Classical biological control: exploiting enemy escape to manage plant invasions

Practitioners of classical biological control of invasive weeds are confronted with a dual expectation: to achieve successful control of plant invaders and to avoid damage to nontarget plants and adverse indirect effects. In this paper we discuss key issues that we consider to be crucial for a safe, efficient, and successful classical biological control project, and that have also caused some recent controversy. These include selection of effective control agents, host specificity of the biological control agents, implications of the genetic population structure of the target populations, and potential impact on native food webs. With regard to improving the success rate of biological control of plant invaders, we first emphasize the importance of a clear a priori definition of success and a more ecosystem-based approach to better document both negative effects of the invasive plant as well as potential positive and negative effects of introducing biological control agents. Secondly, pre-release impact assessment could be improved by better focusing on how to reach high densities of the control agents and by including tolerance to and compensation of herbivory. Thirdly, we advocate a reinforced effort to integrate and combine biological control in combination with existing or potential management options. Finally, we propose various ecological and evolutionary hypotheses in the framework of our topic to document that biological control programmes against plant invaders also offer a great opportunity to gain new insights into basic processes in ecology and evolution.     

Risk and ethics in biological control

All introduced natural enemies present a degree of risk to nontarget species. Since most biological control programs use relatively host-specific natural enemies, the risk to nontarget species is generally very low, particularly from biological control of weeds, which uses extensively tested and validated host-specificity testing procedures to predict risk. However, many of the published comments about risks of biological control are superficial or misleading, often inappropriately lumping risk from all taxa of agents as “the risk of biological control,” and ignore the potential benefits, rather than dealing with species-by-species risk and benefits. Particularly confounding accurate predictions is the common mixing of parameters of hazard and exposure in discussions of risk. In this paper, traditional risk analysis techniques are discussed and adapted for biological control. How people perceive risk is the key to understanding their attitude to risk. Some of the criticisms of biological control relating to inadequate post-release monitoring are valid and the ethical responsibilities of biological control scientists in this area are also discussed. Biological control scientists should address objectively the criticisms of biological control, continue to review and adjust current host-specificity testing procedures and make appropriate changes. This process will result in better science, ultimately delivering more focused programs, and altering the perception of risk from biological control agents by objective observers.     

Success and safety in the biological control of environmental weeds in New Zealand

Biological control, using specialist insect herbivores and plant pathogens, can be a self‐sustaining, cost‐effective and low‐risk tool for the management of environmental weeds. Agents have been recorded attacking non‐target plants in New Zealand and elsewhere, but the effects are usually minor and/or transitory. It seems probable that only two cases, worldwide, will result in significant damage to non‐target plants (representing 0.5% of the nearly 400 insect, mite, or fungal species used in classical weed biocontrol). Both of these cases were predictable from host range testing. Negative indirect, or ‘downstream’, ecological effects from specific weed biocontrol agents are difficult to predict and measure. They are probably insignificant compared to the impacts of the invasive plants that the agents are introduced to control. However, it is necessary to balance the risks associated with any introduction against the environmental benefits from controlling a weed to a predicted level. Recent analyses suggest that success rates are better than generally perceived. For New Zealand programmes, where enough time has lapsed to allow assessment, we calculate a full/partial success rate of 83%. Many of the costs associated with environmental weeds are difficult to quantify. Detailed risk assessment will make biological control programmes more expensive and time‐consuming, so that reliance on non‐biological management methods for environmental weeds may actually increase. The costs of biocontrol programmes against some New Zealand weeds can be kept down by using research already carried out in Australia and other countries, and the process is reciprocal. Developing international consortia of sponsors is also a potential way to fund programmes against weeds shared by several countries.     

