Our Immigrant Insect Faunal By REECE 1. SAILER Dept. of Entomology and Nematology, University of Florida, Gainesville 32611 From the time of Asa Fitch in the .mid-19th century American entomologists have known that many of our most serious crop pests were not native to North America. It is also common knowledge that a very large number of crop pests, present in other countries, have not yet gained entry to the United States. Each year the U.S. Department of Agriculture spends several million dollars on plant quarantine activities designed to insure that these foreign pests are not added to our North American fauna. species that must be added to my list. Also a comparison of numbers shown on my charts and graphs will reveal certain discrepancies. These result from incomplete information concerning many of the species included in the list. Before commenting further on the foreign element of our fauna I would like to say a few words about the native fauna. The most recent estimate I have seen indicates that there are 104,000 named species of insects resident in the 48 contiguous states. Thus, the introduced species comprise only slightly more than 1.3% of the total fauna. Certainly we also have reason to be interested in the origin and history of the native species for, as indicated later, these indigenous species have and will continue to influence the ability of foreign species to successfully invade the United States. In view of the importance of foreign species as pests and of other foreign species as natural enemies of insect pests and weeds or as pollinators of many crops, it would seem highly desirable to know how many are present in the United States. For many reasons it is also desirable to know where they came from, how long they have been here, and how they arrived. Also, while we are painfully aware of those species that are pests, and know something of those that are beneficial, few people, aside from taxonomists, are aware of the large number having little or no known importance. Unfortunately there is no single source from which to obtain such information. While much information is available, it is scattered through literally hundreds of publications. Again, thanks to the work of taxonomists, zoogeographers are able to tell us something of the history of this indigenous fauna. Some species have been present, or more likely descendent from species present, on the North American land mass when it fractured off and drifted away from the great super-continent known as Pangaea. Although a subject of much past controversy, most knowledgeable people now accept Continental Drift as a fact (Hallem 1973, Kurten 1973). Many biologists, including entomologists, now explain many faunal similarities between widely distant continental areas as the result of early occupation of contiguous portions of the ancient super-continent. For several reasons I doubt that more than a few North American species can boast of such ancient lineage. North America is believed to have started splitting off from Europe 150,000,000 years ago. The last land connection across the North Atlantic was broken about 30,000,000 years later. To assume that the many indigenous North American species known also to occur in Europe and Asia owe their present distribution to continental drift implies that they are living fossils unchanged for 120,000,000 years. About 5 years ago I decided to try to rectify this situation partially through compilation of a list of insects and mites, resident in the 48 contiguous states and known to be of foreign origin. Species to be included in the list were those believed to have become established REECE1. SAILER in North America subsequent to 1620. In assembling the list I wish to acknowledge the assistance of many people without whose help such a task would have been impossible. First and foremost are the taxonomic entomologists of the Agricultural Research Service's Systematic Entomology Laboratory and their colleagues of the Smithsonian Institution's Department of Entomology. Secondly, there are the several secretaries and more recently, graduate students who have prepared file cards, tabulated data, worked up charts and graphs and typed manuscripts. Such an assumption also ignores the consequences of climatic change resulting from latitudinal and longitudinal position. It also ignores the catastrophic effects of glaciation that has repeatedly obliterated the biota of much of North America and Europe during the past million years (Flint 1957). In my opinion the intermittent land connection between North America and Asia known as Beringia has had a much greater influence on our insect fauna. Within the past 200,000 years Beringia was a broad 1200 mile wide isthmus of dry land connecting Asia and North America. At the same time much of present-day Alaska was ice-free while most of Canada and much of the United States was buried by a vast continental glacier. In this discussion it is not my intention to present lists of names, but rather