Margaret C. Stanley

Selection for chemical trait remixing in an invasive weed after reassociation with a coevolved specialist.

The interaction between Depressaria pastinacella (parsnip webworm) and wild parsnip (Pastinaca sativa), in its native Europe and in its longstanding nonindigenous range in the midwestern United States, is characterized by chemical phenotype matching, ostensibly mediated by reciprocal selective responses. The first appearance of D. pastinacella on P. sativa in New Zealand in 2004 provided an opportunity to quantify selective impacts of a coevolved herbivore and calibrate rates of phytochemical response in its host plant. Webworms in 2006 reduced seed production up to 75% in New Zealand populations, and in 2007 infestations increased in severity in all populations except one. Most New Zealand populations fall into a furanocoumarin phenotype cluster distinct from European and U.S. phenotypes, although one heavily attacked population clusters with two U.S. populations and one European population long associated with webworms. Multivariate selection analysis substituting realized fitness (with webworms present) for potential fitness (absent webworms) as the dependent variable revealed that reassociation with a coevolved specialist in a nonindigenous area profoundly altered the selection regime, favoring trait remixing and rapid chemical changes in parsnip populations, as predicted by the geographic mosaic theory. That uninfested populations of New Zealand parsnips contain higher amounts of octyl acetate, a floral volatile used by webworms for orientation, suggests that plants that escape from specialized enemies may also experience selection to increase kairomones, as well as to reduce allomones.

Supplementary feeding restructures urban bird communities

Significance Bird feeding is essentially a massive global supplementary feeding experiment, yet few studies have attempted to explore its ecological effects. In this study we use an in situ experimental approach to investigate the impacts of bird feeding on the structure of local bird assemblages. We present vital evidence that bird feeding contributes to the bird community patterns we observe in urban areas. In particular, the study demonstrates that common feeding practices can encourage higher densities of introduced birds, with potential negative consequences for native birds. Food availability is a primary driver of avian population regulation. However, few studies have considered the effects of what is essentially a massive supplementary feeding experiment: the practice of wild bird feeding. Bird feeding has been posited as an important factor influencing the structure of bird communities, especially in urban areas, although experimental evidence to support this is almost entirely lacking. We carried out an 18-mo experimental feeding study at 23 residential properties to investigate the effects of bird feeding on local urban avian assemblages. Our feeding regime was based on predominant urban feeding practices in our region. We used monthly bird surveys to compare avian community composition, species richness, and the densities of local species at feeding and nonfeeding properties. Avian community structure diverged at feeding properties and five of the commonest garden bird species were affected by the experimental feeding regime. Introduced birds particularly benefitted, with dramatic increases observed in the abundances of house sparrow (Passer domesticus) and spotted dove (Streptopelia chinensis) in particular. We also found evidence of a negative effect on the abundance of a native insectivore, the grey warbler (Gerygone igata). Almost all of the observed changes did not persist once feeding had ceased. Our study directly demonstrates that the human pastime of bird feeding substantially contributes to the structure of avian community in urban areas, potentially altering the balance between native and introduced species.