Environmental Health Indicators in New Zealand: Drinking Water—A Case Study

This study examines and quantifies the linkages between population health, environmental risks, and its determinants for drinking water in New Zealand using routinely collected data. It was conducted as part of the national environmental health indicators project in New Zealand. The project is based on the World Health Organization’s (WHO) “Environmental and Health Information System” program. Drinking water quality indicators based on the Driving force–Pressure–State–Exposure–Effect–Action (DPSEEA) framework as part of this program were analyzed to validate the model by quantifying the linkages between the indicators. The results of the model suggested over the study period, the state (drinking water quality) and exposure (water access) indicators are significant independent predictors of the effect indicator (waterborne disease rate). This study suggests that routinely collected data can be structured using the DPSEEA framework and tested quantitatively using standard Poisson regression models, thus, illustrating that the model can be used routinely to provide a basis for consideration of the costs and benefits of any interventions to reduce the burden of waterborne disease. Data quality issues need to be considered if such routinely collected data linkages are to be performed for policy purposes. The online version of this article (doi:) contains electronic supplementary material, which is available to authorized users. Supplemental histograms are available in the online appendix.     

Modelling the use of NPV for the biological control of Asian gypsy moth Lymantria dispar invading New Zealand

1 The accidental introduction of the Asian strain of gypsy moth (AGM) Lymantria dispar (L.) to New Zealand poses a major threat to New Zealand's forestry industry. To aid eradication and control decisions in the event of its establishment, a model was developed for the effect of nuclear polyhedrosis virus (NPV) as biological control for AGM in New Zealand. 2 The model simulates within‐ and between‐season gypsy moth population dynamics, including temperature‐dependent development, density dependence through predation and resource limitation, and interactions with NPV. 3 Following its introduction to New Zealand, AGM is predicted to increase more quickly and have more severe outbreaks than the European strain. In the absence of predators, the model predicts initial outbreaks then damped oscillations to an equilibrium. 4 In the model, a single application of NPV (2500 GPIB ha^?1) at the time of maximum larval density gave up to 80% suppression of peak larval densities in the following year. The same level of suppression was achieved in the absence of predators. 5 In the long term, the model predicted that spraying when an outbreak was just beginning gave best results (a 50–70% reduction of the following two outbreaks). Simulation of threshold spraying resulted in NPV application, on average, every 7 years and suppression of outbreak densities by 40–70%. Following a single application, NPV was maintained in the population as a classical biological control agent, giving approximately a 20% reduction in outbreak densities. 6 Eradication of AGM using NPV was possible if larval densities were very low.     

Indirect nontarget effects of host-specific biological control agents: Implications for biological control

Classical biological control of weeds currently operates under the assumption that biological control agents are safe (i.e., low risk) if they do not directly attack nontarget species. However, recent studies indicate that even highly host-specific biological control agents can impact nontarget species through indirect effects. This finding has profound implications for biological control. To better understand the causes of these interactions and their implications, we evaluate recent case studies of indirect nontarget effects of biological control agents in the context of theoretical work in community ecology. We find that although particular indirect nontarget effects are extremely difficult to predict, all indirect nontarget effects of host specific biological control agents derive from the nature and strength of the interaction between the biological control agent and the pest. Additionally, recent theoretical work suggests that the degree of impact of a biological control agent on nontarget species is proportional to the agent’s abundance, which will be highest for moderately successful control agents. Therefore, the key to safeguarding against indirect nontarget effects of host-specific biological control agents is to ensure the biological control agents are not only host specific, but also efficacious. Biological control agents that greatly reduce their target species while remaining host-specific will reduce their own populations through density-dependent feedbacks that minimize risks to nontarget species.     

新西兰果园规范化栽培技术及发展趋势

报道了新西兰水果主产区果园普遍采用的规范化栽培技术系统,包括果园布局、果树定植与枝条定向牵引、果产品全程质量管理、果园生态环境管理等技术,并分析了该技术的特点、目的和作用,对新西兰果园近年在栽培技术上的优化和创新进行了总结,提出了新西兰果园栽培的发展趋势。     

Evolving MPA Management in New Zealand: Between Principle and Pragmatism

In January 2016 New Zealand released a consultation document proposing a new act on marine protected areas designed to significantly reform current and now dated policy. This article explores those reform proposals in the context of the current regulatory regime, international obligations, and the best practice of selected other states. While the proposed act provides for a much firmer legislative base from which to develop an MPA network to conserve biodiversity and ecosystem function, nevertheless it is limited in geographic and functional scope. As such, it represents a missed opportunity and undermines New Zealand's claims to be an international leader in ocean management.     

