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.     

Risk assessment of an exotic biocontrol agent: Phytoseiulus persimilis (Acari: Phytoseiidae) does not establish in rainforest in southeast Queensland

Abstract The Chilean predatory mite, Phytoseiulus persimilis Athias-Henriot, appeared in Australia in 1978 soon after being introduced into New Zealand as a specialized biological control agent of spider mites. It is known to be naturalized in agricultural habitats in southeast Queensland, Australia, although nothing is known about its distribution in native ecosystems. In order to determine whether P. persimilis is able to invade subtropical rainforest, we placed potted bean plants infested with its preferred prey, the two-spotted mite (Tetranychus urticae C.L. Koch), at 50 m intervals for 200 m on either side of the rainforest-field ecotone at four sites in southeast Queensland. Two, 4 and 6 weeks after placement, five leaves were sampled from each pot and any phytoseiid mites present were identified. The initial experiment took place in the spring and was repeated in summer and in autumn of 1997. At all four sites and in all three seasons P. persimilis rapidly colonized all of the pots in fields. In the rainforest, however, some pots were never colonized and significant populations of the predator developed only in the summer, and then only at the first stations, 50 m into the forest. These results suggest that even when its preferred prey is present, subtropical rainforest is not an appropriate habitat for P. persimilis. In addition, we reviewed extensive collections of phytoseiid mites from native forests and synanthropic habitats in Australia and found P. persimilis records only from fields, glasshouses, gardens, weeds, roadsides and similar disturbed habitats dominated by introduced plants, again suggesting that this biocontrol agent is not a rainforest invader.     

Spatially structured herbivory and primary succession at Mount St Helens: field surveys and experimental growth studies suggest a role for nutrients

Abstract. 1. The 1980 eruption of Mount St Helens (Washington, U.S.A.) created a 60‐km² region of primary successional habitat. Since colonising in 1981, the spatial spread of the legume Lupinus lepidus at Mount St Helens, Washington, U.S.A., has afforded intriguing opportunities to study the effect of trophic dynamics on primary succession. 2. Insect herbivory on this lupine has exhibited striking spatial structure for over a decade, with inverse density‐dependent damage patterns occurring over both small (10–100 m) and large (1–10 km) spatial scales. 3. Hypothesising that lupine nutritional chemistry might underlie the spatial patterns in herbivory, the distribution of elemental macronutrients (nitrogen, phosphorus) across the landscape was characterised. 4. Samples of soil and lupine tissue (roots and leaves) were collected from sites along both local and regional gradients in lupine density. On both large and small spatial scales, lupine leaves from low‐density conditions were significantly more nutrient rich. 5. In addition, in a laboratory growth study native lepidopteran herbivores that specialise on lupines (Gelechiidae: Filatima sp.) performed better when fed leaves from low‐density, high‐nutrient lupines than on diets of low‐nutrient lupine leaves from high‐density areas a few metres away. 6. These data suggest that spatial heterogeneity in lupine nutrient chemistry may underlie the remarkable herbivory gradients witnessed at Mount St Helens.     

Host specificity of the ragwort flea beetle Longitarsus jacobaeae (Waterhouse) (Coleoptera:Chrysomelidae)

Abstract

The ragwort flea beetle, Longitarsus jacobaeae, was tested for host specificity against representative species of native New Zealand Senecio. Adult feeding and oviposition tests were carried out under quarantine with and without a choice of host plants. Larval development was assessed using potted plants. It was concluded that L. jacobaeae is highly specific to Senecio jacobaea and that it is extremely unlikely to be damaging to native New Zealand Senecio species.     

A Molecular and Fitness Evaluation of Commercially Available versus Locally Collected Blue Lupine Lupinus perennis L. Seeds for Use in Ecosystem Restoration Efforts

Dependence on wild seed sources is often impractical for large‐scale habitat restoration programs. Reliance on commercial seed supplies of unknown provenance and fitness is thereby warranted. Little consideration has been given, however, to how the large volumes of seed required should be sourced. We evaluated commercial and locally collected seed sources for potential use in a New York State‐based, landscape‐scale program for restoring blue lupine Lupinus perennis. Through analysis of microsatellite markers we determined that “native” lupine designations by some commercial suppliers were in fact interspecific hybrids and therefore unreliable; at least two commercial sources, however, were genetically as close to native New York populations as native New York populations were to one other. Common garden experiments revealed that seed source influenced first‐year overwintering survival and subsequent height growth of surviving plants; seed sources more closely related genetically to native New York populations survived better and produced more stems per individual in the field in the area targeted for restoration. We conclude that (1) commercial suppliers often but not always offer reliably characterized seed sources of sufficient genetic similarity to native populations to warrant their use in restoration projects and (2) genetic affinity of potential seed stock to native populations is positively related to its fitness in the environment targeted for restoration.     

