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Benefits and Harm Caused by the Introduced Generalist Tachinid

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Benefits and Harm Caused by the Introduced Generalist Tachinid BioControl (2012) 57:277–288 DOI 10.1007/s10526-011-9437-8 Benefits and harm caused by the introduced generalist tachinid, Compsilura concinnata, in North America J. S. Elkinton • G. H. Boettner Received: 11 July 2011 / Accepted: 26 December 2011 / Published online: 24 February 2012 Ó International Organization for Biological Control (IOBC) 2012 Abstract We review North American research on Keywords Non-target impact Á Generalist Á Compsilura concinnata (Meigen), a highly generalist Biological control Á Host range tachinid parasitoid that was introduced in 1906 to control two invasive forest insects: gypsy moth and browntail moth. The impact on gypsy moth in natural Introduction populations in North America has long thought to be minor, a view confirmed by recent work in Canada and Classical biological control has long been a principal Wisconsin, USA. However, research with experimen- tool in the worldwide effort to combat the devastating tally created populations of gypsy moth suggests that effects of non-native invasive species. Classical it may be more important than previously realized. biological control involves locating natural enemies Studies on browntail moth in both naturally occurring of invasive species in their native range and releasing and experimental populations suggest that C. concin- them in the newly invaded habitat. The non-target nata is probably the main reason browntail moth effects of introduced herbivores on beneficial plants disappeared from most of its former range in North have been a concern for many decades in classical America. Research on giant silk moths suggests that biological control programs targeting invasive weeds C. concinnata has become the major source of (Wapshere 1989; Briese 2005). Extensive host range mortality among several species and may be respon- testing has been done since the earliest use of this sible for the notable decline in their densities that has technique. Non-target impacts on native insects are a occurred over the last century. C. concinnata has been much more recent concern. Formerly, there was little touted as a premier example of the generalist natural or no evidence that arthropod predators or parasitoids enemy that should be avoided in classical biological introduced as biological control agents against other control introductions, yet in the case of browntail moth invasive arthropods had any important deleterious its effect has been extremely beneficial. effects on non-target species. As a result, several authors concluded that the technique was generally safe and unlikely to have significant effects on non- target organisms (Coulson et al. 1991; Godfray 1995). Handling Editor: Kevin Heinz However, as pointed out by Howarth (1991), ‘‘absence of evidence is not evidence of absence’’ of such & J. S. Elkinton ( ) Á G. H. Boettner effects. In the last decade, several studies have Department of Plant, Soil and Insect Sciences, University of Massachusetts, Amherst, MA 01003, USA documented attacks on native non-target species by a e-mail: [email protected] number of introduced agents (e.g. Obrycki et al. 2000; 123 278 J. S. Elkinton, G. H. Boettner Henneman and Memmott 2001). As pointed out by predators and parasitoids of gypsy moth from its various authors (Lynch and Thomas 2000; Parry native range in Europe and Asia (Howard and Fiske 2009), however, it is relatively easy to document the 1911). Eventually, twelve introduced species of par- occurrence of introduced predators and parasitoids on asitoids were successfully established on gypsy moth native non-target species. It is much more difficult to in North America (Hajek 2007). One of these species demonstrate significant impact of these natural ene- was C. concinnata, a tachinid with a very broad host mies on the density of their non-target hosts. Our work range that has now been recovered from at least 200 on the effect of the generalist tachinid parasitoid, species of Lepidoptera and Symphyta (Arnaud Jr Compsilura concinnata (Meigen) on giant silk moths 1978; Boettner et al. 2000; Strazanac et al. 2001). The is one of the few examples where such impact has been broad host range of C. concinnata was well understood documented (Boettner et al. 2000). We used life table by the individuals involved in its dissemination, but its studies to show that C. concinnata is causing very high potential impact on native Lepidoptera was not a mortality and decline in densities of experimental significant concern in those days (Howard and Fiske cohorts on a number of our native giant silk moths 1911; Culver 1919; Webber et al. 1926). Introductions (Saturniidae) at several research sites in Massachusetts of C. concinnata and efforts to release it in other parts (USA). Follow-up research by other investigators has of the country against gypsy moth and other target confirmed our findings in some regions (Kellogg et al. species continued over much of the twentieth century 2003), but less so in others (Selfridge et al. 2007; Parry (Sanchez 1996). 