THE LIFE HISTORY AND CONTROL OF CATTLE GRUBS, HYPODERMA LINEATUM DE VILLERS AND HYPODEmU. :SOVIS DE GEER by Donald K. Scharff A THESIS Snbmitted to the Graduate Faculty in partial fulfillment of the requirements for the degree of Master of Science in Entomology at Montana State College Approved: ~~teo 2 Bozeman, Montana _ October, 1949 (l ~-h . . ....'\} t"" ' -2- \ TABLE OF CONTENTS Page .ABSTRACT • 4 INT.BODUOTION • 5 • ACKNOWLEDGEMENTS • 6 LIFE HISTORY OF CATTLE GRUBS 6 .-. . HOSTS • OTHER HYPODERMA. IN CATTLE 7 9 DISTRIBUTION • • North America • • • South America • Europe • India • Africa Asia • Australia, Nev Zealand, Dutch East Indies Hawaii • • 9 10 ll 12 12 • 12 .. • • • • Swnma.ry • 13 13 13 13 14 EOONOMIC IMPORTA.ltCE General • • Damage caused by Heel Flies • Damage caused by ~ra.bs • NUMBER OF LARVAL INsr.A.RS • • 14 15 15 lS DIFFERENTIATION OF sPECIES • 19 :BIOLOGY ~"'D irABITS • ro Heel Flies Route of larvae through the body Larval method of penetration Larval development in the warbles • Larval emergence and pupation • Grubs • ro 21. 23 ~~ 26 SEA.SONAL VARIATIONS Adults • • 27 27 • q~s2q f6g66 -3Page 32 EFFECTS OF NATURAL FACTORS ON CATTLE GRUB POPULATIONS •• Climate Predators • • • • Parasites Natural resistance of host CONTROL • .. ... 32 33 5~ .... .... 35 Heel flies • • • • • • • • • Destruction of eggs and newly hatched larvae • • • • • Immunization • • • • • • . • . • • •. .. .... Preventing oviposition • • • • • • • Combating the grubs in the backs •• The Re~isites of good grub control Specific methods for controlling grubs •• The use of power sprays • • • • • • • • • • • • • • • • • • Tests of cattle grub sprays in Montana Materials and methods • • • • Dis~ssion of results • • • Number of treatments necessar.y • Proper timing of treatments .. LITERATURE CONSULTED ... . . ... . ... ... 35 35 37 37 ~ 41 43 46 46 47 49 51 52 -4- A:B STP.A.C!r !rhe literature on cattle grub biologies and control is reviewed and discussed, and certain criticisms and suggestions are added. Evidence shows that H. bovis, the northern or "late" cattle grub, is much more common in the-western and northwestern United States than bas :previously been ~ro.p:posed. Great difficulty was experienced in distinguishing between H. bovis and H. lineatum in the second (penultimate) larval instar. - -With both species of grUbs the indivi<:luals arriving at the backs in the early part of the season spent an average of three woeks longer in the warbles than those arriving there toward the end of the season. Year to year variations of 30 days in dates of peak grUb abundance are not uncommon. In any one season, variations of 60 days or more are not uncommon between individual herds of cattle in the same general area. TP~s is especially true in mountainous regions. !rhe use of power sprays, using rotenone as the insecticide, and directed at the grubs at the time they are in the warbles, appears to be the only practical method of controlling grubs in large range livestock regions. Pressures of 6oo pounds per square inch are more effective than lo\'rer pressures. The addition of specific wetting agents to the sprays shows promise of increasing control and should be thoroughly investigated. H. bovis is considerably more resistant to currently recommended sprays than is H. lineatum. Preliminary observations show that in parts of Montana twogrub treatments, properly timed, should contact more than 90 percent of the entire infestation of H. ~ and H. lineatum. The control of cattle grc.bs with sprays is unfortunately far from perfect and there is urgent need for much good experimental work on this problem. -5INTBODUCTIO!i During the years 194o to 1948, the interest in cattle grub control increased markedly in the northwestern United States. This increased interest was contemporaneous with the development of cheap and efficient high pressure liquid spray machines. Previous to this period, the recom- mended methods for grub control were not generally applicable to western conditions, where large, comparatively wild, of beef cattle are involved. infre~ently handled herds The hand wash and band dusting methods of grub control were impractical, if not impossible to use. With our present knowledge of cattle grubs and methods for their control, it a.PPears that the use of high pressure rotenone sprays is the most practical method of combating these pests in Montana. The spray recommendations of recent years advise that the animals should be treated at 30-da.y intervals throughout the cattle grub season, which in Montana, is approximately six months in duration (January through June). From the standpoint of beef management and husbandry practices, such a program is highly impractical. The prima.ry purpose of the present research has been threefold: 1. To compile and summarize the information on cattle grab biologies and control. 2. To develop a more practical treatment schedule for large herds of beef cattle. 3. To improve upon sp~ techniques and insecticide ingredients in order to increase the percentage of control from each insecticidal application. -6ACINOllLEDGEMENT S The writer gratefully acknowledges his indebtedness to Dr. J. H. Pepper for the proposal of the problem and for constant help and advice during its development • and to Dr. J. A. Callenbach and Mr. E. :B. Hastings for helpful critici~s and assistance. LIFE HISTORY OF CATTLE GRUBS The life histories of the common cattle grUb (HyPoderma lineatum De Villers), and the northern, or late grUb (EYpoderma ~De Geer) are generally quite similar. The adult heel flies appear from early spring to late summer, and lay their eggs on the cattle. The eggs are placed for the most part on the hairs of the legs and other lower portions of the animal's bod,y. In two to six days, depending upon the temperature, the eggs hatch. The young maggots immediately crawl down the hair to the skin, and burrow directly into the tissues. There is a period of about five months after the larvae enter the body, during which their movements are not too well known. Their travels seem to be exclusively through connective tissues. They 'ltJAY appear in the tissues of the various visceral organs and the spinal canal. A large majority of the larvae of H. lineatum congregate for _several months in the tissues of the gullet, and H. bovis larvae spend a similar period in the spinal canal. The larvae, nov about 15 millimeters in length, begin their final migration, again through the connective tissues. They eventually arrive in the subdermal tissues of the baekp where, almost immediately they make breathing holes through the skin. Cysts are formed around them, but the breathing holes are kept open. After a 6o-day period in the backs, they emerge, drop to the ground, and pupate. -7In one to three months, depending on weather conditions, the adult heel flies emerge to spend a brief but bu~ life terrorizing their bovine victims. HOSTS Cattle are the normal hosts of both species of grubs. :Riley (1892) reports that H. lineatum is not uncommon in American bison. Bishopp et al (1926, 1949) report .that bison harbor cattle gTobs, but do not seem to be as heavily infested as cattle. .Roudabush (1936), however, upon e%lllllina.tion of 13 bison slaughtered in Oklahoma, found.them infested with a total of 150 H. lineatum larvae. Lo (1935) reports that three to flve percent of the buffalo hides inspected from Shensi province, in northwest China, were infested with grubs. Goats and sheep are the occasional hosts of all stages of H. lineatum larvae in India, according to Bhatia (1934), Sen (1934), and 1939a). SoD! (1939, Bishopp et al (1926) and Ditmans (1948) cite several other in- stances of goat and sheep infestation. They were unable, however, to induce complete larval development of H. lineatum in experimentally infested goats or sheep. Barrett and Wells (1948) at necropsy recovered. less than 20 percent of H. months previously. lineatum larvae.tran~lanted into goats four Apparently none had reached the back. Horses are not infrequently found to be subdermally infested vi th H,ypoderma larvae, but there is little information as to whether they successfully complete their development. infestation and normal emergence of one H. Girard. (1939) reports a natural ~ larva from a horse in -8France. Tarlier (1930) and :Baker and Monlux (1939) report H. the brains of horses, causing nervous disturbances. and Monlux ( 1939) , 21 of infestation. k2!!!. in According to Baker 79 riding horses showed positive evidence of grub One percent of horses examined by Potemkin and VedernikoT (1944) were infested with one to four H. bovis larvae per horse, all in the second and third instar. Other infestations in horses are reported and described by Carpenter (1914), Pillars (1923), :Bishopp et al (1926), MOte (192S), Ono (193S), Zottner and Coste (1939, 1939a) and :Badanin and :Badaniva (1945). . Local stockmen report to the writer that grub infes- . tations very frequently render riding horses unusable for periods of several months, apparently because chafing of the saddle breaks the gl'tlb or its enclosing cyst, with the result that an extremely tender, elovhealing infection occurs. :Bishopp et al (1926) give a thorough discussion of grub infestation in man. "They are found usually in the subdermal tissues, where they produce what is often called 1 creeping m;yiasis 1 • There seems to be a tendency for the larvae to work upward and most of them are finally" extracted from the head, face, or upper extremities. These migrations are often extensive and rather rapid, accompanied by considerable pain. In some cases the larvae appear under the mn.cous membranes of the mouth. ir Three cases of HyPoderma infestation of the human eyeball are recorded by Granberg (1913); Von Schmidt zu Wellenburg (191g), and O'Brien and Allen (1939). Other descriptions of human m;yiasis caused by H. linea tum and H. bovis are given by Henricksen (1921), Engel (1923), Style (1924), Andre (1925), Herms (1925), Lanford and Warner (1925), Lawson (1929), -9Wegelin (1930), DosSantos (1932), PavlovSkii and Glazer (1932), Natvig (1935), Smart (1939) and Haseman (194s). The writer found no records of the completion of larval development in humans. Attempts have been made by Bishopp et a1 (1926), Barrett and Wells (194s), Roubaud and Perard (1924) and others cited by Bishopp et al (1926), . . to infest dogs, rabbits, guinea pigs, and mice with H. lineatum and!:_ bovis larvae. The maggots were recoverable up to four months after i&- festation, but all evidence pointed toward death of the larvae before they matured. OTHER HYPODERMA IN CATTLE H. crossi, normally occurring in goats, occasionally infests cattle in India, ~ccording to Patton ~ (1924), and Fletcher and Sen (1930). aeratum, also normally a goat parasite, is recorded in cattle from Cyprus by McDonald (1945, 1946). Infestations of this type are probably not common. H. Diana, a deer parasite, was found in cattl.e in Scotland by Mac Dougall (1934). Attempts at inoculating cattle with larvae of this spe- cies proved unsuccessful. DI STRIBUTIOli The distribution of H. bovis and H. lineatum has received, and apparently continues to receive, a minimum of attention. certainly not as much as the subject deserves. As discussed by Bishopp et al cattle grubs have been known in Europe for more than 200 years. (1926), In Amer- ica., interest in cattle grubs became evident during the latter part of -10the nineteenth centur,r • .A rather complete discussion of cattle gru.b distribution is given by Bishopp et al (1926). Since the date of that work, however, it appears that these pests have spread considerably. North America According to Hearle (1932) (1936), and Gibson and Twinn both spe- cies of grUbs are common wherever cattle are raised in Canada. H. lineatw:l has been common throughout all of the United. States for many years. Regarding H. bovis, Bishopp et al (1949) . . state that its dis- . tribution is limited to the northern states, that it is much more ahund.·ant in the northeastern states than elsewhere, and that it occurs only here and there in the northwestern states. Their map showing the probable southern limits of H. bovis indicates that it should not be found to any extent south of latitude of latitud.e 35° 4o 0 N. west of the state of Missouri, nor south N. east of the Mississippi River. that "both H. lineatum and H. ~are For Oregon, Lindquist and Roth (1926) (1947) Pfadt (1947) reports major pests of cattle in Wyoming". write, "Since Bishopp and coworkers show cooparatively few H: bov:is in the western United States, it is concluded that this species has become firmly established in this area durii:Ig the past 20 years". In Montana. the writer, in e:mmining a typical herd of yearling beef stock, found them to be infested with an average of 17 the spring of H. lineatum per animal and 194S. Sorenson (1945) 17 H. ~ per anima.l during reports that in northern Utah, 199 H. bovis were recorded. from a native yearling steer, and 45 H. bovis -from a native heifer. In California, according to Douglas and Furman -11H. ~is well established as far south as San Luis Obispo (1947), County (latitude 305 found N.). Also in California, Furman and Douglas cattle to be infested with which were H. Russo 35° 3.7 (1948), grubs per animal, 20 percent of E2..!!!· (1927) reports that ~ lineatum is one of the major pests of cattle in San Domingo. There are some areas in North .America where gra.bs do not ordina.ril;r occur. According to Gibson and Twinn (1936), large parts of Sea Island and Lulu Island, British Columbia, are free or nearly free of grubs. Bishopp (1915) states that the Red River Valley of the North in North Dakota and Minnesota is free of grubs. The southern two-thirds of Florida is uninfested according to Bishopp et al (1926). Mills et al {1946) report that R. lineatum does not occur in an area several miles in extent in the vicinity of Bozeman, Montana. The writer surveyed the above area and found its extent to be 20 to 30 sqaare miles. Porto Rico and Jamaica are reported to be free of cattle grubs by Wolcott Von Volkenberg (1932) and Ritchie (1919). (1922), No records are known to the writer of the occurrence of HyPoderma in Mexico or Central America. South America Two reports, from Brazil and Chile, by Palazzolo (1923) are negative, but Doe Santos (1932) (1916) and Sievers reports that two R. bovis iarvae were found. infestizl& a man in :Brazil. This would i.ndiCI'Lte the possibility that R. bovis has become established in South America. -12Europe ,!h ~and H. lineatum are reported £rom Cyprus and Italy to Norvay and Finland, and £rom Britain to the Urals. There is evidence to indicate that there may be some regions in Europe in which one species or the other is not economically important (BiShopp et al, Blagoveshchenskii and Pavlovskii, Bruisova and Pushina, 1932; 1925; Dcyenski, Faber, 194o; Kftrschner, Pacchioni, 1933; 1924; fanski and Obitz, 1931; Degtyarev, 1927; Kurtpinar, ?avlovskii, 1935; 1930; 1928; Boas, 1930; Gannser, 1947; Anonymous, 1928; Brassler, Dinulesco, 1923; Natvig, 1931; 1926; 193S; Doinikov, Gannser-Burckhardt, 1937, 1939; Pricelkov et al, 1924, 1925; Walton, 1926; 1948; Westermarck, Obitz, Roe, 1936; 1938; Ste- 1927). India According to Sen Soni (1938), (1934), Handa (1936), (1939) Chadha and Sen and H. lineatum is common in most of' northwest India, including Sind, Punjab, Northwest Frontier Province, and Baluchistan, and in the Kumaun Hills of' northeast India. H. ~is common in eastern India, including Bengal, Bihar, Orissa, and the Central Provinces. Grubs are widespread in dry, hilly, and sancy areas, and are practically absent f'rom the moister areas and ma.rsh;r districts. Africa Bishopp et a1 (1926), citing others, state that H. ~ occurs in Algeria and Egypt, and that larvae of H. lineatum have been collected in Mozambique, southeast Africa. After many introductions of Hnoderma into Kenya and South Africa, say Bedford (1926) and Lewis (1933), cat- -13tle grubs still have not suc.ceeded in establishing themselves. Asia Sonan (1927} reports that introduced H. bovis did not establish itself in Formosa. In spite of introductions, Lo (1935} states that cattle grUbs have not become a pest in Soochow, Nanking, nor in Central China, but that they are a pest in Shensi province, nor\hwes\ China. Ono (1933, 1936) and Ono and Yama.sa.ld (1934) found that both H. bovis and H. lineatum are common, but that H. lineatum is much more numerous and is a serious cattle pest in Inner Mongolia an~ Manchuria. Austmlia, New Zealand, Dutch East Indies Evidence presented by Miller(1922} and Bi8hopp et al (1926) indicates that HyPoderma have not become established in these areas. Hawaii Sweze.y (1933) gave the first positive evidence that H. lineatum . . had become established. Ten