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The Ecological Consequences of Social Wasps (Vespula Spp.) Invading an Ecosystem That Has an Abundant Carbohydrate Resource

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The Ecological Consequences of Social Wasps (Vespula Spp.) Invading an Ecosystem That Has an Abundant Carbohydrate Resource Biological Conservation 99 (2001) 17±28 www.elsevier.com/locate/biocon The ecological consequences of social wasps (Vespula spp.) invading an ecosystem that has an abundant carbohydrate resource J. Beggs * Landcare Research, Private Bag 6, Nelson, New Zealand Abstract Introduced Vespula wasps have successfully invaded beech (Nothofagus) forests in New Zealand. By collecting honeydew, an abundant carbohydrate resource, wasps can reach high numbers. Honeydew is produced by an endemic scale insect which infests about 1 million hectares of land, 15% of New Zealand's remaining native forest. At the peak of the wasp season, honeydew beech forests had an average biomass of about 3.8 kg of introduced wasps haÀ1 (10,000 workers haÀ1). These abundant invaders reduce the standing crop of honeydew by more than 90% for 5 months of the year and so compete with native species (such as birds and invertebrates) that also consume honeydew. The behaviour of three species of native bird is aected by this reduction in honeydew, but it is unknown if this aects the survival or reproductive success of these birds. Nevertheless, to avoid altering the birds' beha- viour, wasp density should not increase above a level where wasps revisit honeydew threads more than once every 180±400 min. Additionally, the predation rate of wasps on some invertebrate prey species is so high that the probability of an individual surviving through the wasp season is virtually nil. Hence wasps probably reduce or eradicate populations of some invertebrates. Wasp abundance needs to be reduced by 80±90% to conserve some native invertebrate species. Wasps could also aect nutrient cycling in the honeydew beech forest community by reducing the ¯ow of carbon to micro-organisms in the phyllosphere and the soil, which ultimately could aect soil solution chemistry. Current control tools are unable to reduce wasp populations over large tracts of forest. The challenge is to identify and develop new control techniques to achieve widespread control for conservation gains. The impact of introduced social wasps provides a warning of the damage exotic ants could cause if they were to invade honeydew beech forest. New Zealand needs to be vigilant to reduce the risk of an invasion by ants or other social wasps. # 2001 Elsevier Science Ltd. All rights reserved. Keywords: Biological invasion; Competition; Predation; Ecosystem processes; Honeydew; Vespula 1. Introduction vascular plants (Ministry for the Environment, 1997). Some of these aliens have become threats to native spe- Invasion biology and biosecurity is of increasing cies and ecosystem functioning. importance internationally as people and their asso- Increasingly, land-managers have to decide which ciated pests encroach on virtually every habitat and species (or genetically modi®ed species) are safe to ecosystem. The ecological processes and biodiversity of import, and which species should take priority for con- many invaded habitats and ecosystems are under threat, trol eorts. Such decisions are hampered by the lack of in part because of the invasion of exotic species (e.g. a robust theory of invasion biology (Townsend, 1991). Loope and Mueller-Dombois, 1989; Ramakrishnan and To build such a theory requires a greater knowledge not Vitousek, 1989). Throughout New Zealand, more than only of what makes a species invasive, but also how 4300 introduced species have become established in the introduced species aect native species (Simberlo, 1990). wild, including 90 species of vertebrates, perhaps 2200 Certain classes of introduced species are prone to species of invertebrates, and more than 2000 species of have eects throughout the receiving community (Sim- berlo, 1990). Many social insects are in this category because they are polyphagous, and their reproductive * Corresponding author and dispersal characteristics make them particularly E-mail address: [email protected] eective invaders (Moller, 1996). They can pose a threat 0006-3207/01/$ - see front matter # 2001 Elsevier Science Ltd. All rights reserved. PII: S0006-3207(00)00185-3 18 J. Beggs / Biological Conservation 99 (2001) 17±28 to their host community, causing changes in community coated with a black sooty mould [probably Capnocybe composition (Howarth, 1985; Wojcik, 1994; Moller, novae-zealandiae (Hughes); Morales et al., 1988] which 1996), and this has the potential for ¯ow-on eects to is maintained by the honeydew. The mould provides ecological processes. food for many arthropods (Morales et al., 1988), which Two species of introduced Vespula wasp have estab- in turn are consumed by other animals. It is probable lished in New Zealand Ð German wasps [Vespula ger- that honeydew promotes the growth of a variety of manica (Fabr.), Hymenoptera: Vespidae] arrived in other micro-organisms. This has been well demon- 1945, but were displaced in some