Assessment of risk of attack to safflower by Ceratapion basicorne (Coleoptera: Apionidae), a prospective biological control agent of Centaurea solstitialis (Asteraceae)

Ceratapion basicorne (Coleoptera: Apionidae) is a prospective biological control agent of yellow starthistle (Centaurea solstitialis, Asteraceae: Cardueae), which is an important invasive alien weed in the western United States. Previous studies have shown that it is possible for this insect to oviposit on and complete development on safflower (Carthamus tinctorius) under no-choice laboratory conditions; however, it has never been reported as a pest of safflower. Field experiments were conducted at three sites in eastern Turkey during 3 years to evaluate the risk of attack on safflower by this insect in its native range. At two sites where C. basicorne was the only apionid observed, no safflower plants were attacked despite high attack rates on yellow starthistle test plants (48–98% of plants infested). At a third site, where C. basicorne and three other species in the same genus; C. scalptum, C. orientale, and C. onopordi were present, 8–26% of safflower plants were infested, but none of the insects reared from safflower during 3 years were C. basicorne. Other authors have reported rearing C. basicorne from field-collected plants of only Ce. solstitialis, Ce. cyanus, Ce. depressa, and Cnicus benedictus. Our results indicate that C. basicorne does not attack safflower under field conditions and that its introduction would not pose a risk to this crop.     

Conceiving Reproduction: New Reproductive Technologies and the Redefinition of the Kinship Narrative in New Zealand Society

This paper focuses on technologies for assisting conception and considers how normative notions of conception and family formation have informed the rejection, incorporation and adaptation of technologies and their regulation in New Zealand. Drawing on a textual analysis of primary written materials and secondary sources generated between 1965 and 2004, the paper reveals how these technologies help various groups in New Zealand society make explicit their understandings of relatedness, identity and social and cultural belonging. Whilst biological/genetic connection and social connections inform understandings of kinship, reproduction, parenthood and social structure for many New Zealanders, the boundaries between the biological and social are often blurred and precedence of one over the other is very much context dependent. The incorporation of the new reproductive technologies has been an inherently politicised process, with barriers to access to these technologies emerging in relation to ethnic identity and race, sexuality, socioeconomic status, gender, and marital status.     

Extinction and recolonization of coastal megafauna following human arrival in New Zealand

Extinctions can dramatically reshape biological communities. As a case in point, ancient mass extinction events apparently facilitated dramatic new evolutionary radiations of surviving lineages. However, scientists have yet to fully understand the consequences of more recent biological upheaval, such as the megafaunal extinctions that occurred globally over the past 50 kyr. New Zealand was the world''s last large landmass to be colonized by humans, and its exceptional archaeological record documents a vast number of vertebrate extinctions in the immediate aftermath of Polynesian arrival approximately AD 1280. This recently colonized archipelago thus presents an outstanding opportunity to test for rapid biological responses to extinction. Here, we use ancient DNA (aDNA) analysis to show that extinction of an endemic sea lion lineage (Phocarctos spp.) apparently facilitated a subsequent northward range expansion of a previously subantarctic-limited lineage. This finding parallels a similar extinction–replacement event in penguins (Megadyptes spp.). In both cases, an endemic mainland clade was completely eliminated soon after human arrival, and then replaced by a genetically divergent clade from the remote subantarctic region, all within the space of a few centuries. These data suggest that ecological and demographic processes can play a role in constraining lineage distributions, even for highly dispersive species, and highlight the potential for dynamic biological responses to extinction.     

Hepatitis B in a high prevalence New Zealand population: a mathematical model applied to infection control policy

Background

Chronic hepatitis B (CHB) is a vaccine preventable disease of global public health importance. The prevalence of CHB in New Zealand's Tongan population is over 10%, a level consistent with endemic infection, which contrasts to the low overall New Zealand prevalence (<0.5%). Despite the introduction of infant vaccination in 1988, coverage among Tongan children is estimated to be only 53%.

