adjacent areas that may identify factors leading to such differences in abundance. The northern pine processionary moth Thaumetopoea pinivora presently occurs at outbreak densities within a well‐defined area of approximately 3000 ha on the island of Gotland, south Sweden. We investigated whether cohorts of young larvae (first and second instar) had higher growth rate and survival inside or outside the outbreak area. 2 Group‐feeding appears to promote outbreaks in certain insect groups. Because T. pinivora larvae are highly social, we also compared larval performance between groups of different sizes inside and outside of the outbreak area: ‘small’ (33 eggs/group) and ‘normal’ (100 eggs/group). 3 Averaged over group size, whole colony mortality through the first two instars was two‐fold higher in the non‐outbreak area compared with the outbreak area. Mortality of individual larvae in the surviving colonies, however, did not differ between the two areas. There were only small differences in food quality (toughness, nitrogen content) between the areas, with no detectable effects on larval performance. 4 Larval relative growth rate did not differ between reduced and normal‐sized groups, which is surprising given that growth rate is known to increase with group size in other group‐feeding lepidopterans. 5 Reduced group size negatively affected larval survival, particularly in the outbreak area; by contrast, normal‐sized groups survived equally well in the two areas. Wood ants (Formica spp.) were more common outside the outbreak area, and appeared to be the main cause of colony mortality at low larval density. A different result was observed with regard to per‐capita mortality, which was higher in the outbreak area. We speculate that this could have been due to solitary predators being locally specialized on T. pinivora in the high‐density area.     

Exploring stable pattern formation in models of tussock moth populations

1. The western tussock moth ( Orgyia vetusta ) at the University of California Bodega Marine Reserve (Sonoma County, California, USA) exhibits dense, localized populations in the midst of extensive habitats where variation in host plant quality or predator abundance is unable to explain the restricted extent of the outbreaks.
2. Two primary features suggest that the host patterning is intrinsically generated: (i) female tussock moths are wingless, producing a low effective dispersal distance for the hosts; and (ii) the tussock moth population is attacked by several species of widely dispersing wasp and fly parasitoids.
3. We consider a set of spatially explicit host–parasitoid models to examine whether intrinsically generated patterns are possible within this system. These models include a spatially extended Nicholson–Bailey model to examine general features of pattern formation in host–parasitoid systems, and two system-specific models, an individual-based simulation and a population-level analytic model, to examine the details of this empirical system.
4. Both stable patterning and rapid extinction of the host population are initial-condition dependent outcomes of the general and specific models, implying that an intrinsically generated stable host pattern is feasible within the tussock moth system.
5. Stable patterning is enhanced by a large parasitoid-to-host dispersal ratio, local host resource limitation, and increased parasitism at the host patch's edge.     

A native nitrogen-fixing shrub facilitates weed invasion

Invasions by exotic weedy plants frequently occur in highly disturbed or otherwise anthropogenically altered habitats. Here we present evidence that, within California coastal prairie, invasion also can be facilitated by a native nitrogen-fixing shrub, bush lupine (Lupinus arboreus). Bush lupines fix nitrogen and grow rapidly, fertilizing the sandy soil with nitrogen-rich litter. The dense lupine canopy blocks light, restricting vegetative growth under bushes. Heavy insect herbivory kills lupines, opening exposed nitrogen-rich sites within the plant community. Eventual re-establishment of lupine occurs because of an abundant and long-lived seed bank. Lupine germination, rapid growth, shading and fertilization of sites, and then death after only a few years, results in a mosaic of nutrient-rich sites that are available to invading species. To determine the role of bush lupine death and nitrogen enrichment in community composition, we examined nutrient dynamics and plant community characteristics within a site only recently colonized by lupine, comparing patches where lupines had recently died or were experimentally killed with adjacent areas lacking lupine. In experimentally killed patches, instantaneous pool sizes of exchangeable ammonium and nitrate nitrogen were higher than in adjacent sites free of lupine. Seedlings of the introduced grass Bromus diandrus accumulated 48% greater root biomass and 93% more shoot biomass when grown in a greenhouse in soil collected under experimentally killed lupines compared to B. diandrus seedlings grown in soil collected at least 1 m away from lupines. At the end of the spring growing season, total above-ground live plant biomass was more than twice as great in dead lupine patches as in the adjacent lupine-free grassland, but dead lupine patches contained 47% fewer plant species and 57% fewer native species. Sites where lupines have repeatedly died and reestablished during recent decades support an interstitial grassland community high in productivity but low in diversity, composed of mostly weedy introduced annual plants. In contrast, at a site only recently colonized by bush lupines, the interstitial grassland consists of a less productive but more diverse set of native and introduced species. We suggest that repeated bouts of lupine germination, establishment, and death can convert a rich native plant community into a less diverse collection of introduced weeds.