years later :Beaumont (1943) found that 26 percent of the cattle examined from the Island of Hawaii were inf'ested with H. linea tum. Summary With the possible exceptions of Mozambiqu.e, and :Brazil, no records are known to the writer of the occurrence of Hzyoderma farther south than about latitude 16° N. H. lineatum is recorded from San Domingo at 16°, Hawaii at 19o, India at 25°, and Cyprus at 35° north latitude. -1~ H. ~is reported from California at Africa and Egypt at 35°, India at 20°, and l!'reneh West 30° to 32° north latitude. It appears that cattle gru.bs have not a.s yet established themselves in economic numbers in the tropics or the southern hemisphere. Na.tvig (1937) reports H. lineatUill as far north as latitude 66° N. in Norway. ECONOMIC IMPO~CE General Biley (1890) aptly describes a gru.bby carcass, as follows: 1 ••• beyers of the highest class meat, who supply hotels and restaurants will not on any account pnrchase carcasses showing traces of warble attack ••• The appearance known as Licked....Beef, which, resnlting from the presence of the grub, may be described as a moist or ro.nning surface of a greenish-yellow color, is certainly unwholesome in look, if not in fact. The description of such meat as given in the Farmers' Review ••• is almost sufficient to tarn one against beef altogether.• Smith (1948) says, "When hides are removed from grub-infested cattle, there is a mass of yellowish gelatinous meat around the grub hole that must be trimmed out as inedible. In a slaughter test conducted for the National Live Stock Loss Prevention Board by a Chicago paCker, there vas a loss of $1,248.131 on 29 badl7 infested carcasses, and for the lot of 57 steers averaging 1,254 pounds live weight, the loss amounted to $22 per head. Twenty-nine of the 57 carcasses were lowered one full gra,de by the trimming-out process, constituting a loss of 2!~ per lb., or a total of $512.50. Many of the loins were so badly damaged tha.t the7 had to be boned which meant a loss of $10 per cwt. on 1~ of the carcass weight, or $1.10 per cwt. on the en.tire carcass. 0! the 57 cattle, 44 hides were graded f2 which amoulited to a loss of $37. The detailed breakdown of this damage shows that the loss of meat Talue is often far greater than the hide damage. This, to be sure, was a rather bad lot, but it does go to show vbat actual slaughter losses are in -15such cases. There was a waste of at least 14,000,000 lbs. of meat trimmed out of all grubby cattle carcasses in the United States in 1948 causing a loss of approximately $13,000,000 which includes degrading loin Cllts mutilated. u Smith (1948} estimates the annual loss due to cattle grUbs in the United States at well over $100,000,000. :Barret and Wells (1948) quote estimates as high as $300,000,000 per year. It is very difficult to arrive at a really accurate figure on total damage due to grubs because of the varied ways in which cattle are affected by them. Damage caused by Heel Flies l3ishopp et al (1926) say, "HYpoderma~ causes the most excite- . . ment to stoCk and hence, where it occurs, produces the greatest damage in this respect. Losses caused by this anno)"ance include marked reduc- tion in milk flow, failure to put on :flesh normally, mechanical inJU%7 due to the wild efforts of the animals to escape attacks, and not infrequent!)" loss from the miring down of cattle when they rosh into ponds or mn.d holes, or their injury or destruction when ronning over embankments or cliffs. 1 Bishol'P et al (1949) estimate that milk production is reduced 10 to 25 percent during periods of heel fly activit)". Roe (1938) states that in Cyprus during the heel fl7 season agricultural operations are impeded owing to the frequent impossibility of using cattle for draft purposes. Damage Caused b7 Grubs Soni (1938), checking weekly weights on five heavily infssted _and two unin:f'ested bulls, found that the infested animals lost weight from- ·.. -15aFebrua.ry until mid-March then graduall;y gained thro'llgh April (most of the grubs had emerged b;y mid-March). There was no appreciable weight loss in the two uninfested animals. Mohler kept on 16 (1939) describes an experiment in which weight records were cattle held in a fly proof encloBtlre, and 18 check animals. The experiment lasted about 2 years, and the results showed that the exposed animals gained an average of 514 pounds against 550 pounds for the protected ones. Gunderson (1945), reporting on weight tests on cattle, treated twice for grubs, at 219 head of grubby 30 da.;y intervals, found that they gained an average of 0.14 pounds per animal per dq over similar untreated gru.bb;y animals. Munro et al The tests ran for an average period of (194 7) 53 da1's. report that noninfested steers averaged 43 pounds greater gain per steer, during the wintering period, than grubby steers treated twice for the control of grUbs. According to Trullinger et al (1949), grtib control tests at the Oklahoma Experiment Station resulted in 0.1 pounds extra gain per dAy on steers and 0.5 pounds extra milk per dq on dairy cattle over check animals. Smith (1949) ~s, "In 1948 a State Experiment Station made a test in which 10 steers in one lot, sprayed with rotenone • • • gained on the average one-fourth pound each per da.;y more than the same quality of 10 other infested steers untreated fed the same ration. It is significaut also that the treated cattle required 10~ less grain per 100 lbs. of gain in weight than the untreated cattle during a 6-months' feeding period. 11 -16BiShopp et al (1926) cite work by Boas, in which a cow, daily producing 30 to 32 pounds of milk, increased the yield to 4o pounds a few days a:Cter SO grubs had been extracted from her back. They also cite an experiment by Sch8ttler and Glllser in which the grubs were half the animals in a feeder herd. extrac~ed from These animals showed a gain in weight of 5.16 percent over the infested portion of the herd. Moderate grub infestations do not materially affect the yield of milk in dairy cows, according to Julienne (1947) and Mohler (1948). . ~ ' ~ Ride damage by grubs is by far the easiest of all grub damage to evaluate. Practically all nations and countries with economic numbers of cattle grubs have figures which show the percentages of total hides that are grubby, the devaluation of hides caused by grubbiness, and the yearly loss to the leather industry from Hypoderma infestation. Gibson and Twinn (1936) give the following figures for western Canada: 25-~ percent of all hides harvested are grubby. 41.4 percent of all hides harvested during grub season are grubby. 70 percent of all hides harvested in April are gru.bby. Smith (1949) states that at least 33 percent of all cattle hides proanced in the United States contain :five or more grub holes and are sold at the agreed discount of $.01 per pound. These holes are in the thick- est and best part of the hide used for sole leather. Coppens (1913) cites a statement by Godfrind, chemist to the War Ministry at Brussels, to the effect that the wearing property of warbled hides as leather is only 30 percent of that of sound hides. 0 1Flaherty and McLaughlin (1930) and Simmons (1939) made histological studies of the effects of grubs upon cattle hides. They found that cellular infiltrations and grul> debris are commonly walled off in the process of healing. These leave weak -17spots in the scar tissue and corresponding weak spots in the finished leather. Simmons states that even a perfectly healed cyst presents a defective hide for leather production. This is due to the fact that in the healing process fibers are laid down which are more uniformly parallel to the epithelium than those in normal tissue. As pointed out by 0 1 Flaherty and McLaughlin, polydirectional fibers are the source of the tissue's strength, while unidirectional structures are a source of weakness. The penetration of the skin by the ne\~ly hatched larvae produces what Hadwen (1919) calls cypodermal rash. The points of penetration be- come extremely sore, tender and inflamed. They generally eXIl.de a yellow- ish serum. Simmons (1937), in a histological stu~ of esophagi infested with congregations of H. lineatwn, states that the larvae typically occur in the submucosa, causing inflammatory swellings into which there is a very striking infiltration of leucocytes. Sometimes, accar ding to Scilmid (1941), when the larvae are quite numerous, the