Invasion of the gall mite Aceria genistae (Acari: Eriophyidae), a natural enemy of the invasive weed Cytisus scoparius,into California,U.S.A. and predictions for climate suitability in other regions using ecological niche modelling

Scotch broom (Cytisus scoparius (L.) Link) is a European shrub that has naturalised in several countries worldwide and is recognised as an invasive weed in much of western North America. The mite Aceria genistae (Nalepa) is a coevolved, gall-inducing herbivore associated with Scotch broom in its native range and has been intentionally introduced as a classical weed biological control agent of C. scoparius in Australia and New Zealand. An adventive, never intentionally introduced, population of A. genistae was discovered in Washington and Oregon, U.S.A. in 2005. Surveys for A. genistae in California resulted in the discovery of the gall mite in 11 counties, with a widely scattered distribution. Molecular and morphological assessments confirm the mites collected from galls in California are A. genistae. Whether natural or anthropogenic, the estimated rate of long range dispersal for A. genistae from Washington or Oregon to California ranges from 39 to 62?km/yr. Niche model predictions indicate that A. genistae will continue to expand its distribution throughout much of the Scotch broom-invaded lands of California but areas supporting the weed in the Eastern U.S.A. appear less suitable. Modelling evidence also indicates that portions of Chile and Argentina are suitable for colonisation by A. genistae, also suggesting that expansion of the mite is possible in areas of Tasmania, southeastern Australia, and New Zealand where the mite was released. The environmental safety of A. genistae in relation to non-target plants and the influence of herbivory on Scotch broom fitness are discussed.     

Reintroduction of Wild Lupine (Lupinus perennis L.) Depends on Variation in Canopy,Vegetation, and Litter Cover

We experimentally examined the effects of canopy, vegetation, and leaf litter cover on the demography of Wild lupines (Lupinus perennis) in a central North American oak savanna spanning 9 years. We also compared the distribution of Wild lupine across the landscape to results predicted by the demographic experiments. With less canopy cover, soil temperatures were warmer and seedlings emerged earlier. Seedling survival increased 14% with each additional leaf grown. Seedling survival was four times greater in openings and partial shade than in dense shade. Seedling survival was also influenced by interactions between canopy cover and vegetation cover, between canopy cover and leaf litter, and among canopy cover, vegetation cover, and litter cover. In openings, seedlings had higher survival when vegetative cover was present, suggesting a positive shading effect on survival, but with greater canopy cover vegetative cover reduced survival. Seedling survival was greater for plants that experienced herbivory, a result that was probably related to plant size and quality rather than having been eaten. Survival of lupines to 9 years after seed planting was greatest in the partial shade, moderate in openings, and least in dense shade. Wild lupine cover across the landscape was greatest when litter cover was low and canopy cover and ground layer cover were moderate. Reduction of canopy cover by burning or cutting, and reduction of leaf litter by prescribed burning will benefit the reintroduction of Wild lupine by increasing light, reducing litter cover, and creating disturbances; however, the reduction of vegetation cover in openings may hinder lupine reintroduction.     

Risk posed by the invasive defoliator Uraba lugens to New Zealand native flora

• 1 The risk posed to New Zealand native flora by the recently‐established pest of Australian origin Uraba lugens Walker (Lepidoptera: Nolidae) (gum leaf skeletonizer) was assessed. Weed biological control host range testing methods were applied to identify those New Zealand plant species potentially at risk. Native plants tested were primarily in the Myrtaceae, the family that contains all the Australian hosts of U. lugens.
• 2 Experimental methods included no‐choice larval feeding assays and field cage and laboratory oviposition trials. Difficulty in ascertaining reliable oviposition preference data from cage oviposition trials limited the confidence with which the invader's field host range could be predicted.
• 3 Field surveys of plants attacked by U. lugens in the infested area supported the initial predictions but only some of the at‐risk native Myrtaceae were present in the area.
• 4 The risk to native New Zealand plants is presented in terms of two mechanisms: development of self‐sustaining populations and temporary spill‐over of solitary larvae. Development of self‐sustaining populations of the pest within native forests is deemed to be highly improbable.
• 5 Temporary spill‐over impacts are most likely in urban areas within mixed species plantings or boundaries between native/exotic forests and coinciding with the mobile larval life stages. Spill‐over impacts from U. lugens have only been recorded to date on Metrosideros excelsa.