Aims

To estimate the population benefit of additional public health control measures besides ‘business as usual’ infant vaccination for hepatitis B in high prevalence populations.

Methods

A mathematical model of hepatitis B virus (HBV) transmission was used to predict future CHB prevalence in the New Zealand Tongan population under different infection control strategies.

Results

Prevalence of CHB is predicted to plateau at 2% in the New Zealand Tongan population if coverage remains at current levels, which are therefore insufficient to achieve long-term elimination of HBV. The critical proportion of immunisation coverage for elimination of the virus is estimated to be 73%. The effect of screening for HBV carriage and early disease management was unable to be quantified, but is likely to reduce the population burden of HBV infection and thus contribute to accelerating elimination.

Conclusions and recommendations

Mathematical models are a useful tool to forecast the future burden of CHB under a range of control strategy scenarios in high prevalence populations. Serosurveillance and targeted vaccination has similarly arrested HBV transmission in time-series prevalence studies from Taiwan and Alaska. Such a policy may demonstrate similar efficacy in New Zealand ethnic groups with endemic HBV infection.     

Persistence and stability of lotic invertebrate communities in New Zealand

1. Persistence and stability of lotic invertebrate communities were determined at an annual time scale over a 9‐year period (1990–98) at 26 river sites over the northern half of New Zealand. In addition, a number of water quality variables were measured monthly and flow information collected continuously over the same period at each site. 2. The aim of the study was to describe the levels of interannual variability in invertebrate communities, and relate community changes to variability in environmental conditions. The consequences of this temporal variability for the sensitivity of predictive models were also investigated. 3. Levels of change in environmental conditions varied significantly between years, but were relatively similar across sites. In contrast, community persistence (similarity between years in species assemblage composition), and stability (similarity between years with respect to relative abundance of species in the community) both varied significantly between sites, but changes between years were similar. Community stability was highest at sites with relatively harsh flow conditions (high coefficient of variation, high relative size of floods), and was also greater in communities dominated by Ephemeroptera. 4. Relationships between change in environmental conditions and changes in community composition and structure were relatively weak for most individual sites. However, when average levels of change for each of the 26 sites were used, communities showed greater persistence under conditions where flow conditions remained relatively constant. Water quality changes had no significant effect on community persistence when assessed for all 26 sites combined. 5. Results from this study suggest that lotic invertebrate communities fluctuate around a relatively stable state, at least over a 9‐year period. However, the extent of interannual variation in community composition and structure observed, along with the relatively low degree of cluster fidelity observed within a single region, suggests that predictive models based on reference site conditions extrapolated over several years should be applied with caution in New Zealand streams.     

Distribution of macroinvertebrates in relation to physical and biological variables in the littoral zone of nine New Zealand lakes

Macroinvertebrates play a key role in the littoral zone of lakes. Macroinvertebrate community composition is closely linked to habitat conditions. To date, there have been few attempts to relate macroinvertebrates to habitat factors in lakes. In this study, nine mainly oligotrophic lakes from throughout New Zealand were surveyed for macroinvertebrates. The lakes were selected to represent a range of suspended sediment loading and lake level regimes. Within each lake, several sites were selected to provide a range of exposure to wave action. A multiple regression approach was taken to relate macroinvertebrate community composition and habitat characteristics. The results of the analysis suggest that the littoral zone of the lakes we studied could be divided into four general habitats. The first is the wave wash zone characterised by coarse substrates and macroinvertebrate taxa usually associated with lotic environments, such as Ephemeroptera and Plecoptera. The second habitat is associated with macrophytes and is limited at the top by wave action and at depth by light attenuation. In this zone, the snail Potamopyrgus antipodarum is dominant, along with Trichoptera and Odonata. At the base of the macrophytes is the detrital habitat characterised by fine, organic rich sediments and dominated by chironomids, oligochaetes and Trichoptera. At depths below the macrophyte zone, fine sediments are found, and bivalves such as the freshwater mussel Hyridella menziesi are common. While macroinvertebrate abundance can be highly variable, some general predictions of community structure can be made based on a few key environmental factors. Abundance of snails Odonata and Trichoptera was positively related to macrophyte biomass. Some macroinvertebrate groups such as oligochaetes, chironomids, snails and bivalves were more common in fine substrates, while Ephemeroptera were characteristic of coarse substrates. Detrital biomass was important for most of the macroinvertebrate groups studied showing a positive relationship for oligochaetes and Trichoptera and a negative relationship for Ephemeroptera and Plecoptera.     