swelling ~ become severe enough to hinder swallowing. H. ~ congregating in the spinal canal sometimes causes paralysis, myelitis, and other nervous disorders (Schmid, 1941; Sergent, 1939; Ano:nymous, 1947; Popescu-l3aran, 1939) .. There appear to be some variations in opinion on the matter of anaphylaxis. Hadwen and l3ruce ( 1917) report having produced severe anapby- laxis in cattle by injecting ~oderma extracts into the jugular veins. They also state that the same symptoms may be caused by breaking cypoilermal grubs and their qsts in the back. Hadwen (1922), describes 11 Eose -18- Fever", which he says, appears to be identical with eypodermal anaphyla::rls. Re cites numerous cases of it in Europe, and also warns against rough or careless methods of extracting grubs. MacDougall (1930) and :Blagoveshchenskii and Pavlovskii (1930) report that numerous grubs broken in the backs caused an anaphylaxis. The latter two authors also state that the subcutaneous injection of grub extract bas no effect on the animals. The fact that Jensen (1924) reports eight fatal cases of "Rose Fever" in Denmark after one year of intensive nationwide grub extraction, indicates that some da.nger of anaphylaxis may be expected from band extraction of grubs. liatvig (1937) observes that animals in poor condition are usually more heavily infested than healthy ones. pressed by stockgrowers, who not This is an observation often uncommo~ animals is conducive to grubbiness. e~ feel that poor condition in It bas not been determined definitely whether heavy infestations of grubs are the result of unthriftiness or the cause of it. 'l!he preponderance of evidence, however, as set forth in the preceding paragraphs, would indicate that gro.bbiness could be and probably is a contributing factor to the poor condition of the host. EmmER OF LARVAL IN&rARS The number of larval stages in R. · bovis and R. lineatum has been a matter of considerable argument for at least ha.lf a centur;y. de~cribed what were apparen~ly four stages. Ri1B7 ( 1892) Then for a quarter of a cen- tur;y, four stages were generally recognized (Xnipling, 1935) until Laake (1921) described what he considered to be anOther, fifth, stage. the publication of Laake's description of the supposed new stage Since in~ -19lineatum and Phibbs' (1921) description of a corresponding new stage in R. ~. five stages have been recognized by most investigators (Laake 1921, 1924; Warburton, 1922; Hadwen and Fulton, 1924; :Bishopp et al, 1926; Mote, 1928; Case, 1929; Carpenter, 1924; Ono, 1933; Knipling, 1935}. Knipling (1935) cites others who hold to the older theory of four and states further that investigators in Germa~ generally recognize three stages. He then convincingly shows, through numerous measurements of cephalopharyugeal sclerites, counts of the spines on various segments and measurements of spiracles, that what were formerly thought to be the first three stages are actually only one. Bishopp, Laake, and Wells (1936) nov reverse their former stand, and refer to three iarval stages, as- do L8a.ke et al (1942). Since 1935 most workers recognize only three stages (ROudabush, 1936; Natvig, 1937; Smart, 1939; Zottner and Coste, 1939; Snipes and Cooper, 1942; Mathysee, 1945; Lindquist and Roth, 1947; Westermarck, 1947). A few still hold to five: (Ono, 1938; Potemkin and Vedernikov, 1944; Stewart, 1944; Furman and Douglas, 1948). Dinulesco (1938) speaks of four stages. Rase~ man (1941) questions the three-stage theory in favor of one or tvo additional stages. DIFFERENI!IATION OF SPECIES Using Laake 1 s (192i) -methods, it has been consistently possible for the writer to differentiate between H. bovis and!!.:. lineatum in the first and last larval stages. Regarding the second (penultimate) stage Laake ( 1921) says, "••• the characters of the stigmal plates permit, with absolute certainty, the differentiation (of species) of the fourth (actually the second) as well as the fifth (third) larval insta.r 11 • The writer has been -20- consistently unable to duplicate Laake's success on second stage larvae from Montana. Natvig (1937a) also reports that it is sometimes difficult to distinguish between !h. lineatum and R. bovis by the characters of the posterior stigmal plates. BIOLOGY AND HABITS Reel Flies Adult heel flies generally emerge from their puparia early on bright sunny mornings (Carpenter, et al, 1914). They are sexually mature, and may mate within. an hour after emergence (Carpenter, et al, 1914; Bishopp et al, 1926; and others). Copulation lasts only a few minutes. Bishopp et a1 (1926) observed the l~ing of fertile eggs 20 minutes after copulation. ing. bo~ The mouthparts are degenerate and apparently incapable of function- The adult fly, upon emergence, bas enough energy reserves within its to last throughout its lifetime. Length of adult life varies, accor~ ing to different authors, from two to 25 days, depending on temperatures. Various figures are available as to the number of eggs laid. MacDougall (1935) reports that 6oo to SOO may be deposited; Gibson and Twinn (1936) estimate an average of 4oo per fly; Bishopp et a1 (1949) estimate 500 per fly. Oviposition takes place most commonly on warm, suney, quiet dqs. R. linea tum lays chiefly on the hairs of recumbent stock (Hadwen, 1917, 1919) , and has often been observed to actually place its eggs on the bairs along the ground line while standing on the ground. Several eggs are usually placed on a single hair by R. lineatum while R. singly. ~ places them The latter species attacks the animals more viciously and -21- persistently, causing a considerably greater amount of terror in herds of cattle. The reason for the wild running of cattle when attaCked by heel flies has never been conclusively established. A single adult H. ~ has been observed by Hadwen (1915) to put a whole herd of cattle into hea~ long flight. Capturing the single fly put an end to the cattle's excite- ment. It is generally the opinion of investigators (Faber, 1927; Davies and Jones, 1932; Bishopp et al, 1926, and others) that adult heel flies have very little tendency to migrate more than a mile from one area to another. MacDougall (1935), however, states that there is a record of one heel fly that flew nine miles. Descriptions of eggs, of hatching, and of skin penetration by young larvae have been adequately covered in other literature (Hewitt, 1914; Had~en, 1919; Warburton, 1922; Bishopp et al, 1926). Route of Larvae through the Bo~ Discussing H. lineatum Hadwen and .Fulton (1924) state, - 11 Wherever the . . eggs are laid the underlying tissues show the tracks followed by the larvae•. These investigators found young larvae in the connective tissues of the diaphragm, duodenum, esophagus, pericardia! sac, and heart. Bishopp et al (1926), citing Stub, report that he observed an infiltrated area, caused by grubs, in the superficial connective tis~e on the inside of the right tibia, which continued over the shoulder, around the muscles of the neck, to the tissue of the esophagus where it enters the thoracic cavity. They cite the finding of Hypoderma. larvae in connective tissues of the spleen, pouch, intestine$, thoracic cavity, loin muscles, hip muscles, baCk -22muscles, spinal column, and kidney vicinity. Hadwen ( 191 7) says, liThe larvae follow connective tissue exclusively and no larvae have been discovered in the muscular tissue 11 • He explains that connective tissues offer the least resistance to the passage of the larvae. The majority of H. lineatum migrate in connective tissues by various routes, from the point of invasion to the esophagus. A few may wander over the various organs of the body and never reach the esophagus. They- remain on the esophagus for a :period of several months, and then migrate through the connective tissues to their subde~ positions in the back (Schmid, 1941; BiSho:pp et al, 1926; Stub, 1920; Warburton, 1922; Hadwen, a.nd others). Schmid (1941) reports that .!h bovis migrates to the spinal canal along the nerves, and from there along the muscles to the back. According to Nauck et al (1948) H. ~ follows the course of the large nerves to the epidural fat of the spinal canal. Haberman et al (1949) report that Gebauer found. larval traces of H. bovis in the epineural connective tissues along nerve trJ.Dks, from the foot to the spinal canal of calves. Haberman et al (1949) summarize their investigations as follo\tS: 11 