the orders to which they belong, where they came from, how they arrived and what influence they now have as components of our fauna. I should also point out that while I have been able to record 1,385 species as of foreign origin, the list is still far from complete. Almost without exception each new monographic treatment or review of some sizeable genus or larger group of North American insects will include one or more 1 Pr~sid~ntial addr~ss : Entomological Washington, D. C., Nov=b~r 28, 1977. Society of Obviously, the climatic conditions prevailing in Siberia, Alaska and the connecting land bridge are critical considerations if this route is to be accepted as a major factor in explaining faunal relationships of North America and Asia. Even as dry land, the Bering Sea area is likely to be associated in the minds of most people with igloos and Eskimos mushing across the tundra behind a dogteam. Today the region is a formidable ecological barrier America, 3 ESA 4 Vol. 24, no.1 BULLETIN 1978 1300 V) 1100 w V w a.. V) 900 u.. 0 et: W d:l 700 ~ ::> Z UJ > - 500 I- 4: ...J ::> ~ ::> 300 V 100 1640 1700 1800 1860 1900 1940 1977 TIME Accumulation FIG. I.-Number of Immigrant Species of exotic species of insects and mites resident in the 48 contiguous states at time intervals from 1640-1977.' to insects of the temperate zones. But with cold water of the Arctic Ocean blocked off from the North Pacific, the southern coast of Beringia would have been swept by a warm ocean current with an effect similar to that of the present Atlantic Gulf Stream on the British Islands and Scandanavia. That plants and animals did move freely across this land bridge is evidenced by the present fauna and flora of Alaska. My own unpublished list of Alaskan Hemiptera shows that 80% of the species are of Old World origin. Clearly Alaska has served as a beachhead and staging area for invasion of North America by plants and animals that first evolved in Asia. There is another consequence of Asian species' access to Alaska. At the time these animals and plants migrated across Beringia, the vast North American ice sheets had obliterated or driven the New World flora and fauna far to the south. Later, as this flora and fauna followed the retreating ice sheets north, it found access to Asia blocked by the physical barriers of climate and the Bering Sea. But perhaps of equal or greater importance, northward movement of New World species was also blocked by the presence of the invading Old World species of plants and animals. The Bering land bridge has therefore functioned as a valve that tended to admit Asian species to North Amer'All figures prepared by Scott R. Yocom. ica but deny North American species access to Asia. As one consequence of this valve effect there are more than 500 species of Diptera in the United States that are also known to occur in temperate Europe and Asia, but are not now found in Alaska and northern Canada (Stone et al. 1965). These species almost certainly originated in the Old World and their presence now in central and eastern United States is strong evidence that the Bering Sea land bridge at some time had a climate not too different from that of the present day Smoky Mountain area of North Carolina. During the 15,000± years since the ice sheet retreated from our northern borders, the fauna and flora of North America settled into the biomes and life zones that characterized the continent when the first European colonists began to arrive on its eastern shores. Their arrival marked the beginning of an ecological revolution that has not only affected the biota of North America but even the physiography of the continent. Beginning with the Age of Discovery, geographic and ecological barriers that isolated islands, continents, and inter-continental areas have been breeched by commerce. In the 1;nited States natural ecosystems have been displaced by agro-ecosystems over much of the land surface. The remainder has in one way or other been influenced by our modern industrial civilization. Vol. 24. no.1 1978 5 ESA BULLETIN Now having in a most general and superficial manner covered the origin of the indigenous insect fauna of the 48 states, we will look at what has happened since 1620. I think we can safely assume that several species of insects were on board the Mayflower when it tied up at Plymouth Rock. Some would most certainly have gone ashore with the little band of 102 Pilgrims. There is a good chance that some of our stored product pests can trace their lineage to the Mayflower. If not, it is because they were eaten by the Pilgrims during the near famine conditions that the colony experienced in its first year. United States from all parts of the world. None-the-Iess, comparatively few foreign species had become established in the U.S. by 1800. Distance and time were still formidable barriers. Only species that lived on man himself or his domestic animals and those that lived in stored products could survive the trip. The one important exception were