Biogeographic area relationships in southern New Zealand: a cladistic analysis of Lepidoptera distributions

ABSTRACT. Recently, attention has been directed toward the application of cladistic techniques to reconstruct the history of areas from species distribution data. In this study, hypotheses of area relationships for southern New Zealand are generated from lepidopteran distribution data analysed at two taxonomic levels. Data are shown to possess cladistic structure and area relationships presented here are consistent with the geological history of the southern region of New Zealand. Our results suggest a recolonization of inland lowland regions from the south following a period of extinction during the early Pliocene. Analysis of selected data including only flightless or locally endemic species resulted in little resolution of area relationships but topologies were significantly congruent with a total species dataset. Hypotheses generated from this study are open to testing with congruence analysis using independent species phylogenies.     

Evolution of biological dispersal corridors through a tectonically active mountain range in New Zealand

Aim To assess the geological evolution and biogeographical implications of low mountain passes. In particular, we question the common biogeographical belief that major mountain belts form impervious physical barriers to biological dispersal, and that related taxa found on opposites sides of mountains are necessarily a result of vicariant tectonic processes. Location The Southern Alps of New Zealand form a long (500 km) narrow mountain belt at the oblique collisional Pacific–Australian tectonic plate boundary. High mountains were uplifted during the Pliocene (2–5 Ma) and uplift has continued to the present day. Methods We integrate previous work from several disciplines to obtain an overview of inter‐relationships between plate tectonic processes, geomorphology and biogeography along the main mountain barrier in New Zealand, and then extend this approach to other major mountain belts. Results The Southern Alps initially formed a barrier to at least some biological dispersal, including vicariant formation of separate species of freshwater non‐migratory galaxiid fish on either side. However, the high mountain barrier was breached in several places when passive transport of topography occurred, from the low‐erosion rain shadow on the eastern side towards the high‐erosion, high‐rainfall western side. This tectonic transport resulted in the capture of eastern rivers by west‐draining rivers, leaving low passes at the topographic divide. These low‐elevation corridors permitted biological dispersal across the mountains, although continued uplift raises these passes. A new set of passes has formed in the northern part of the mountains where younger faults are cutting across the older mountain topography. These potential dispersal corridors are becoming lower with continued erosion, and more common as the defining structures migrate southwards. Main conclusions Biological dispersal across the Southern Alps may be facilitated by numerous mountain passes, especially via the new passes formed by cross‐cutting faults. More low‐lying corridors existed than is readily apparent now, as old river capture‐related passes have been blocked by ongoing uplift. The dynamic mountain‐building and erosional environment typified by the Southern Alps occurs in all the world’s collisional mountain belts, such as the Andes, Himalayas, European Alps and North American Cordillera. Sister taxa occurring across mountain belts are not necessarily a result of vicariance driven by the rise of the mountains, as numerous passes may have permitted intermittent dispersal. The evolution of low passes may have been more prevalent than is currently appreciated, suggesting that topographically complex mountain ranges might be more effectively viewed as dynamic filters within a probability landscape rather than as static and impervious high‐altitude barriers to all but the rarest of biological dispersal events. In some cases, the biological disjunctions observed across mountains may more directly reflect habitat differentiation driven by orographic mountain development that has limited the probability of trans‐alpine dispersal success.     