forests by common strated for micro-organisms in the phyllosphere and the wasps [V. vulgaris (L.)] which arrived in the late 1970s soil in northern hemisphere communities where honey- (Clapperton et al., 1989, 1994). Common wasps are now dew is present (Dighton, 1978; Dik et al., 1991; Stadler abundant in New Zealand beech (Nothofagus spp.) for- and MuÈ ller, 1996; Stadler et al., 1998). Micro-organisms ests infested by the honeydew-producing native scale aect several ecosystem processes in Europe, such as insects Ultracoelostoma assimile (Maskell) and U. brit- throughfall of dissolved organic carbon, and nitrogen tani (Morales; Margarodidae: Homoptera; Thomas et ¯uxes (Stadler and Michalzik, 1998; Stadler et al., 1998; al., 1990). The term `honeydew beech forest' refers to Michalzik et al.,1999). It is possible that honeydew beech forest infested with the scale insects. Honeydew, plays a similar key role in nutrient cycling in New also an important food resource of many native verte- Zealand honeydew beech forests. brate species (Gaze and Clout, 1983; Clout and Gaze, Honeydew is by far the most abundant nectar-like 1984; Beggs and Wilson, 1987; Moller and Tilley, 1989), resource in beech forest and is consumed by a range of provides an abundant food resource that allows social birds, lizards and invertebrates (Fig. 1; Gaze and Clout, wasps to reach very high numbers and have a major 1983; Clout and Gaze, 1984; Beggs and Wilson, 1987; predation impact on some native invertebrate species Moller and Tilley, 1989). For instance, kaka (Nestor (Harris, 1991; Toft and Rees, 1998; Beggs and Rees, meridionalis meridionalis), a native parrot, can obtain 1999). Therefore, wasps may also have a competitive their daily energy requirement by feeding on honeydew eect on native species, especially birds. for about 3 h (Beggs and Wilson, 1991). In the northern Common wasps in honeydew beech forest have been hemisphere more than 250 invertebrate species are the focus of more than a decade of research to docu- known to consume honeydew (Zoebelein, 1956), but ment their spread, population dynamics and impact, there are relatively few published accounts of native and to develop tools to reduce their abundance. While invertebrates feeding on New Zealand honeydew. In the initial impetus for the work was fuelled because of many parts of the world ants form the dominant group their direct eect on humans, there is now increasing in the guild of invertebrate honeydew consumers (HoÈ ll- concern over their impact on natural habitats. The dobler and Wilson, 1990). In New Zealand ants do feed principal cause of this impact in honeydew beech forest on honeydew, but they are not numerous (personal is the plundering of a key resource Ð honeydew. The observation), nor has their relationship with honeydew aim of this review is to summarise what is known about the impact of wasps in beech forests, to identify critical gaps in our knowledge, and to identify important les- sons from the wasp invasion of New Zealand, particu- larly with respect to potential future invaders. 2. The honeydew resource Honeydew is the sugary exudate produced by scale insects that infest black beech (Nothofagus solandri var. solandri), mountain beech (N. solandri var. cliortioides), red beech (N. fusca), and hard beech (N. truncata), but rarely silver beech (N. menziesii). Encapsulated second and third larval instars insert their stylet into the phloem vessels of the host tree, and excrete excess carbohydrate via a waxy anal ®lament protruding from the tree (Mor- ales et al., 1988; Morales, 1991). This honeydew is rich in carbon (mostly fructose, sucrose, glucose and oligo- Fig. 1. Overview of the major interactions in¯uenced by introduced Ves- saccharides) but low in protein (Grant and Beggs, 1989). pula wasps in beech forest infested with honeydew producing scale insects. Honeydew provides a major energy source at several Many of the interactions involve predation and competition. Dotted lines trophic levels (Fig. 1). Honeydew trees are typically indicate interactions that have received little or no scienti®c study. J. Beggs / Biological Conservation 99 (2001) 17±28 19 been studied. Furthermore, there are only a few species (6 300 000 ha; Newsome, 1987). Increasing the rainfall of native ant in New Zealand (Valentine and Walker, limit by 500 mm per annum only slightly increased the 1991). In many New Zealand honeydew beech forests, area estimate (by 2%), but decreasing rainfall by 500 the most dominant honeydew feeders are now intro- mm decreased the area estimate by 18%. This lower duced social wasps (Moller and Tilley, 1989). Other limit excluded most of the West Coast (where honeydew exotic insects feeding on honeydew can also become is present, e.g. Kelly, 1990), so a 3000 mm rainfall limit numerous, e.g. feral honeybees (Apis mellifera (L.), gives a distribution closer to the observed one. I have Hymenoptera: Apidae) up to 21.7 mÀ2 tree trunk and observed the scale insects at higher altitudes than the bumble bees (Bombus spp., Hymenoptera: Apidae) up criterion used here (800 m a.s.l.), so the estimate may be to 4.2 mÀ2 (Moller and Tilley, 1989).
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