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The Canadian Entomologist The Canadian Entomologist Vol. 107 Ottawa, Canada, March 1975 No. 3 BIOLOGICAL CONTROL ATTEMPTS BY INTRODUCTIONS AGAINST PEST INSECTS IN THE FIELD IN CANADA B. P. BEIRNE Pestology Centre, Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia Abstract Can. Ent. 107: 225-236 (1975) This is an analysis of the attempts to colonize at least 208 species of parasites and predators on about 75 species of pest insects in the field in Canada. There was colonization by about 10% of the species that were introduced in totals of under 5,000 individuals, 40% of those introduced in totals of between 5,000 and 31,200, and 78% of those introduced in totals of over 31,200. Indications exist that initial colonizations may be favoured by large releases and by selection of release sites that are semi-isolated and not ecologically complex but that colonizations are hindered when the target species differs taxonomically from the species from which introduced agents originated and when the release site lacks factors needed for introduced agents to survive or when it is subject to potentially-avoidable physical disrup- tions. There was no evidence that the probability of colonization was increased when the numbers of individuals released were increased by laboratory propagation. About 10% of the attempts were successful from the economic viewpoint. Successes may be overestimated if the influence of causes of coincidental, actual, or supposed changes in pest abundance are overlooked. Most of the successes were by two or more kinds of agents of which at least one attacked species additional to the target pests. Unplanned consequences of colonization have not been sufficiently harmful to warrant precautions to the extent advocated by Turnbull and Chant but are sufficiently potentially dangerous to warrant the restriction of all colonization attempts to biological control experts. It is concluded that most failures were caused by inadequate procedures, rather than by any weaknesses inherent in the method, that those inadequacies can be avoided in the future, and therefore that biological control of pest insects has much unrealized potential for use in Canada. Introduction Biological control in Canada evolved recently and rather quickly from a distinct discipline that was applied virtually exclusively by government experts into a pest management procedure that is being applied increasingly by non-specialists (Beirne 1973). Most past attempts at permanent control of pest insects in the field in Canada by colonizing introduced biological control agents failed. Primary reasons for failures were that most of the introduced species failed to colonize and most of those that did colonize failed to lower the pest status of the target species significantly. The future use of biological control is liable to be limited if the high proportion of past failures was primarily because of weaknesses inherent in the method. But the potentialities for use are good if those failures were primarily because of avoidable human errors in applying the procedures. Attempts made against pest insects in the field in Canada over the past 90 years were analyzed from the viewpoint of determining which of those alternatives occurred, by deducing probable causes of successes or failures to colonize or to control pest damage. This analysis does not cover attempts made against pests in closed environ- ments such as greenhouses or against noxious weeds. These were in general more successful than the attempts against pest insects in the field. The analysis is based on information given in the reviews by McLeod (1962), McGugan and Coppel (1962), LeRoux (1971), and Beirne and Kelleher (1973), unless other sources are referred to. 276 THE CANADIAN ENTOMOLOGIST March 1975 The Colonization Attempts A colonization attempt is all the releases that were made of a particular species of introduced biological control agent. At least 208 species of parasites and predators of pest insects have been the subjects of field colonization attempts in Canada. The total is inexact because some attempts were inadequately recorded, some supposedly single species were later found to be several, and some individual species may have been released under several names. The immediate purpose of the colonization attempts was to establish pest-killing organisms as permanent residents of areas that they did not inhabit already. The attempts to colonize individual species fall into four classes on a basis of their outcomes, as follows. (1) Undetermined: attempts were made too recently to determine whether or not permanent colonizations have resulted. There are about 12 species in this category at time of writing. (2) Unclear: colonization attempts that were largely redundant because later information showed or strongly indicated that the introduced species were already present (at least 37 species). The introduction of new strains of existing species might theoretically have useful consequences by increasing their genetic diversity, but there does not seem to be a clear example of this type of consequence in Canada. (3) Failed: no evidence of permanent establishment (about 115 species, or about 55% of the total). Some might be still unrecognized successes, as was the sawfly parasite Pleolophus basizonius (Grav.) which was not recovered in some areas until 12 years after the most recent nearby attempts to colonize it. (4) Successful: apparent permanent establishment in that the introduced species were found several years after the most recent attempts to colonize them (up to 44 species, or about 21% of the total). Some, however, could have been the subjects of redundant colonization attempts that cannot clearly be identified as such. For instance, the apparently successful colonizations of four species of parasites of the holly leafminer Phytomyza ilicis (Curt.) must always be suspect because the existing parasites in Canada do not seem to have been surveyed before the colonization attempts and could have been imported unwittingly withP. ilicis within the holly leaves. Analyses indicate that the successes and failures were much influenced by quantities and origins of the material released and by characteristics of release sites. Influences of the Numbers Released The median total number released of all 159 introduced species, excluding species that were subjects of attempts with undetermined or unclear results, was about 5,000 individuals. The median was about 3 1,200 for the 44 species that became established, that is, colonized. Of the species released in totals of under 5,000, 10% (9 of 98) became established; of the 5,000 to 31,200 group, 40% (13 of 33); but of the over 31,200 group, 78% (22 of 28). It is clear that the greater the number released the greater the likelihood of successful colonization and that if the numbers were below some minimum (about 5,000 individuals) the probability of success was small. A release is all the individuals of a species that were liberated at a particular place and time. There are indications that the average number per release had an influence on the probability of establishment. Data on numbers per release are available for species introduced into forest e'nvironments up to 1958 (Graham and Jones in McGugan and Coppel 1962). Analyses indicate that about 60% of species that averaged over 800 individuals per release became colonized but only about 15% of those that averaged less than 800 per release did so. Volume 107 THE CANADIAN ENTOMOLOGIST 227 The same data may imply that the greater the number of releases per species the greater the possibility of establishment: about 70% of the forest species that were each released more than 20 times became established as compared with about 10% of those released less than 10 times each. However, these figures may be somewhat deceptive because early success or failure of a species to become established each generated its own form of feedback. Early successful colonization induced a conviction that the species was a "good" colonizer, thus encouraging additional and perhaps redundant colonizations of the same species. This tendency probably inflates estimates of the number of individuals that are normally needed to ensure successful coloniza- tion. Thus the median number of 31,200 individuals of successful colonizers is probably in excess of the minimal numbers actually required. Early failures of colonization induced a conviction that the species involved were "poor" colonizers, thus tending to discourage additional recolonizations. This tendency probably caused some colonization attempts to be abandoned prematurely, before the species was represented in the colonial environment by a large enough number of individuals for colonization. These feedback processes, however, did not occur sufficiently extensively and frequently to account for the fact that releases of large numbers made establishment a probability and that releases of small numbers made it merely a possibility. It seems likely that many of the failures of attempts to colonize exotic parasites or predators in Canada were owing to inadequate procedures rather than to inherent deficiencies of the organisms. This implies that many of the failures of the past could be successes in the future if the species will be reintroduced in sufficiently large quantities to make colonization a probability. A principal reason why numbers of individuals should exceed a certain minimum in each colonial site if the probability of colony success is to be maximized is
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