     

Plant and host effects on the leafroller parasitoid Dolichogenidia tasmanica

Parasitism by the braconid wasp Dolichogenidia tasmanica of first instar larvae of the lightbrown apple moth Epiphyas postvittana, established on four different species of potted host plants, was assessed after 2 weeks of field exposure in an apple orchard. Parasitism varied significantly between larval host plants (apple 58%, broom 59%, clover 30%, poplar 19% ). Parasitism on potted apple seedlings of the co-evolutionary host, E. postvittana, was compared in a field trial with that of two native New Zealand leafroller species (to which D. tasmanica has had exposure for 5 decades only). Parasitism varied significantly with larval host (E. postvittana, 83%; Ctenopseustis herana, 58%;Planotortrix octo, 26% ). Larval collections were made from mature apple trees and identification of larvae was achieved by DNA analysis for the leafroller species using PCR-RFLP of ITS1 + ITS2, and for the parasitoid by specific PCR of partial 18S. Parasitism under natural field conditions on mature apple trees was not different between larval hosts (mean 32.5% ). In laboratory studies, more P. octo larvae departed in response to parasitoid probing behaviour than E. postvittana, which is likely to contribute to the difference in parasitism rates. This study conclusively shows that D. tasmanica parasitises native New Zealand leafrollers, despite their different evolutionary origins.     

Effects of tall fescue turf on growth and nitrogen fixation potential of the woody legume Lupinus albifrons

The effect of tall fescue turf on growth, flowering, nodulation, and nitrogen fixing potential of Lupinus albifrons Benth. was examined for greenhouse and field grown plants. No allelopathic effect was observed for lupine plants treated with tall fescue leachates. The nitrogen-fixing potential measured by nodule dry weight and acetylene reduction rates was not significantly affected by tall fescue turf.Both the greenhouse and field studies showed that the growth, sexual reproductive allocation and number of inflorescences were significantly reduced when lupine plants were grown with tall fescue. The root-length densities of tall fescue turf and lupine monoculture were measured. The tall fescue turf had 20 times higher root-length density (20 cm cm^-3 soil) than the lupine plant monoculture. This suggests that intense competition at the root zone may be a dominant factor which limits the growth of the lupine plants.The flowering characters of the lupine plants were improved by phosphorus fertilization. Transplanting of older lupine plants into the turf substantially alleviated the tall fescue turf competitive effect.     

Potential risk of accidental introduction of Asian gypsy moth (Lymantria dispar) to Australasia: effects of climatic conditions and suitability of native plants

1 The potential risk of the establishment of the Asian strain of the gypsy moth (AGM) (Lymantria dispar) in New Zealand and Australia (Australasia) was assessed from a study of the insect's host range and potential distribution. In New Zealand, viable eggs of AGM have been continuously intercepted on cargo from Asia, and therefore there is a high probability of accidental introductions of AGM to Australasia. 2 We predicted potential distribution ranges of AGM based on climatic conditions. Asian gypsy moth is predicted to be able to persist in N and SE New Zealand and SE and SW Australia. 3 Using three populations of AGM and 59 species (seven families) of plant (55 from Australasia and four from elsewhere), we also conducted laboratory trials to examine the ability of AGM larvae to complete development on native plants from Australasia. Asian gypsy moth was able to complete development on 26 out of the 55 native species tested. Furthermore, larval performance on at least five species of Australian native plant was as good as on AGM's preferred host species (Quercus pubescens and Q. robur). 4 Larval performance of AGM was poor on all but one species of New Zealand native tree species (Nothofagus solandri), and therefore the risk of establishment in the indigenous forests of New Zealand is considered to be low. 5 Given the suitability of some Australian plants and the climatic suitability for the establishment of AGM, this insect should be treated as a serious quarantine threat and managed accordingly, particularly in Australia.     

The nutritional value of narrow-leafed lupine (Lupinus angustifolius) for fattening pigs

The aim of this study was to determine the nutrient digestibility of seeds of four varieties of narrow-leafed lupines (Lupinus angustifolius) and the possibility of soya bean meal (SBM) substitution by lupine seeds alone and in combination with rapeseed meal (RSM) in the diets of pigs. The seeds of the lupine varieties Kalif, Sonet, Zeus and Boruta were analysed. The apparent total tract digestibility (ATTD) was determined on 50 cross-bred pigs using the difference method with titanium dioxide as a marker. The substitution of SBM by lupine seeds alone (at 0 – 100%) was tested on 60 pigs (20–105 kg body weight (BW)) and by a combination of lupine seeds and RSM on 180 fattening pigs (35–80 kg BW). The chemical composition of lupine seeds differed considerably, especially in terms of crude protein and mineral content. All seeds contained less than 0.05% alkaloids and 9.3% oligosaccharides in dry matter. The ATTD of protein ranged from 70% to 74%, those of ether extract from 36% to 55% and those of gross energy from 77% to 84%. The entire replacement of SBM by lupine seeds (var. Sonet) did not have a negative effect on the performance of grower and fattener pigs. The substitution of SBM by a combination of lupines and RSM reduced the performance of growing and finishing pigs significantly.     