0ur observations on the occurrence of Hypoderma ~ in the spinal canal of cattle indicated a distinct difference in the migration r~~te follo~red by the larvae of the two species. Only the larvae of H. ~ occurred regularly- in the spinal ca.nal whereas those of R. lineatum were rarely found in the spinal canal. On the other hand, only H. lineatum occurred regularly- in the esophageal region. The increase Til the average length of the larvae and in the number collected betveen September a."1d June indicated that the larvae graduall7 ac~~ate in this region prior to appearing in the subdermal regions of the back. This occurrence corresponds with the well known acc-ll.tiUla.tion of H. lineatum larvae in the submucosa of the esophagus". - It is to be noted that Bishopp et a.l (1926) found only one larva of R. bovis in 114o larvae removed from the gtulets of 563 cattle in regions -23where H. bovis occurs. It is also to be noted that of 9S2 larvae removed by Haberman et al (1949) from 293 infested spinal canals, 975 were identified as R. E2.!!! and oniy seven as H. lineatum. A review of Haberman 1 s data shows that these seven H. lineatum were collected in December and February in Wisconsin. Those are months when it would be perfectly logical in Wisconsin for H. lineatum to be making its final migration to the akin of the back from other parts of the body. The average length of Haberman 1 s seven H. lineatum larvae (14.6nm.) corresponds very well with those of Bishopp et al (1926) made on H. lineatum which were read;r to leave the esophagas and migrate to the back. Larval Method of Penetration Terada and Ono (1930) report finding a toxin resembling globulin in first stage ¥zpoderma larVae. 0 bypodermatoXin" an enzyme. Ono (1932, 1932a, 1933) gives it the name ~. B:~ describes it ~s a green, viscous fluid, apparently It is found in the midgut which is dialated so as to occuw nearly three-fourths of the bod;r cavity. and has a proteolytic action. It can dissolve fibrin readily Ono states that it is certain that perfor- ation of the Skin of cattle is caused by the dermolytic action of "hypodermatoxin". Simmons (1939) agrees that the appearance of the tissues suggests gradual dissolution rather than mechanical perforation. The toxin is not present in second and third stage larvae. Larval Development in the Warbles According to Biahopp et al (1926), Ono (193S), and others, the first . . stage larva perforates the Skin of the back very soon after it arrives in -2~ tis~es. the sUbdermal Migration now ceases and a cyst of dense connective tissue, infiltrated with leucocytes, is formed about the grUb (Simmons, 1939). The grub feeds on the pus and bacteria that accumulate within the cyst (Simmons, 1939a; Ne~ov, 1933; Ono, 1933). Two molts occur while the gru.b is in the back. The length of the period spent in the warbles varies sonewbat according to di£ferent authors as is shown in table I. Bishopp et al (1926) found in Texas that the time spent in the warbles by R. lineatum. averages slightly loDger in the case of larvae coming up to the backs in the early part of the season. The average period of those which appeared in the backs in October and early November was about seven days longer than that of those which appeared during the latter part of December and in Jannar,r. Under Montana conditions during two grub seasons, the writer found that the first H. linea.tum. to arrive in the backs remained there for a. period approximately three weeks longer than was the case with the last ones that came up. -- This was also true of H. bovis. this difference has not been determined. The cause of Su.ch a phenomenon is of consider- able importance in a beef-raising state like ~iontana, where the Sll.ceess of a control program might require reducing the number of insecticide application to two. Larval Emergence and Pupation When the third stage larvae complete their growth they work their way out through the holes which have been kept open for two months, and drop to the ground. This normally occurs between 5:00 and 10:00 A.M. They seek a sheltered spot among t:rash, or if the soil is soft, they may burrow down -25- Table I. Total developmental period of grubs in the backs of cattle. lnve stigator H. lineatum H. bovis Number of Number of days Number of ""'Nuiber of days specimens in back specimens in back studied Min. Max. Av. studied Min. Max. Av. ~lagoveshchenskii and Pavalovskii(l930) . 56 Obitz (1935) Soni (1938) 51 Nauck et al (1948) 51 Gebauer (1940) 53 72 77 :Bishopp et a1 (1926) 350+ 35 89 54.5 approx. 6 sCharf'f 209 36 85 54.5 174 stu~) (present 65 77 72.8 -26into it for a distance of an inch or two. Pupation may take place within 12 hours if temperatures are favorable, or may be delayed for several days (Massonnat and Vinet, 1913; GlAser, 1914; Gebauer, 1939; Bishopp et al, 1926; and others). The length of the pupal period is dependent upon temperature. references were consulted on this matter. Twenty The shortest period, 10 days, was reported by Ono (1938) and Gebauer (194o) for R. lineat'Wll; the longest, 80 days, by Pfadt (1947), also for H. llneatiun. The average for both species is probably between 30 and 50 days, but since the rate of metamorphosis is so dependent upon temperature, the pupal period will vary with the temperature of' its micro-environment. SFASORA.L VARIATIONS For a period of from 9 to 11 months~·cattle grabs are in a comparative- ly constant environment within the body of their host. It seems reasonable to assume that in this environment their average rate of' development should be fairly constant. During the prepupal, pupal, and adult periods, however, they are at the complete mercy of the elements. As shown above, it is theoretically possible, under most favorable weather conditions, for oviposition to occur within two weeks after emergence of grdbs from the back. On the other hand, it is again possible that oviposition might be delayed to a~ period of 14 weeks or more. Thus it is possible to have a year-to-year variation of three months in the peak period of fly activity and the subsequent period of gra.b appearance in the baclcs. seasonal variations probably never occur. &.1ch vide -27Adults The season of adult activity will var,y considerably according to the climatic conditions. ]ishopp et al (1926) report heel fl;y emergence in southwestern Texas as early as December and in iev York as late as September. Where H. lineatum and H. several weeks earlier. ~ occur together, the former is usually The period of adult activity 1a generally an esti- mate made on the basis of observations of cattle showing ~toms fly attack, a method which lends itself to considerable error. of heel Mills et al (1946) report that cattle in Montana have shown tY"Pical symptoms of heel f17 terror in late Februar,y. The writer observed. the same symptoms in a herd in 1948 at least one week before there had been an;y larval emergence in the area. In that instance at least, the heel fly symptoms were caused b;y something other than heel :flies. Since cattle grubs remain in the hosts' backs for periods of from five to eleven weeks, a much easier and more tangible study of H. E..2.!!! and !:.. lineatum seasonal variations can be made through observations of the grabs in the warbles. Grubs :Bather thorough studies have been made b;y ]ishopp et al (1926), Mote (1928), and. Haberman et a1 (1949) on the seasonal histor-.r of cattle gru.bs . . in the esophagus and spiDnl canal. Year to ;year variations of grub appea~ce and grub abundance in the backs of cattle have been reported b;y only a few investigators. ing reports are from ]ishopp et al (1926). The follow- At Uvalde, Te:xas, H. lineatum ordinarily appears in the backs about September 15, but in 1920, it did -28not appear until October 9. At Dallas, Texas, first appearance in 1914 and 1918 was about October 10; in 1916 it was October tember S. JO; and in 1923, Sep- At Herkimer, N. Y., H. lineatum first appeared in the backs on March 1, 1920 and on Febru.aey 6, 1922. Walton (1925) states that the date of first larval appearance varies widely with climatic conditions. The vri ter found H. linea tum in the backs of one Montana herd on February 10, 1948; the following year there were none before March 10. It appears evident that year-to-year variations of a month or more in grub appearance dates may not be at all uncommon. Appearance dates of gruhs vaey widely