those long-lived forms carried in ballast of ships. As agriculture expanded, commerce increased and time in transit diminished, opportunities for foreign insects to gain entry and become established increased exponentially. Fig. 1, in at least an approximate manner, shows how the number of established foreign species accumulated during the past 350 years. According to my most recent and still far from complete list the total number now stands at 1385. This includes all categories-those that were introduced intentionally as well as those that arrived accidentally. The vertical axis of the graph shows number of species and the horizontal axis, time. The line connects points that show the number of species known to have been present at the end of each time interval. Because of the paucity of data, intervals of 20 years are shown up to 1860. After that date each interval is 10 years. Several things are immediately evident from the graph. Certainly reinforcements arrived with every ship bringing additional people and supplies from England. However, it is unlikely that any crop pests were among the foreign species that date from this early period. Most would not have survived the long sea voyage. Any that did would have found conditions for establishment of a population extremely hostile. Prospects improved as the human colonists expanded their numbers and established gardens, farms and orchards. As the human settlements grew, the trickle of commerce between Europe and America turned into a torrent. At first limited to England, commerce was soon established with other European countries and by 1800 commodities were reaching the Major Orders ~o~o2o - Hymenoptera: -- Coleoptera: ~ Lepidoptera "0"'0"'0 '0°0°0 300 Homoptera : 272 295 : ;~~~1.Acarina: al ~ w U w 79 80 : 86 Heteropt era 55 200 Q.. (/) Minor Orders I.L. 0 ~ Orthoptera 27 Psocoptera 11 Thysanura w a:l Dermapte ~ :::l Neuroptera Z 120 Thysanoptera ............. Diptera V) 327 100 : 11 ra 8 4 Anoplura : 2 Araneida : 2 E mbioptera Isoptera: : 2 2 Siphonaptera Total 2 = 1385 ORDERS Immigrant FIG. 2.-Number states. Species of species In each of the Orders represented by Order in the exotic arthropod fauna of the 48 contiguous Vol. 24, no. 1 1978 ESA BULLETIN 6 100 RESIDUAL VI w THYSANO 90 HETEROPTERA U w a... VI ORDERS PTERA 01 PTERA 80 HOMOPTERA lL. 0 LEPIDOPTERA 70 ACARINA c:: w ~ ~ ::l Z ....• HYMENOPTERA 60 COLEOPTERA 50 <l: I- 0 I- 40 •..•... 0 30 I- Z w U c:: w a... 20 10 0 1820 1840 1860 1880 1900 1920 1940 1960 1977 TIME Change in Proportional Representation of Species FIG. 3.-Proportional by Order Since 1820 changes in Order composition of established exotic species at time intervals from 1840-1977. First there was a very slow accumulation of species until 1860. Between 1860 and 1920 number added during each 10-year interval increases almost exponentially. This is clearly correlated with an exponential increase in Commerce that occurred during this same period. In 1920, there is an abrupt change to a still ascending but straight line. This must reflect the deterrent effect of the Plant Quarantine Act of 1912. This act was not fully implemented until 1920 (Marlatt 1920). Since that time additional species have been recorded at the rate of about 12 per year. Some of these might well have been in the United States prior to 1920 and escaped notice until a later decade. Also, on an average, 3 of the 12 are intentionally introduced beneficial species. This leaves an increment of 9 accidentally introduced foreign species that are annually incorporated into our fauna. Obviously, plant quarantines have not proved an insurmountable barrier to invading species. However, the data also suggest that plant quarantines have prevented the entry of a very large number. Had foreign species continued to arrive at the accelerating pre-1920 rate, instead of 682, we might expect at least twice this number would have gained entry and, based on past performance, 20% or 286 would have been pest species of significant importance. These would certainly have included the Mediterranean fruit fly and many of the serious crop pests that so far have been unable to invade the U.S. Next, I would like to look at who these fellow immigrants are and how they got here. In Fig. 2, the number of species belonging to each of the Orders is shown by means of a bar graph. Here we see that Homoptera takes first place with more than 23% of the total number. Coleoptera with 21%, Hymenoptera with 20% and Lepidoptera with 9% include 65% of the remaining species. Diptera stands 5th in number of species, closely followed by Thysanoptera and Acarina. In aU, 17 orders are represented. Sixteen, if you belong to the school that places Homoptera and Heteroptera as suborders of Hemiptera. For obvious reasons, it serves my purpose here to treat theme separately, and since I would otherwise prefer to place them together under Hemiptera, I have used suborder designations. Fig. 3 indicates the changes that have occurred in the kinds of