Diversification of lupine Bradyrhizobium strains: evidence from nodulation gene trees

Bradyrhizobium strains isolated in Europe from Genisteae and serradella legumes form a distinct lineage, designated clade II, on nodulation gene trees. Clade II bradyrhizobia appear to prevail also in the soils of Western Australia and South Africa following probably accidental introduction with seeds of their lupine and serradella hosts. Given this potential for dispersal, we investigated Bradyrhizobium isolates originating from a range of native New World lupines, based on phylogenetic analyses of nodulation (nodA, nodZ, noeI) and housekeeping (atpD, dnaK, glnII, recA) genes. The housekeeping gene trees revealed considerable diversity among lupine bradyrhizobia, with most isolates placed in the Bradyrhizobium japonicum lineage, while some European strains were closely related to Bradyrhizobium canariense. The nodA gene tree resolved seven strongly supported groups (clades I to VII) that correlated with strain geographical origins and to some extent with major Lupinus clades. All European strains were placed in clade II, whereas only a minority of New World strains was placed in this clade. This work, as well as our previous studies, suggests that clade II diversified predominately in the Old World, possibly in the Mediterranean. Most New World isolates formed subclade III.2, nested in a large "pantropical" clade III, which appears to be New World in origin, although it also includes strains originating from nonlupine legumes. Trees generated using nodZ and noeI gene sequences accorded well with the nodA tree, but evidence is presented that the noeI gene may not be required for nodulation of lupine and that loss of this gene is occurring.     

A modelling approach to estimate the effect of exotic pollinators on exotic weed population dynamics: bumblebees and broom in Australia

The role of mutualisms in contributing to species invasions is rarely considered, inhibiting effective risk analysis and management options. Potential ecological consequences of invasion of non‐native pollinators include increased pollination and seed set of invasive plants, with subsequent impacts on population growth rates and rates of spread. We outline a quantitative approach for evaluating the impact of a proposed introduction of an invasive pollinator on existing weed population dynamics and demonstrate the use of this approach on a relatively data‐rich case study: the impacts on Cytisus scoparius (Scotch broom) from proposed introduction of Bombus terrestris. Three models have been used to assess population growth (matrix model), spread speed (integrodifference equation), and equilibrium occupancy (lattice model) for C. scoparius. We use available demographic data for an Australian population to parameterize two of these models. Increased seed set due to more efficient pollination resulted in a higher population growth rate in the density‐independent matrix model, whereas simulations of enhanced pollination scenarios had a negligible effect on equilibrium weed occupancy in the lattice model. This is attributed to strong microsite limitation of recruitment in invasive C. scoparius populations observed in Australia and incorporated in the lattice model. A lack of information regarding secondary ant dispersal of C. scoparius prevents us from parameterizing the integrodifference equation model for Australia, but studies of invasive populations in California suggest that spread speed will also increase with higher seed set. For microsite‐limited C. scoparius populations, increased seed set has minimal effects on equilibrium site occupancy. However, for density‐independent rapidly invading populations, increased seed set is likely to lead to higher growth rates and spread speeds. The impacts of introduced pollinators on native flora and fauna and the potential for promoting range expansion in pollinator‐limited ‘sleeper weeds’ also remain substantial risks.     

Multispecies interactions among a plant parasite,a pollinator and a seed predator affect the reproductive output of an invasive plant,Cytisus scoparius

Interactions between species pairs are almost always mediated by interactions with other species. The outcomes of these multispecies interactions are often difficult to predict and are rarely studied. In addition to their theoretical importance, multispecies interactions are also important for management situations. Where multiple agents are used to control invasive species, interactions between agents may either enhance or reduce the impacts on the target species, or may simply have additive effects. In this study, conducted in a Mediterranean‐type woodland in Australia, we examined how the interaction between an invasive legume, Cytisus scoparius (Leguminosae), its pollinator, Apis mellifera and a seed predator, Bruchidius villosus (Bruchidae), are modified by a native hemiparasitic vine, Cassytha pubescens (Loranthaceae). The parasite had a direct negative effect on C. scoparius, reducing flowering by 50% and consequently fruit and seed production. Despite having fewer flowers, infected plants had the same proportion of ‘tripped’ flowers, an indirect measure of pollinator visitation, as uninfected plants; although fruit formed on infected plants it was more likely to abort prematurely. Seed predation by B. villosus was lower on parasite‐infected C. scoparius plants than in uninfected plants. Although Ca. pubescens had an antagonistic effect on B. villosus, in consort, the two agents reduced overall seed production by an average of 62%. The acquired parasite Ca. pubescens was more effective in reducing reproductive output than the introduced seed predator, B. villosus, and shows potential as a biocontrol agent for C. scoparius. We documented a subadditive effect of two biological enemies on the invasive species, where the acquired parasite had a stronger effect than the introduced seed predator.     