in aey one season vi thin compare.tively small individual areas. Smith et al (1941) say, "In El Paso county, Colo., the first grab openings appeared in one herd, located at an altitude of about 5,000 ft. on December 4, 1938. In another herd, 50 miles distant, but located on a high mountain plateau at an altitude of about 7,000 ft., the first grub openings were seen on January 15, 1939." Here, in a distance of 50 miles, is a difference of 42 dais in seasonal development. Shull et al (1945) found in Idaho that the first gru.bs appear from late January in . . . the lover Salmon River area until mid-April in the upper Snake River area.. This is a difference of nearly 90 days· in a distance of about and a change in elevation of approximately 3500 feet. 3~ miles Pfadt (1947) states that H. lineatum first appeared in the backs at Cheyenne, Wyoming on Januaey 10, 1944. At Laramie, 1000 feet higher and 50 miles away the first appearance was on February 2, 1944. Here, there is a difference of 23 dqs in a distance ·of 50 miles. The date of gru.b appearance cannot be correlated in all instances, however, vith elevation. In Table II are listed the appearance dates of -29- Table II. First appearance dates of H. lineatum and H. bovis in Gallatin County,"T!ontana. -- Herd number and name I. (Feistner) II. (Storey) III. (Axtell) IV. (Hash) · V. (Uonforten) VI. (Carter) IIIA. {Axtell) IVA. (Hash) Number of cattle examined Elevation Date grubs were first observed in in backs feet H. lineatum H. ~ 18 ~100 32 29 16 4800 4900 27 5000 5&00 ~2 12 21 ~950 49oo 4950 Jan. Jan. Dec. jan. Dec. z:ra.r. Dec. Jan. 12, -1949 14, 1949 Feb. 8, 1949 29, 1948 21, 1949 30, 10, 26, 5, 1948 1949 1947 1948 lt.ar. 19, 1949 Feb. 3, 1948 Mar. 4, 1948 -30~ lineatum in six herds under observation during the 1948-1949 grub sea- son, near Bozeman, Montana. Referring to Table II: Herd number V, 6oo feet lover thrul number VI, and less than 25 miles distant, was Herd number I, 57 ~s 1500 earlier. days earlier in grub appearance. feet lover than number VI, and Herd number I, 25 miles away, was 71 14 900 33 miles distant, vaa feet lower the.n number V, and about days~ in grub appearance. Herds numbered II to V were located within an area five mileo long by two miles vide. ~iahopp et al (1949) have prepared a map entitled, ~Approximate date lines of first appearances of grubs in backs of cattle in the United States•. A comparison of some of these da.te linea vi th data already supplied above bythe writer indicates that sol!le of those lines may be in error by as much as 30 to 90 da.ys in parts of Utah, Wyoming, Idaho, and Montana. It would. seem that errors as large as those could hardly be included rl thin the range encompassed by the term, "approximate". The first appearance of H. bovh in four herds (Table II) during the 1948 and 1949 grub seasons vas from nine to appearance of H. lineatum in those herds. R. lineat\U'I first appeared. by Hadven (1917), 58 days later than the :t'irst The average was 33 dars a!ter This period agrees very well with that reported but is considerably at variance with the corresponding data supplied by :Bishopp et al (1926) and Mills et al (1946). The appearance and energence dates of grubs in the backs are listed in Table III on four herds in the vicinity of Bozeman, V.ontana during the and 1949 cattle grub seasons. 1948 Table III. Appearance and emergence dates of cattle grubs in the backs of cattle, Gallatin County, Montana, 19l~g and 1949 grub seasons. Herd number and name Elevation in feet IliA. (Axtell) l49oo 4950 4100 56oO IVA. (Hash) I. (Feistner) VI. (Carter) II IA. IVA. I. VI. (Axtell) (Hash) (l!,eistner) (Carter) 4900 4950 4100 56oO Grub First season appearance H. 194S Dec. 2~ 1943 Jan. 5 1949 Jan. 12 1949 Mar. 10 1943 194S 1949 1949 Feb. Mar. Feb. Mar. Last appearance linea.tum Feb. 23 Feb. 20 Mar. 13 Apr. 20 H. bovis 3 Apr. 19 4 May 12 g May 11 19 June 4 First emergence Last em ergence Mar. 3 Apr. 11 Mar. 15 Apr. 11 Mar. 13 May 5 Apr. 20 Jun. 4 S:eread in days of: AppearEmerganoe en co 59 45 65 41 39 27 4S 45 I \j.l Apr. May Apr. May 29 July 1 12 July 12 13 June 24 20 Aug. 3 76 69 92 77 63 61 72 75 1-' I -32EFFECTS OF NATUBAL FACTORS ON CATTLE GRUB POPULATIONS Climate The role of climate as it affects seasonal variations in appearance of heel flies and cattle grubs has already been discussed. Climatic conditions are also an important factor in reducing cattle grub populations. Natvig (1937) reports that flies and grubs are fewer in Norway following cold, wet summers, and that in the more northern parts of the country th~ seem to have been totally exterminated in unfavorable years. · I:a. Bri\ish Columbia, according to Radven (1915), the spring of 1913 was very wet; as a result, very few H. lineatum were found in the backs in 1914. Similar situations are reported and discussed by Radwen (1919) 1 !J.acDougall (1925), Walton (1925, 1928), Blagoveshchenskii and Pavlovskii (1930), Obitz (1936) and Bruce (193S). Extended periods of cool rainy weather lengthen the prepupa1 and pupal periods, and practically prevent oviposition. Bruce (1938) states that H. . E2.!!.! will not mature in soils with a .· moisture content of more than 22 percent, and that H. lineatum will not mature if the moisture content is higher than 10 percent. The same vet soils, he found, when amply drained, would permit co:a:plete development and adult emergence. Citing Donaghue, Bru.ce states that during the spring and vegetative season in the Red River Valley of the North in North Dakota, Minnesota and Manitoba, the moisture content of the soil is above 25 percent. Under average climatic conditions, cattle grubs are entirely absent from this 12,000 square mile area. Soni (1938) vri tes that in India cattle grubs are most prevalent in sandy areas where rainfall is scanty. Sen (1934) states that !!.:_ lineatum is practically- absent from moist Madras, -33Assam, .Burma, and Ceylon. According to .Bishopp et al (1926) no grubs are found in cattle on low sw.mpy ground. Cocpa.ratively small areas, grub- free as a result of soil moisture relations, are probably not uncommon throughout most countries where cattle grubs are a problem. The general dissemination of cattle grubs throughout the tropics and the southern hemisphere has not occurred, in spite of the fact that many grubby cattle have been shipped into various points within these areas. Oase (1929), .Bishopp (1915) and ::Sishopp et al (1926) express the belief - ~ - ~ _,._ that climatic barriers, principally hot weather and its effects on pupae and adults, are at least partly responsible for this lack of spread of the two species. Predators Robins have been observed by the writer to exhibit a marked likiDg for mature H. bovis in the active prepupal stage. ::Sishopp et al (1926) re- port that both robins and chickens will greedily devour cattle grub larvae. Walton (1924) states that jackdaws commonly visit cattle and extract the grubs. Goodrich (194o) states that in 1937, 193S, and 194o, there were Jllai1Y reports of starlings devouring grubs in the backs of cattle. The starliDgs were feeding not only on the grubs, but also on the surrounding flesh, making large and deep wounds. According to Lucet (1914) and Stegmann (1920) moles and other small rodents feed upon mature larvae and pupae. Parasites Obitz (1937) reports two H. E.2!!..! pupae parasitized by ichneumonids, -34one by nematodes, and several others by bacteria and. fungi. Lucet (1914) expresses the opinion that certain fungi destroy the pupae. Natural Resistance of Host It is common knowledge among stockgrowers that cattle grubs are mch more numerous in the backs of calves and yearlings than in older animals. Experiments by Peter and Gaehtgens (1934) Show that the treatment of rabbits with an aqueous extract of cattle grubs produces specific antibodies, the presence of which can be proved by antigens. In the same way, epecific antibodies were determined in the serum of infested cattle, a negative result being obtained with the sera of calves that could not have been infested. The antibodies were found in all young cattle that bad been exposed to infestation and bad grubs-in the backs, but in ol4 ~-free animals, the serological reactions were positive in some cases and negative in others. In the latter, it could not be determined whether antibodies had ever existed. The authors conclude, therefore, that the reaction with