foreign insects gaining entry and becoming established during the past 350 years. This graph actually starts with those present in 1820 and shows the proportional representation of the major orders at each of the later time intervals. You will note the early dominance of Coleoptera. These are species that for the most part arrived in ship ballast. After 1860 Homoptera took the lead by a wide margin. The explanation for this remarkable invasion of homopterous species is obvious and well documented. Following the Civil War the steamship greatly reduced the time required to cross the Atlantic. At the same time, growing affluence of Americans created a demand for nursery stock available in Europe. This traffic continued to grow through the end of the century. At the same time private individuals and the U.S. De- Vol. 24, no.1 1978 7 ESA BULLETIN partment of Agriculture were searching the world for new crop plants. An indication of the magnitude of this traffic is provided by Fairchild (1945) who was head of the l:nited States Office of Plant Introduction. In a book about travel in the tropics he mentions that nearly 200,000 named species and varieties of plants were introduced from all over the world by this agency of the Department of Agriculture. I can now look over my list of immigrant species and confidently label at least fifty Homoptera as having been brought in by plant explorers. Well before 1900, entomologists of the U.S. Department of Agriculture and those of California and several Western other States were aware of the problem involving both nursery stock and plant exploration (Howard 1897). Efforts to obtain Plant Quarantine legislation were finally successful in 1912 but not fully operational until 1920. By administrative order, the USDA had instituted quarantine inspection of all plant material imported for Department use in 1906 (Weber 1939, p. 3). The effectiveness of these measures appears to be reflected by an immediate leveling off and later drop in the number of species of Homoptera recorded as newly established in the United States. In more recent years, the Hymenoptera have taken the Palearctic 721 c '.p> t.. 0 C ~ 0 -0 N c: 0 III ~ C ~ c :::> Origin FIG. 4.-0rigins of Immigrant Fauna of exotic arthropod fauna showing number of species contributed by each biogeographic region. 8 Vol. 24, 110.1 1978 ESA BULLETIN Oceania .<; Oc eania World FIG. 5.-Biogeographic lead. This, I am happy to say, is the result of intentional introduction of beneficial species. You will note an expansion in the Diptera band about 1910. This results from similar introductions primarily against the gypsy moth. The Acarina show a marked increase during the years 1910-30 and reflects a different factor-taxonomic activity. Prior to that time, there was no one able to identify mites. Work of one taxonomist active in this period resulted in the first reported presence of most of these foreign species. Without having delved beyond numbers and order names, we have now covered the questions of how many of what kinds, as well as when and how they arrived. Now we will see where they came from. Fig. 4 provides a pie graph showing the percent of species coming from the major biogeographic regions of the World. You will note that more than 50% came from the Western Palearctic. This means Europe. The Eastern Palearctic and Oriental Regions account for more than half the remainder. Fig. 5, a map of the biogeographic regions of the world, shows the location of these regions in relation to the United States. The number of species each has contributed is clearly correlated with similarity of climate and crops but even more with amount of commerce. Until Regions regions of the world. very recently most commerce has been with Europe. Since World War II, Japan and Southeast Asia have become increasingly important. However, the importance of Europe as a continuing source of invading species is indicated by data for species recorded as established since 1970. Of 68 listed for this period, 40 came from Europe while only 10 came from the Eastern Palearctic and Oriental regions combined. During the same period, II are recorded as coming from Mexico, the West Indies or other areas of Central and South America. Economic and Ecological Consequences In 1958 the English ecologist, Charles Elton, published a book that should be in the library of every entomologist. The title of this book is "The Ecology of Invasions by Animals and Plants." In this book Elton uses the term "ecological explosions" to describe the effect on both natural and agroecosystems that so often results from invasion of a system by organisms of an alien system. He draws on examples from all plant and animal groups as well as from all parts of the world. He concludes that thanks to the activities of one dominant organism-man, the entire world biota is undergoing a convulsive change more drastic than any that have occurred in the past million years. The United States has experienced more than its share of such