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Competition Between the Introduced Wasps Vespula Germanica and V. Vulgaris in Honeydew Beech Forest, North-Western South Island

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Competition Between the Introduced Wasps Vespula Germanica and V. Vulgaris in Honeydew Beech Forest, North-Western South Island "'''''''k,",'''' between the introduced e'"'-"'-"=~= and y. ""-='''&=~ beech forest, north-western Island, New Zealand. Richard J. Harris ,;::: A thesis submitted in partial fulfillment of the requirements of Doctor of Philosophy in Zoology, University of Canterbury 1992. Hz .:; I ~ c( Z To Barbara and Brian I TABLE OF CONTENTS Abstract Chapter Introduction 1 Chapter Study sites Documentation of competition between Vespula vulgaris and V. gennanica in honeydew beech forest The influence of habitat use and foraging on the 29 replacement of Vespula gennanica by V. vulgaris The protein diet of Vespula vulgaris and V. gennanica in 38 honeydew beech forest Chapter Honeydew foraging: harvesting a limited resource 53 Chapter 7: Variation in the quality of Vespula vulgaris queens and 75 its significance in population dynamics Chapter 8: Population dynamics and competitive exclusion among 98 vespid wasps in honeydew beech forest: a conceptual model Acknowledgments 108 References 110 Appendix 1: Harris, R.J. 1991: Diet of the wasps Vespula vulgaris and 119 V. gennanica in honeydew beech forest of the South Island, New Zealand. New Zealand Journal of Zoology 18: 159~ 169. Appendix 2: Harris, R.J.; Thomas, C.D. & Moller, H. 1991: The influence 130 of habitat use and foraging on the replacement of one introduced wasp species by another in New Zealand. Ecological Entomology 16: 441-448. Appendix 3: Harris, R.J.; Moller, H. & Tilley, J.A.V. 1991: Weather- 138 related differences in attractiveness of protein foods to Vespula wasps. New Zealand Journal ofEcology 15: 167-170. Harris, R.I. 1989: An entrance trap to sample foods of 142 social wasps (Hymenoptera: Vespidae). New Zealand Journal of Zoology 16: 369-371. Prey items taken to four nests by V. VUlgaris foragers at Spooners Range in 1989. ABSTRACT New Zealand was colonised by the German wasp, Vespula germanica (F.) in the 1940s and it became established throughout the country, The common wasp, V. vulgaris (L) colonised in the late 1970s and is still expanding its range, Wasps are now common in honeydew beech forests of the South Island and can reach high densities, The behaviour and ecology of V. vulgaris and V. germanica were studied from 1988 to 1991 to investigate competitive interactions in honeydew beech forests of the north-western South Island. v: germanica disappeared from areas of honeydew beech forest within 3-5 years of the first arrival of v: vulgaris and has not returned to them. A change in the relative abundance of the two species occurred in spring following the arrival of V. vulgaris, and was consistent with the predictions of Archer's (1985) model of wasp population fluctuation. In north-western South Island honeydew beech forests where both species occur, the more abundant species dominates honeydew trunks. Aggressive interactions may take place on this high quality, potentially defensible sugar resource. The two wasp species show different foraging patterns that provide the potential for local coexistence. Although both are generalist feeders, v: germanica is more commonly found foraging for protein amongst the forest litter, whereas v: vulgaris forages more on shrubs and tree saplings. Approximately 15% of foragers returning to sampled nests carried animal prey. The proportion of protein in the diets of wasps decreased sharply after rain. V. germanica collected more Orthoptera and large Hymenoptera, whereas V. VUlgaris collected more Hemiptera and Lepidoptera. An estimated 0.8 and 4.8 million prey loads, or 1.4 and 8.1 kg per hectare per year was taken into nests in western and northern South Island honeydew beech forests, respectively. Wasp foraging and rainfall reduced the honeydew standing crop. As standing crop decreased, more wasps occurred on honeydew trees, they became less active, spent more time lapping the tree surface, and fed at a slower rate. v: vulgaris was more active and fed more efficiently than v: germanica. III Variation in queen size was found between wasp colonies and sites. Small queens were present in juvenile queen populations but were under represented in the reproductive population suggesting that small queens have a lower probability of survival. A conceptual model of population regulation and species replacement involving an overcompensating control mechanism is presented. Variation in queen quality is a consequence of differential larval nutrition in autumn when the food resource fluctuates with changes in wasp density and environmental conditions. The competitive advantage that V. vulgaris has in harvesting honeydew means that foragers spend less time obtaining carbohydrate