IMPORTANCE OF SEED INGESTION TO AN AVIAN FRUGIVORE: AN EXPERIMENTAL APPROACH TO FRUIT CHOICE BASED ON SEED LOAD

Abstract Frugivorous birds may be able to reduce the cost of processing seeds by discarding seeds, selecting fruits that have a high pulp-to-seed ratio, or by choosing fruit in which seeds are packaged in a way that the frugivores gut can void them more rapidly. A preference for fruit based on pulp-to-seed ratio or seed composition within a fruit is likely to have important implications for plants and evolution of seed size. We tested whether captive Silvereyes (Zosterops lateralis) discriminate among artificial fruit on the basis of seed presence by presenting birds with artificial fruit with or without a seed. In the first experiment, fruit were translucent so that birds could see which fruit contained a seed. In the second experiment, the visual cue was removed. When Silvereyes were presented with a choice between translucent, artificial fruit with or without a seed, they showed a strong preference for fruit that did not contain a seed. However, when the visual cue to seed presence was removed, preference for seedless fruit was still significant, but markedly reduced. We also tested seed-size preference of Silvereyes in the field in Victoria, Australia. Seeds from a fruit commonly consumed by Silvereyes, fragrant saltbush (Rhagodia parabolica), were recovered from Silvereye faecal samples and their volumes measured. Comparisons were made between seed volumes of fruit consumed by Silvereyes and those within fruit available on the plant. Silvereyes consumed significantly smaller seeds than the mean size available on saltbush plants. When Silvereyes were presented with a cereal-based diet containing artificial seeds (designed to mimic large fruit containing many small seeds), they avoided seed ingestion and were able to consume proportionally more cereal than seeds, even when on a high seed-load diet (30%). Seed dispersal by Silvereyes may be inefficient for plant species with large fruit containing many small seeds, because Silvereyes in this experiment were able to avoid ingesting seeds.

DOES SEED PACKAGING INFLUENCE FRUIT CONSUMPTION AND SEED PASSAGE IN AN AVIAN FRUGIVORE

Abstract Seed packaging is one fruit characteristic that may influence post-ingestional fruit processing in avian frugivores. We tested the response of a facultative frugivore, the Silvereye (Zosterops lateralis), to fruit containing different forms of seed packaging. Wild-caught, captive Silvereyes were presented with artificial fruit containing either one large seed or three small seeds of equivalent total volume, and their consumption rates were recorded over 90 min. In a second experiment, the seed transit times (ingestion to excretion) for similar large-seeded and small-seeded fruit consumed by Silvereyes were recorded. Silvereyes consumed significantly more large-seeded fruit than small-seeded fruit. The transit time of seeds was also significantly shorter for large-seeded (mean = 22 min) than for small-seeded fruit (mean = 29 min). Thus seed packaging had a significant influence on the rate at which fruit were processed. Silvereyes were able to consume more large-seeded than small-seeded fruit because the seeds in large-seeded fruit were defecated faster than those in small-seeded fruit. It is likely that Silvereyes can compensate for the costs of seed ingestion through having a rapid gut passage rate and hence an increased fruit consumption rate. The gut of Silvereyes showed morphological characters intermediate between insectivores and specialist frugivores. The dimensions of the intestine and gizzard were like those of insectivores and the gizzard was substantially larger than those of specialist frugivores. Facultative frugivores appear to have few morphological adaptations to frugivory and we argue that this facilitates plasticity in the diet and the processing of insects when fruit is not available. ¿Influencia el Empaquetamiento de Semillas el Consumo de Frutos y el Tiempo de Tránsito en Aves Frugívoras? Resumen. El empaquetamiento de semillas es una característica que puede influenciar el procesamiento de frutos post ingestión en aves frugivoras. Pusimos a prueba la respuesta de un frugívoro facultativo (Zosterops lateralis) a frutos con distintas formas de empaquetamiento de semillas. Individuos de Z. lateralis silvestres fueron capturados y expuestos a frutos artificiales de volumen equivalente con una semilla grande o con tres semillas pequeñas y su tasa de consumo fue registrada durante 90 minutos. En un segundo experimento, se registró el tiempo de tránsito (de ingesta a defecación) de frutos similares con semillas grandes y pequeñas. Z. lateralis consumió una significativamente mayor cantidad de frutos con semillas grandes que de frutos con semillas pequeñas. El tiempo de tránsito de semillas grandes fué significativamente menor (promedio = 22 min) que el de semillas pequeñas (promedio = 29 min). Por lo tanto, el empaquetamiento de semillas tuvo una influencia significativa sobre la tasa a la que los frutos fueron procesados. Z. lateralis fue capaz de consumir más frutos con semillas grandes ya que sus semillas fueron defecadas más rápidamente que las de frutos con semillas pequeñas. Es probable que Z. lateralis pueda compensar el costo de la ingesta de semillas con un paso rápido a través del tracto digestivo lo que se traduce en un aumento en la tasa de consumo de frutos. El tracto digestivo de Z. lateralis muestra caracteres morfológicos intermedios entre los de insectívoros y frugívoros especialistas. Las dimensiones de los intestinos y la molleja son similares a las de insectívoros y la molleja es sustancialmente más grande que la de los frugívoros especialistas. Los frugívoros facultativos parecieran tener pocas adaptaciones morfológicas a la frugivoría. Sugerimos que esto facilita una mayor plasticidad en la dieta y en el procesamiento de insectos cuando existe poca disponibilidad de frutos.