2009). In addition, however, we have also shown that C. concinnata was probably responsible for the decline and extirpation of browntail moth, a serious Effects of C. concinnata on gypsy moth populations forest defoliator that invaded New England in the first decade of the twentieth century, but subsequently Despite the establishment of C. concinnata and eleven disappeared over most of its newly invaded range other introduced parasitoid species, gypsy moth has (Elkinton et al. 2006). Our review expands upon and continued to spread into southern and mid-western updates an earlier review (Elkinton and Boettner regions of the US. Frequent outbreaks have continued 2004) that focused on the impact of C. concinnata to occur throughout the introduced range in northeast- on giant silk moths. Here we add more recent work ern North America. Even in its native range in Europe on that topic and review additional research on and Asia, where gypsy moth has a richer fauna of C. concinnata in North America. Furthermore, we parasitoids and other natural enemies, outbreaks and add an important dimension to the story, which is the defoliation by gypsy moth occur in many, but not all, impact of C. concinnata on browntail moth. regions with forests of appropriate host trees. Few long term studies have been done to document the impact of parasitoids on gypsy moth anywhere in the world, and Introduction of C. concinnata to North America their overall role in gypsy moth dynamics remains ambiguous (Elkinton and Liebhold 1990). A study by The gypsy moth was introduced to North America in a Sisojevic (1975) in the former Yugoslavia appeared to suburb of Boston, MA, USA in 1868 (Forbush and show a classic host-parasitoid oscillation between Fernald 1896). By the late 1880s, it had become a gypsy moth and three tachinid species, of which serious defoliator in eastern Massachusetts, and this C. concinnata was one. The level of parasitism damage triggered a substantial eradication effort based recorded by Sisojevic and in other European studies on hand removal of gypsy moth egg masses and was notably higher than that observed for the same widespread application of lead and copper arsenate parasitoid species on gypsy moth in North America insecticides to infested trees. Despite this effort, the (Reardon 1976; Elkinton and Liebhold 1990). area infested by gypsy moth continued to expand A comprehensive eleven-year study of parasitism in across eastern New England in the first decade of the naturally occurring populations of gypsy moth in New twentieth century. The eradication effort was then Jersey (USA) was conducted by Williams et al. abandoned, and in 1905 the United States Department (1992). In that study, there was no hint of any direct of Agriculture embarked on a major effort to introduce or delayed density dependence of parasitism by 123 Benefits and harm caused by C. concinnata 279 C. concinnata on gypsy moth, and overall levels of highest densities (Fig. 1). We have confirmed these parasitism by this species never exceeded 20%. results in many subsequent experiments (e.g. Ferguson Parasitism by other species was also quite low and et al. 1994). It is important to understand, however, that only weakly density dependent. This study confirmed the data collected by Williams et al. (1992) represented the conclusions drawn by earlier investigators: that variation in parasitism in a single population followed parasitoids played a limited or equivocal role in the over several years (temporal density dependence), population dynamics of gypsy moth in North America whereas the data from our experiments represented (Campbell 1975; Reardon 1976; Elkinton and Lieb- variation in parasitism among plots or populations all in hold 1990). Several more recent studies have recorded the same year (spatial density dependence). Population parasitism of gypsy moths in populations along the ecologists have disagreed about the conditions under advancing front of the gypsy moth invasion in North which spatial density dependence leads to temporal America. Gray et al. (2008) recorded 32% parasitism density dependence, but most agree that the latter is by C. concinnata in one year among gypsy moths that required for a natural enemy to stabilize the densities of had recently invaded Wisconsin (USA). In another its host. We do not know why the levels of parasitism Wisconsin study, Hajek and Tobin (2011) recorded recorded by Gould et al. (1990)weresomuchhigher parasitism by C. concinnata that varied between 1.6% than that recorded by Williams et al. (1992), but we and 11.9%, Timms and Smith (2011) showed that suspect that it is due to differences in spatial scale. In the parasitism by C. concinnata on gypsy moth in Ontario, hectare-sized, experimentally created populations, we Canada varied between 2.5% and 5.9% in successive suspect that C. concinnata aggregated to the higher years. In all three of these studies, C. concinnata density populations from forest areas outside the plots.
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