Suitability of Agonopterix ulicetella (Lepidoptera: Oecophoridae) as a Control for Ulex europaeus (Fabaceae: Genisteae) in New Zealand

The larvae of Agonopterix ulicetella (Stainton) (Lepidoptera: Oecophoridae) feed on the green foliage of gorse, Ulex europaeus L., and this insect is a potential biological control agent of this weed in New Zealand. The biology of the insect is described and its known parasitoids are listed. In experiments to measure oviposition preference, 46 plant species from 11 families were exposed to adult moths. Gorse was highly preferred over other plants, and there was no oviposition on 33 species tested. Eggs were found on Spartium junceum, Chamaecytisus palmensis, Lupinus arboreus, L. polyphyllus, Genista tinctoria and occasionally on eight other species. In experiments to measure the ability of first instar larvae to feed on 70 test plant species, 59 did not support development beyond the first instar and only seven species supported development to the pupal stage. These results show that under laboratory conditions this moth can lay its eggs and complete development on five members of the tribe Genisteae other than gorse. A. ulicetella was released in New Zealand in 1990 but has not yet established.     

A comparison of realised niche relations of species in New Zealand and Britain

Summary Many plant species prominent in the native vegetation of the dry shingle banks at Dungeness (Britain) are also prominent as exotics in the dry Upper Clutha catchment (New Zealand). To examine the realised niche relations of these species, vegetation was sampled in the two areas. Inverse classification and ordination were used to determine the relative beta niches of the species in the two areas. There was little agreement; it seems that the exotic species in the Upper Clutha were pre-adapted to different niches from those in their native range.     

Dendroprovenancing: A preliminary assessment of potential to geo-locate kauri timbers in northern New Zealand

After 1840, New Zealand kauri (Agathis australis) became the focus of a large-scale timber industry in the upper North Island, which converted trees into timber that was then used in construction. Cities such as Auckland and Wellington relied on importing kauri to meet local demand for construction and other uses. Kauri timbers from buildings and in-ground features, mostly in Auckland city, have been collected for tree-dating and master chronology development. Although the use of timber at archaeological sites is understood, the geographic origin of the timber from within the natural range of kauri (north of 38 °S) is largely unknown. This limits interpretation of archaeological wood and constrains use of the tree-ring data in dendroclimatology. In this paper, we consider the potential to provenance kauri timbers used in 19^th and early 20^th century New Zealand buildings, using a combination of documentary sources, t-value mapping, and refined statistical matching using spatial patterns of correlations. Analysis of documentary sources for a test period of 1861–1865 CE indicates that there is sufficient information about the kauri timber industry to provide a geographic context for provenancing kauri in the upper North Island. The use of t-values for provenancing may be confounded by the relatively small size of the kauri growth region and a lack of sufficient spatial differentiation in growth patterns between the sub-regions. However, a new approach of using indexed residual chronologies and field correlations has promise for geo-locating timber in a relatively small region. The research highlights the value of establishing kauri provenance to New Zealand dendroclimatology, dendroarchaeology and environmental history.     

Population density and geographical range size in the introduced and native passerine faunas of New Zealand

The current avifauna of New Zealand comprises species with two distinct origins: those that evolved in New Zealand or colonized naturally from neighbouring landmasses, and those that were deliberately introduced to the islands by European settlers. Elsewhere, it has been shown that for species introduced to New Zealand from Britain there is a positive interspecific correlation between the geographical range sizes attained in both countries. Since positive relationships between abundance, measured either as population size or density, and geographical range size are a near ubiquitous feature of assemblages of closely related animal species, this suggests that species’ abundances may also be so correlated between the two countries. Here, data for 12 passerine bird species introduced to New Zealand from Britain are used to compare population densities and density–range size relationships in their native and alien ranges. In addition, the density–range size relationship for 12 passerine bird species that can be considered native to New Zealand is compared to that for the introduced species. The geographical range size and the mean and maximum densities of introduced species in New Zealand were significantly positively correlated with those values for the same species in Britain. However, in no case was the relationship between mean density and range size significant. While not statistically significant, density–range size relationships for introduced species are similar in New Zealand and Britain, but those for introduced and native species in New Zealand are quite different. Implications of these patterns are discussed.