Predicting plant invasions under climate change: are species distribution models validated by field trials?

Climate change may facilitate alien species invasion into new areas, particularly for species from warm native ranges introduced into areas currently marginal for temperature. Although conclusions from modelling approaches and experimental studies are generally similar, combining the two approaches has rarely occurred. The aim of this study was to validate species distribution models by conducting field trials in sites of differing suitability as predicted by the models, thus increasing confidence in their ability to assess invasion risk. Three recently naturalized alien plants in New Zealand were used as study species (Archontophoenix cunninghamiana, Psidium guajava and Schefflera actinophylla): they originate from warm native ranges, are woody bird-dispersed species and of concern as potential weeds. Seedlings were grown in six sites across the country, differing both in climate and suitability (as predicted by the species distribution models). Seedling growth and survival were recorded over two summers and one or two winter seasons, and temperature and precipitation were monitored hourly at each site. Additionally, alien seedling performances were compared to those of closely related native species (Rhopalostylis sapida, Lophomyrtus bullata and Schefflera digitata). Furthermore, half of the seedlings were sprayed with pesticide, to investigate whether enemy release may influence performance. The results showed large differences in growth and survival of the alien species among the six sites. In the more suitable sites, performance was frequently higher compared to the native species. Leaf damage from invertebrate herbivory was low for both alien and native seedlings, with little evidence that the alien species should have an advantage over the native species because of enemy release. Correlations between performance in the field and predicted suitability of species distribution models were generally high. The projected increase in minimum temperature and reduced frosts with climate change may provide more suitable habitats and enable the spread of these species.

The response of captive silvereyes (Zosterops lateralis) to the colour and size of fruit

Birds are important consumers and dispersers of the seeds of fleshy fruit and some have been shown to be selective in their choice of fruit. However, our knowledge of how birds respond to a variety of fruit characteristics is poor. Some birds are known to avoid green fruit or consume them less than fruit of other colours. The fruit of many plant species are green when they are unripe and contain low concentrations of sugars and high concentrations of secondary compounds. In this study, captive silvereyes (Zosterops lateralis) were presented with a choice of red, white and green artificial fruit. Half of them were given these fruit with equal sugar concentrations (15%) and the other half were given the choice but with the green fruit having a sugar concentration twice that of the other two colours (30%). Green fruit were not strongly avoided by silvereyes and were actually preferred when they had a higher sugar concentration than the other two fruit types (red and white). Sugar concentration was therefore a more important determinant of fruit choice than colour. Fruit size is also known to affect fruit choice in some bird species. Small fruit are easier to consume than larger fruit, but contain less pulp. To maximise energy gain, birds should consume fruit that are large, but not large enough to incur high handling costs. Silvereyes in this study were presented with a choice between 20 artificial fruit 4 mm in diameter (large fruit) and 20 artificial fruit 2 mm in diameter (small fruit), both of which they are able to consume. In this experiment, silvereyes exhibited a strong preference for large fruit over small fruit. In general, larger fruit contain more pulp and therefore more energy than smaller fruit. However, other fruit traits, such as seed load, are likely to influence fruit choice by silvereyes in the wild and result in a trait hierarchy.