Coastal sand dune stabilization in the Pacific Northwest

Coastal dunes are stabilized in three stages: (1) The initial stage uses sand-stilling grasses established vegetatively. For this purpose, European beachgrass,Ammophila arenaria (L.) Link., is most used, followed by American beachgrass,A. breviligulata Fern., or American dunegrass,Elymus mollis Trin. Large solid plantings must be made with the spacing and number of plants per hill adjusted to the site conditions. Plantings, using Nitrogen fertilizer, are made during the dormant season. (2) Secondary permanent grasses and legumes are seeded 2 years later when sand movement slows and the initial grasses begin to lose their vigor. Most used are seaside lupine,Lupinus littoralis Dougl., purple beachpea,Lathyrus japonicus Willd., seashore bluegrass,Poa macrantha Vasey., and native red fescue,Festuca rubra L. (3) Final control is with woody plants that are well adapted to rough areas. Scotch broom,Cytisus scoparius (L.) Link. Hooker willow,Salix hookeriana Barratt, Nootka rose,Rosa nutkana Prese., and shore pine Pinus contorta Dougl., have been extensively used.Presented at the Seventh International Biometeorological Congress, 17–23 August 1975, College Park, Maryland, U.S.A.     

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.     

Introduced Scotch broom (Cytisus scoparius) invades the genome of native populations in vulnerable heathland habitats

Cytisus scoparius is a global invasive species that affects local flora and fauna at the intercontinental level. Its natural distribution spans across Europe, but seeds have also been moved among countries, mixing plants of native and non‐native genetic origins. Hybridization between the introduced and native gene pool is likely to threaten both the native gene pool and the local flora. In this study, we address the potential threat of invasive C. scoparius to local gene pools in vulnerable heathlands. We used nuclear single nucleotide polymorphic (SNP) and simple sequence repeat (SSR) markers together with plastid SSR and indel markers to investigate the level and direction of gene flow between invasive and native heathland C. scoparius. Analyses of population structures confirmed the presence of two gene pools: one native and the other invasive. The nuclear genome of the native types was highly introgressed with the invasive genome, and we observed advanced‐generation hybrids, suggesting that hybridization has been occurring for several generations. There is asymmetrical gene flow from the invasive to the native gene pool, which can be attributed to higher fecundity in the invasive individuals, measured by the number of flowers and seed pods. Strong spatial genetic structure in plastid markers and weaker structure in nuclear markers suggest that seeds spread over relatively short distances and that gene flow over longer distances is mainly facilitated by pollen dispersal. We further show that the growth habits of heathland plants become more vigorous with increased introgression from the invaders. Implications of the findings are discussed in relation to future management of invading C. scoparius.     

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.     

Combined effects of drought stress and psyllid herbivory on the invasive weed Scotch broom,Cytisus scoparius

Increases in the frequency and duration of droughts under global climate change could have implications for plant–insect interactions, and could either increase or decrease the effects of weed biological control agents. In this study, we used greenhouse and field experiments to examine the impacts of drought stress on the abundance and impacts of the adventive psyllid Arytainilla spartiophila Forster (Hemiptera: Psyllidae, Arytaininae) on its target weed Scotch broom, Cytisus scoparius (L.) Link (Fabaceae), in California, USA. The psyllid impacted plant growth in both field and greenhouse experiments, whereas drought stress affected plant growth in the greenhouse only, suggesting that other factors besides water availability may be more limiting for plants in the field. Effects on psyllid survival were consistent with the plant vigor hypothesis, which predicts that herbivores will perform better on vigorously growing plants; psyllid numbers were lower on drought-stressed plants in the greenhouse and were correlated with plant growth in both the greenhouse and the field. In the greenhouse, the combined effects of the psyllid and drought stress were additive, indicating that the effects of the psyllid were consistent across unstressed and drought-stressed plants. Although the psyllid is unlikely to control Scotch broom on its own, results suggest that it may work in conjunction with drought stress to suppress Scotch broom.