explosive invasions. There is little Vol. 24, no.1 1978 ESA BULLETIN need to enumerate them here. The gypsy moth, European cornborer, Japanese beetle, oriental fruit moth, European pineshoot moth, and alfalfa weevil are examples. However, not all invading species cause disruptive effects in their new habitat. Actually, they run the gamut from catastrophic importance through no known importance to highly beneficial. Fig. 6 provides a rough indication of the economic impact of the immigrant species. Of the 1379 for which I have assigned economic importance, 236 are in the category of important pests-species that annually cause some monetary loss; while 566 fall in a minor pest category. By minor pests, I have in mind species that are known to feed on economically useful plants but seldom cause significant damage. Also included are species of nuisance value as well as species that are parasites of beneficial forms. Species of no known importance account for 21% of the total and an equal number are assigned to the beneficial category. The latter include parasites, predators, pollinators, dung feeders, and weed feeding species. With closer scrutiny of these species, several interesting aspects of the immigrant fauna are evident. First, the important pest species constitute only 350/0 of the estimated 700 species of important arthropod pests present in the U.S., yet this 35% includes species responsible for 50% of all insect pest losses. A second important point relates to quarantine strategy. If we had undertaken to predict the economic importance of each of the 236 spe- Number of Immigrant Species to the 48 States FIG. 6.-Economic 9 importance of exotic species showing number represented in each category. ESA 10 BULLETIN cies in advance of their arrival we would have been wrong 2 times out of 3, the reason being that we would have relied on evidence of economic importance in the countries of origin. As a result we would not have anticipated the importance of such species as the Japanese beetle, the citrus blackfly, and the cereal leaf beetle. From this knowledge it appears that accurate prediction of any foreign species' potential importance as a pest in the United States requires that we know what factors are responsible for lack of importance in the country of origin. Undoubtedly, these factors are, more often than not, natural enemies that will be left behind when the species invades the U.S. I can only conclude that any plant feeding insect is potentially dangerous to American agriculture if it is found in a foreign country to be associated with one of our crop plants. The beneficial immigrant species also fall into 2 subcategories-those that were intentionally introduced and those that came in as accidents of commerce. As an entomologist specialized in introduction of beneficial insects, I find it disconcerting that of 271 beneficial immigrants, 118 are accidental introductions. These include many valuable species such as the parasite of San Jose scale, Prospaltella perniciosi Tower, and the alfalfa seed pollinator, Megachile rotundata (F.). On the other hand, beneficial insect introduction activities during the past 90 years have contributed only 2 species to the injurious species list and these are of negligible consequence. Future Prospects Originally, I became involved in this immigrant species subject because of assignment to a task force charged with review of USDA Quarantine Inspection program. The objective was to determine how effective this program was and how it might be improved. I concluded that one means of determining effectiveness would be a retrospective view of our immigrant insect fauna. Secondly, it seemed that a comparison of the immigrant fauna of the 48 states with that of Hawaii would be useful. As indicated earlier, it is evident that quarantine restriction activities, while not altogether stopping entry of inj urious foreign species, have served as an effective deterrent and considerably reduced the probability that a foreign plant feeding species will gain entrance to the United States. It was with the view of determining the magnitude of this deterrent effect that I undertook the 48 states-Hawaii comparison. Hawaii has only 0.8% of the land area of the 40 contiguous states and essentially only one port of entry for foreign commerce. More man hours of quarantine inspection effort per unit of commodity are expended at Honolulu than at any of the other U.S. ports of entry. Other things being equal, one would reason that fewer foreign insects should gain entry to Hawaii than to the 48 contiguous states with their vastly greater area, more diverse environments, and a vastly greater amount of commerce that arrives through more than 30 ports of entry. However, from even a preliminary review of the data it is evident that other things are not equal. Instead of the annual increment of 12 foreign species for the continental states, 16 are being added to the Hawaiian fauna (Mitchell 1975). It is also of interest to look