and more time meeting larval food demands. This increases the relative fitness of V. vulgaris queens, and results in V. vulgaris densities increasing relative to those of V. gennanica. Eventually, V. gennanica is eliminated from patches of honeydew beech forest. Introduction 1 CHAPTER 1: INTRODUCTION Competition w _____ ., Competition can be defined as the negative effect of one organism on another by consuming, or controlling access to a resource that is limited in availability (Keddy 1989). The presence of interspecific competition has now been established both experimentally and through observations in a great variety of both natural and artificial systems (see reviews by Weins 1977, Connell 1980, Schoener 1983, Roughgarden 1983, Keddy 1989). However, there continues to be often heated debate as to its role in structuring animal communities (Wiens 1977, Connell 1980, Schoener 1983, Roughgarden 1983). One of the reasons for this is that competition is exceedingly difficult to study in natural communities (Schoener 1983), partly because reduction or avoidance of competition is almost always advantageous (Pianka 1976). Many field studies have relied on comparing behaviour and resource use patterns of naturally occurring allopatric and sympatric species. However, such studies tend to be poorly controlled since other environmental factors that can be expected to affect an animal's behaviour and distribution (e.g. physical environment, predators, parasites) probably differ (Pianka 1976, Connell 1980). The results of field experiments indicate that a great variety of factors (e.g. survival, growth, foraging success, food type or size) can be affected by interspecific competition, although most can be categorised as either behavioural or physiological (Schoener 1983). Even when population size of a species is affected, the degree to which species composition of the community is altered is often not investigated (Schoener 1983). From the early laboratory work of Gause (1934, 1935) on yeast and Paramecium came the axiom that "complete competitors cannot coexist". This led to the development of niche theory, which predicts that competition will be stronger between specialists than generalists, and between closely related species and those occupying the same fundamental niche (Hardin 1960). It has also been argued that competition is Introduction 2 most symmetric among closely related taxa, but that similar and symmetric competitive abilities should lead to long term coexistence, since neither species will have a clear advantage over the other (Aarssen 1983, Agren & Fagerstrom 1984, Goldberg 1987). Keddy (1989) suggests a possible reconciliation of these two contrasting predictions. At one extreme very similar species coexist precisely because they are so similar (Keddy 1989) and stability comes from intraspecific competition exceeding interspecific competition (Pontin 1982). As the organisms become increasingly different, so the possibility of asymmetric competition increases, leading to dominance and competitive exclusion (Keddy 1989). As organisms become sufficiently different they coexist by means of niche differentiation. Although niche overlap (Le. the sharing of resources between two or more species) is clearly a prerequisite for exploitative competition, it is harmless in a competitive sense if the resources are not acting to limit the consumers (Hart 1983). Similarly, interference competition is unlikely to evolve unless there is potential for overlap in the use of limited resources (Pianka 1976). Ecological theory provides a number of suggestions as to the means by which potential competitors may avoid or mitigate this competition (Schoener 1974, Terborgh & Robinson 1986). Partitioning of resources and habitats may take place and can be measured by the degree of overlap in the size and type of food utilisation, timing of foraging activity, microhabitat use, foraging strategies, or by any combination of these factors. Field evidence of the competitive exclusion of one species by another, has been observed with certainty in the field on very few occasions (Varley et al. 1973, Pontin 1982). Most instances involve parasitic Hymenoptera that have been introduced successively into a region, and have competed for the same host species; e.g. Opius spp. in Hawaii (Bess et al. 1961), Aphytis spp. in Southern California (Varley et al. 1973). Examples have also been documented of one ant species replacing another when introduced into a new habitat (Way 1953, Haskins & Haskins 1965, Crowell 1968). In contrast, the absence of a species from a particular situation today may be a reflection Introduction 3 of "the ghost of competition past" (Connell 1980). so, actual mechanisms of replacement, must remain unknown, and can only be inferred from observation (Schoener 1983, Keddy 1989). To surmise that exclusion is caused by competition rather than by some other mechanism is virtually
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