antigens may be used as a proof of infestation, but not of a naturally acquired immunity. Hadwen (1916, 1917) and Hadwen and Fulton (1924) present evidence to show that immunity, or naturai resistance to cattle ~b invasion increases with the age of the animals. Newly hatched larvae, penetrating the skin, cause skin lesions, anaphylactic in nature, and accompanied by swelling, dermatitis and more or less exudation of serum. These akin lesions, they report, are confined almost entirely to older animals, calves showing only slight effects. The writer has observed that varying percentages of grubs will die in different animals after their arrival in the back. In some animals, all -35grobs that perforate the skin of' the back will mature and emerge normally. In other animals, on the other hand, all grubs that reach the back will die within 10 days after the breathing pore has been made. In one instance, 30 R. lineatu.m and two R. bovis were observed to be destroyed this way in the back of' a three-year-old Guernsey cow. Table IV shows the total mortality of' gra.bs in the backs of' four herds of' cattle under observation in Montana. The natural resistance of' cattle to grubs varies widely, even one-year-old animals which are experiencing their first grubs. a.l!!Ong In some herds that are infested with large numbers of' grubs, there will be individuals present which are practicall;y grub free. will show the af'oreDentioned 11 There will be others which dela;yed resistance 11 (where the gra.bs die after reaching the back) • CONTBOL Heel Flies The short life and elusive habits of' the heel flies, together with the fact that they do not feed, make the direct control of' the flies a virtual impossibUit;y according to our present knowledge. No feasible method of' repelling the flies to prevent their oviposition has as yet been discovered or devised (Bishopp et al, et al, 1926; Mote, 1928; Mills 1926; Soni, 1946). Destra.ction of' Eggs or Newly Hatched Larvae Imes and Schneider (1921) and Mohler (1923, 1926) used shallow wading vats containing various chemicals, and by systematic and consistent use of' this method, were able to reduce grub infestations in succeeding seasons. Table IV. Natural mortality of grubs in backs of cattle, Gallatin County, Montana. :Breed of Cattle Age in Years IVA.. (Hash) Mostly Guernseys 3-7 IliA. (Axtell) Shorthorns Herd number and name VI. (Carter) Holsteins I. (Feistner) Holsteins Number Total number of Number of' grubs cattle Grub grubs that appear- that died before Percent examined Season ed in backs maturin~ Morta.litz 18 1948 151 111 73 9 1948 169 56 33 3-5 9 1949 115 31 27 3-7 17 1949 234 lto 19 Mostly yearlings l VI 0"\ I -37Eradication is not :probable, however, and the method is wanting in many respects. It would be impractical in a range cattle countr,r. Immunization A few immunization exper.iments have been made, but to date they have been negative in results (:Boubaud and Perard, 1924; Stefanski and Obitz, 1935). Preventing Oviposition Coppens (1913) . reports that there will be very limited numbers of gru.bby animals if all cattle are kept under shelter during the heat of the ~ (10:00 A.M. ~ 5:00 P.M.) in the fly season. the use of shelters as a control meagure. Haseman and Roland (1941) Selan Walton (1913) {1925), recommends Mote (1928) and recommend the use of abundant shelters and ponds for the protection of the animals during fly time. Seldom will heel flies oviposit on animals in shade or on animals standing in water. These methods wUl reduce gru.b infestations to a large degree i f pro:perl.7 used, but the)" will not result in eradication. Combating the Grubs in the l3acks The fight against cattle grubs finall7 concentrates upon the grubs during the period when they are present under the skin of the animal's back. During this :period, the infestation is visible ant!. apparently most vulnerable to the control efforts aimed at destroying it. It is during this period that the greatest success in combating the pest has been achieved. Eradication has been demonstrated in small areas of certain foreign -38countries, and significant reductions have been obtained in Europe and North America. Mote (1928), citing ~oas, reports a systematic grub ex- traction campaign undertaken by a dairy association in Denmark. On a farm in the center of the control area total grub counts over a period of seven years, beginning in 1889 were as follows: 832, 215, 65, 229, 64, 0, 0. Complete eradication was not accomplished toward the edges of the area, but the number of grubs was greatly reduced. Coppens (1913), reporting on the systematic destruction of grubs in approximately 4,000 dairy cattle, says that in 1902, 22,394 grubs were killed, and in 1910 only 3,875 were found. In 1910, the average infest- ation of the treated cattle was 0.11 grubs per head, and for imported cattle, 16.0 per head. He states further that local eradication of the pest would have been po·ssible in the absence of imported animals. Carpenter and Hewitt (1920) observe that on· Clare Island, County Mayo, Ireland, three miles-from-the mainland, successfui extermination of cattle grubs was achieved in five years, and probably could have been done in two years. In Denmark a law was enacted in 1923 reqa.iring the destruction of all cattle grubs. As a result, according to Faber (1927), ~oas (1928) and ~homsen (1935), the percentage of grub-damaged hides in Denmark dropped from 23 percent in 1922 to four percent in 1925, to three percent in 1932, and even to 0.1 percent in the Danish Islands. the Denmark situation further. He says Petersen (1945) discusses that in 1939 and 191K> eradication was suspended due to an outbreak of foot and mouth disease, then vas :put into effect again. In 1941, 303,550 animals were infested and by 1944 -39the infestation had been reduced to 99,747 animals, a reduction of 67 percent in n~er of animals infested. Davies and Jones (1932) give an account of control experiments in Wales, where derris powder was used. One to two. years after treatment, in- festations were reduced from an average of 13 grtibs per animal to about one per animal . In Bavaria ZaUbzer (1932) describes a cooperative control campaign in which the total grub infestation was reduced 37.3 percent (29,790 grUbs extracted in 1929 and 3, 808 in 1932). hook. The grUbs were extracted with a crochet Zau.bzer feels that the infestation could have been eradicated in two years, with proper cooperation of the cattle owners. In May, 1938, the government of Cyprus approved a compulso17 grub eradication program for the Island. McDonald (1945, 19%), reporting on the campaign, states that from the fall of 1943 to Decembe~ 1945, only 52 cases of warble infestation '-tere observed. During this period, only one H. bovis was found, - - the rest of the identi£iable larvae being H. aeratum which is normally found in goats. Gibson and Twinn (1936) report complete grub eradication in Canada on 50 to 80 milk cattle, and more than a 95 percent reduction in grubs on 750 beef animals in five years. In another test on 2,000 ~ttle, grub numhers were reduced 37 percent in four years. In a control experiment in Ohio, Mote {1928) reports 73 percent reduction in grUb numbers in three years. Coapton.(l931) states that an iso- lated herd of 26 Holsteins in Illinois was entirely freed of grUbs in three seasons. In the central zone of a control area in Colorado, a five year -l!ocontrol program reduced average grub infestations from animal by 1933, according to Mohler infestation was reduced to 6.4 4.9 (1933, 1934, grUbs-per head. and In 35 to 1935). 