at the total insect fauna of the Hawaiian Islands in comparison with that of the 48 contiguous states. Zimmerman (1948) estimated that there were 6,000 species of insects in the Hawaiian Islands. Of these, only 3,722 were endemic, the remain- Vol. 24, no.1 1978 ing 2,278 being of foreign ong1l1. Thus, we find that Hawaii is not only acquiring new immigrant species more rapidly but already has almost 900 more than the 48 contiguous states. As another way of looking at the foreign element of the 2 areas, 38% of the Hawaiian fauna but only 1.3% of the 48 states has been introduced accidentally or intentionally by man. From this comparison of Hawaii and the 48 states I can only conclude that environmental resistance is a powerful deterrent to invading species. In fact, it would appear to be much more important than Plant Quarantine Inspection. By environmental resistance I have in mind the kind of problem facing the tropical insect that finds itself on the grounds of the Kennedy Airport in New York. Or, the plight of the insect hitchhiker that leaves a plane in Miami only to be confronted by a hungry horde of predatory ants, spiders, birds, and a variety of other potential enemies. Then, assuming the insect is lucky enough to have escaped the sharp eyes of the quarantine inspectors and the jaws of potential predators and to find a niche in which to live and start a family, it may still have a problem. The niche may already be occupied, leaving no room for the invader. Finally, there is another barrier that has received little attention but one I am convinced is important. This is inbreeding depression. Most colonizing species arrive in very small numbers. Relieved of natural enemies and with other environmental factors favorable, the invading species may increase its numbers rapidly for a few generations and then suffer a drastic reduction in fecundity, fertility, and general vigor. This I believe to be a consequence of the genetic load of deleterious recessive allelic genes carried by the founding individuals. As a result of inbreeding, a portion of the population will soon become homozygous for these deleterious recessive genes. The resulting depression of reproductive potential combined with environmental resistance may result in extinction of a well established population. At least 10 immigrant species once recorded as established have disappeared, possibly for this reason. In several cases, introduced beneficial species were involved. By comparison, only 6 species have been eradicated by regulatory action and one of these-the citrus blackfly-is back. Thus, in its efforts to prevent entry of new pests into continental United States, Plant Quarantine is really only the first line of defense. It is backed up by powerful natural deterrents. This does not mean that I depreciate the Quarantine effort. Quarantine activities have obviously reduced the rate of new entries and have prevented establishment of many potentially dangerous pests but it is still a numbers game. There is a certain probability that any foreign species capable of living in the United States will gain entry and become established. For some, the probability is so high that we can expect that they will be discovered and hopefully eradicated within a year or two. For other species the probability is so low that they may never make it. In the case of Hawaii, it appears that environmental resistance to invading species is so low that the Islands act almost as a faunal vacuum and will continue to do so until all available ecological niches are filled. Here, the barrier provided by Plant Quarantine is of much greater importance per unit of agricultural production than for the 48 continental states. Aside from protecting Hawaiian agriculture and the Islands' native fauna and flora from invading species, there is an additional reason Vol. 24, no.1 1978 ESA BULLETIN for maintaining the strictest possible plant Quarantine procedures in Hawaii. Because of their vulnerability to invading species, the Islands comprise a dangerous stepping stone and staging area for invasion of Asiatic species into the continental states. The hazard of invasion by foreign pests has also been greatly increased by the opening of such airports as those of Chicago and Kansas City to international traffic. Insect hitchhikers arriving at New York find a generally hostile urban environment. By contrast, a corn or SOybean insect landing at the Kansas City International Airport finds itself in the midst of corn and soybean fields. If the dangers posed by these internal centers of foreign traffic are to be minimized, they will require expenditure of a proportionately greater quarantine and surveillance effort. It would also be sound policy to establish land USe policies that would result in establishment of ecological barriers around such airports. Finally, my personal interest in immigrant species is their importance in the field of biological contro\. Based on both theory and experience, the discovery of a