Emerging threats in urban ecosystems: a horizon scanning exercise

As urbanization intensifies, urban ecosystems are increasingly under pressure from a range of threats. Horizon scanning has the potential to act as an early warning system, thereby initiating prompt discussion and decision making about threat mitigation. We undertook a systematic horizon scanning exercise, using a modified Delphi technique and experts from wide-ranging disciplines, to identify emerging threats in urban ecosystems. The 10 identified threats were generally associated with rapid advances in technology (eg solar panels, light-emitting diode lights, self-healing concrete) or with societal demands on urban nature (eg green prescriptions). Although many of the issues identified are also technological opportunities with recognized environmental benefits, we have highlighted emerging risks so that research and mitigation strategies can be initiated promptly. Given the accelerated rate of technological advancement and the increasing demands of urbanized populations, horizon scanning should be condu...

Assessing the risk of invasive ants: a simple and flexible scorecard approach

Abstract.Numerous ant species are being transported around globe through international trade, many of which become invasive. To mitigate the potential impacts of invasive ant species, an assessment system is needed which is simple, can be tailored for specific-user and regional requirements, and has the potential to integrate a variety of additional information. Here we present such a system, which has been used in New Zealand to determine which ant species pose the greatest threat to natural ecosystems. However, the scorecard system could be applied widely around the globe for assessing the threat of invasive ant species.

Evolution of tolerance in an invasive weed after reassociation with its specialist herbivore

The interaction between the European wild parsnip Pastinaca sativa and its coevolved florivore the parsnip webworm Depressaria pastinacella, established in North America for over 150 years, has resulted in evolution of local chemical phenotype matching. The recent invasion of New Zealand by webworms, exposing parsnips there to florivore selection for the first time, provided an opportunity to assess rates of adaptive response in a real‐time experiment. We planted reciprocal common gardens in the USA and NZ with seeds from (1) US populations with a long history of webworm association; (2) NZ populations that had never been infested and (3) NZ populations infested for 3 years (since 2007) or 6 years (since 2004). We measured impacts of florivory on realized fitness, reproductive effort and pollination success and measured phenotypic changes in infested NZ populations relative to uninfested NZ populations to determine whether rapid adaptive evolution in response to florivory occurred. Irrespective of country of origin or location, webworms significantly reduced plant fitness. Webworms reduced pollination success in small plants but not in larger plants. Although defence chemistry remained unchanged, plants in infested populations were larger after 3–6 years of webworm florivory. As plant size is a strong predictor of realized fitness, evolution of large size as a component of florivore tolerance may occur more rapidly than evolution of enhanced chemical defence.

Invasive interactions: can Argentine ants indirectly increase the reproductive output of a weed?

The direct and indirect interactions of invasive ants with plants, insect herbivores, and Hemiptera are complex. While ant and Hemiptera interactions with native plants have been well studied, the effects of invasive ant–scale insect mutualisms on the reproductive output of invasive weeds have not. The study system consisted of Argentine ants (Linepithema humile), boneseed (Chrysanthemoides monilifera monilifera), and sap-sucking scale insects (Hemiptera: Saissetia oleae and Parasaissetia nigra), all of which are invasive in New Zealand. We examined the direct and indirect effects of Argentine ants on scale insects and other invertebrates (especially herbivores) and on plant reproductive output. Argentine ants spent one-third of their time specifically associated with scale insects in tending behaviours. The invertebrate community was significantly different between uninfested and infested plants, with fewer predators and herbivores on ant-infested plants. Herbivore damage was significantly reduced on plants with Argentine ants, but sooty mould colonisation was greater where ants were present. Herbivore damage increased when ants were excluded from plants. Boneseed plants infested with Argentine ants produced significantly more fruits than plants without ants. The increase in reproductive output in the presence of ants may be due to increased pollination as the result of pollinators being forced to relocate frequently to avoid attack by ants, resulting in an increase in pollen transfer and higher fruit/seed set. The consequences of Argentine ant invasion can be varied; not only does their invasion have consequences for maintaining biodiversity, ant invasion may also affect weed and pest management strategies.