new foreign pest in the United States should trigger 2 kinds of response. (1) Immediate surveys and studies to determine feasibility of eradication and (2) initiation of foreign work needed to find and import natural enemies. If eradication succeeds-fine. The foreign work will generate information likely to be useful at a future date. If eradication does not succeed, several years lead-time on eventual control will be gained. Based on past performance, foreign species of insects will continue to gain entry to the United States. Improved quarantine inspection procedures may reduce the number, but more likely, any increase in the effectiveness of quarantines will be offset by the effect of increased Commerce and tourist traffic with foreign countries. Certainly, every economically feasible effort should be made to prevent additional species of insect pests from becoming established in the United States. In conclusion, it is my opinion that our best hedge against the dangers posed by foreign species is intelligence. The more information we have about the world insect fauna the more likely we are to be able to devise means of preventing entry of dangerous species and to take appropriate counteraction against any that do invade the United States. We need current information about the status of pests in other countries. But, of greater importance, we need detailed knowledge of the insects and mites associated with crop plants, forests, and other commodities in all foreign countries with which we have any significant commerce. Much of this knowledge is available 11 and needs only to be collated and incorporated into a computerized information bank from which it could be readily retrieved. There would remain a much larger body of needed knowledge that should be the subject of a special program. This would involve work in foreign countries either by American entomologists or through contract with foreign institutions. Insect taxonomists would play a key role in such a program. Such a program would not be cheap, nor completed quickly but the long-term savings to American agriculture would far exceed its cost. REFERENCES CITED Elton, C. S. 1958. The Ecology of Invasions by Animals and Plants. Methuen and Co. Ltd. London. Reprinted as a Science Paperback, 1977. Distributed in the USA by Halsted Press, a Division of John Wiley and Sons, Inc. New York. Fairchild, D. 1945. Garden Islands of the Great East: Collecting from the Philippines and Netherlands India in the Junk "Cheng Ho." New York. Flint, R. F. 1957. Glacial and Pleistocene Geology. John Wiley and Sons. New York. Hallam, A. 1972. Continental drift and the fossil record. Scientific American 227: 56-66. Republished as Chapter 9, p. 81-90 in "Eocology, Evolution and Population Biology. Readings from Scientific American." W. H. Freeman and Co., San Francisco. 1973. Howard, L. D. 1897. The danger of importing insect pests. Yearbook of Agriculture, 1897. P. 527-52. Kurten, B. 1969. Continental drift and evolution. Scientific American 220: 54-64. Republished as Chapter 10, p. 90-100 in "Eocology, Evolution and Population Biology. Readings from Scientific American." W. H. Freeman and Co., San Francisco. 1973. Marlatt, C. L. 1920. Federal plant quarantine work and cooperation with State officials. J. Econ. Entomo!. 13: 179-80. Mitchell, W. C. 1975. Pers. comm. In paper presented at 59th Ann. Meeting Pacific Branch, Entomo\. Soc. Am., Portland, Ore. June 24-26, 1975. Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson. 1965. A Catalog of the Diperta of America North of Mexico. Agric. Handb. No. 276. USDA, Washington, D. C. Weber, G. A. 1930. The Plant Quarantine and Control Administration-its history, activities and organization. Service Monograph of the U.S. Government. No. 59. Brookings Institution, Washington, D. C. Zimmerman, E. C. 1948. Insects of Hawaii. Vo!. 1. Introduction. Honolulu. HOUSTON MEETING DEADLINE The deadline for the submission of titles of presented papers at the Houston meeting is July 1. The submission form is printed elsewhere in this issue of the BULLETIN. For members planning a symposium, their ideas should be submitted to the appropriate Section Chairman by April 1, 1978. See the information sheet referred to above for more complete information. A reminder: The privilege of presenting submitted papers at an Annual Meeting of the ESA is limited to members in good standing only. ESA member sponsorship of non-mem- ber entomologists is not permitted. Not more than two submitted papers, including joint authorship, will be accepted from any member. No member shall be permitted to read more than one submitted paper. Non-entomologist members of related societies are permitted joint authorship with ESA members and are extended membership privileges on individually sub· mitted papers. Member and non-member abuse of the privilege of presenting papers at ESA Annual Meetings has necessitated the establishment of these requirements.
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