1935, 5.1 In grabs per 1934 the it increased to owing to importation of infested cattle, invasion of heel flies, and lack of cooperation among cattle owners. tained a reduction of Ma.tcysee (1945) in one year ob- 60 percent in grub numbers on, approximately 600 dairy cattle in a four-square-mile area (New York). He obtained approxi- mately SO percent kills in his ezperiments. (1945) 1947, and 1948) Miller and Simms (1946, discuss control experiments in Colorado and New l-iex:tco. On a 150 square mile area, H. lineatum numbers vera reduced 60 to SO percent in three years using a rotenone wash. On an area 200 miles square in liew Mexico, a spray treatment which killed 77 percent of the a grubs resulted in 5S percent reduction in grub numbers the following year. The Requisites of Good GrUb Control A study of the work of those who have successfully controlled cattle grubs shows that there are two fundamental reqo.irements that a successful campaign must meet. First, all treatments applied must have as their objective a maximum kill of grubs throughout the control area. It appears possible to eradi- cate gru.bs in one season if the exterminators are thorough enough in their work. Second, isolation of treated animals from infested ones is an absolute essential if reinfestation is to be prevented. If grUbby animals never come any nearer t:han one-half mile from grub-free areas, there is probably -41little danger of rein!estation. All imported grubby animals should be systematically !reed of their grubs before entering a grub-free area. In a large rangeland area where there are communal pastures, all animals that use such a pasture should be treated. During heel fly season, grub-free animals should not be taken through or into an area that held grubby animals at any time during that season. Cattle grub control is far from easy and many a poorly executed campaign bas resulted in no significant rednction in cattle grub numbers. Specific Methods for Controlling Grubs Removal of individual grubs from their cysts was one of the first effective methods used in grtib control, and is still one of the most effective i f systematically done. Squeezing, and the use of various sorts of hooks and suction methods have been recommended (Coppens, 1913; Selan, 1913; 1917; Carpenter and Hewitt, 1920; Seymour and Jones, 1921; Broderson, Bene, 1922; Gabert, 1924; MacDougall, 1924; Mote, 1928; Case, 1930; StefanSki and Obitz, 1935; Haseman and Roland, 1941). Scores of chemicals in various formulations, and methods of application have been tested on cattle grUbs. and practical. Only very few have proved to be effective Most investigators have reached the conclusion that ground derris or cube root is the most effective and practical material known at the present time for the control of grubs. The new organic insecticides, chlordane, to::mphene, DDT, benzene hemchloride and methoxychlor, in preliminary tests, bave proved to be either ineffective or quite variable in their grub killing ability (Sergent, Furman and Douglas, 1947; Telford, 1947; Westermarck, 1948}. 1946; -42The wash or scrub method of applying derris and cube root to grubs has for many years been an effective and accepted method of control (Wells et al, 1922; Bishopp et al, 1926; MacDougall, 1924, 1930; Davies and Jones, 1932; Remien, 1933; StefanSki and Obitz, 1935; Gibson and Twinn, 1936; Mohler, 1938, 1941; Smith et al, 1941; Stewart, 1944; Miller, 1945; Snipes Kills of et al, 1948). 6o to 100 percent are usually reported. The most commonly used formulae consist of 8 to 16 ounces of powdered cube or derris root (four to five percent rotehone content) in 1 gallon of water, to which is added two to four ounces of soap or other detergent. The material is applied to the backs and worked in vigorously by hand or with a stiff brush. The dusting method by which derris or cube powder mixed with a diluent dust is sprinkled on the backs and worked in vigorously by hand, has gained mu.ch favor in recent years. This is especially tru.e in areas where cold weather treatments are necessary and the application of is preS1lllled to be injurious to cattle under such conditions. li~ds Kills with the dust are generally not quite as good as with the cube or derris washes. Bishopp et al (1926) report 95 percent kills with undiluted derris powder, . while Bishopp et a.l (1930) obtained erratic results in extensive tests with derris powders. derris. Howard (1924) reports a 98 percent kill with. undiluted Laake (1942) recommends the mixing of equal parts of wettable sulfur vi th the powdered roots. Since 1942 there has been considerable difference of opinion as to which diluents are best. that tripoli earth is most effective, yielding sulfur is least effective. 6o Miller (1944) states percent kills, while Annand (1945) reports tba.t tripoli earth, vol- -43canic ash, and pyrophyllite are much more effective than talc or sulfur. Mathysee {1945) (4.2 found that ground cUbe powder part, to pyrophyllite three parts, gave 77 percent rotenone) one percent kills, and, he ~s. Telford (1947), using cube root (4.5 resulted in satisfacto17 control. percent rotenone) one part, and ca.lci11m carbonate two parts, obtained 90 percent kills. cUbe root (1.25 Snipes et al {1948) reports 66 percent mortality with percent rotenone) in tripoli earth. The injection of various liquid materials into the individ.u.a.l cysts bas given high mortalities but is indeed a laborious process (Bishopp et al, 1926; Mote, 1928; Dinulesco, 1934; Westermarck, 1947, and others). A somewhat similar method, the application of insecticidal ointments bas been tested widely and found effective, although again it is laborious (Wells et al, 1922; Howard, 1924; Bishopp et al, 1926; Compton, 1931, and others). The Use of' Power 'Sprays The first recommendation for use of a power ~ray or for cattle grub control was made by Wells spray applied at (1942). He advocated the use of a bard, driving 250 pounds per square inch, using the formula: CUbe or derris powder Wettable sulfur Water (5 percent rotenone) 5 pounds 10 pounds 100 gallons The material, he explained, penetrates the anima.l 1 s coat well, and dislodges the scabs over the grub holes. The high pressure spray method quickly' became popular and millions of cattle have been sprayed with ing degrees of control obtained. var.~- This method of control would seem to be the only practical one developed to date for the treatment of range cattle. Ma.tw range animals are ve17 wild, and it would be virtually impossible to -44handle them efficiently b7 hand treating methods. The speed o! treatment vi th a power spray machine 1 s of great importance in making the control o! grubs in range cattle practicable. sp~a Unfortunatel7, in practice the control of cattle gntbs by u~ll7 far from 100 percent effective. 11 At the present time there are m&ny conflicting recollli:lendations in the literature regarding the ingredients of spra7s and. in application techniques. In all investigations knovn to the writer the investigators, vi th two exceptions ( Stevar~, 1944; J'u.rma.n and Douglass, 194S) have used a cone-shaped. spray which makes a round pattern on the animal's back. Stewart ~s, nEarl7 trial tests demonstrated tbnt a nozzle delivering a flat, fan-shaped spray provided much better penetration through the hair than a nozzle delivering a cone-shaped spray". Douglass also used the fan-shaped pattern, and a pressure of pounds. Jlu.rman and 350 to 4oo Rupp (1948) states that 75 pounds pressure resulted in 75 to 85 percent kills. Snipes et al (1948), making five tests on onl.r 25 animals and 324 grubs, apparently obtained an almost perfect correlation between spray pressure and percentage of grubs killed. report 73 percent control. At intervals of At 200 pouda preaw.re thq 50 pounds, they increated press- ures up to 4oo pounds, with gradually increasing percentages of control. Their reported kill at 4oo pounds pressure vas 97.5 percent. Shull et ol. (1945) and Anna.nd (1945) • recommend. a minl1n1m pressure o! 4oO pounds. Stewart (1944) used 250 to 300 pounds presware in his experimental work. !ishopp et al (1949) recommend spra7 pressures of 4oo to During the first year or tvo that power spraya vere 500 pounds. use~ for grub control most workers agreed that 5 pounds of powder bearing 5 percent Table V. Power Author we-if&-· \191+2J Snipes (1942) Stewart (1944) spr~ ingredients and pressures tested or recommended by different workers for cattle grub control. Ingredients per 100 gallons of water Ground derris or cube root Pressure used Percentage ( 5% rotenone)____ ~ _ W~t_ting _age_nj; ~ _____l]l_._ I'_~r _s_g_.___!n~--~k_ill re-ported 11Effeetive 11 5 lb. Wettable sulfur 10 lb. 250 5 lb. 10 lb. Wettable sulfur 10 lb. 350 11 90 Wettable sulfur Detergent 10 lb. 300 72 l pt. lb. None 4oo "Good Xi11 11 lb. None 4oo 11 Effective 11 Shull et al (1945) 7?3 11! Snipes et al (1948) 5 lb. Wettable sulfur 10 lb. 4oo 11 Furman and Douglas (1948) 10 lb. \fettable sulfur Diglycol laurate 10 lb. 4oo 56 Furman and Douglas 10 Anonymous {1944) or more 11 1 :pt. 90 or more" lb. None 4oo 59 (1948) Furman and Douglas (1948) 7Ir 1b. None 4oo 53 Eishopp et al (1949) 7i lb. None 500 "Effective" Knowlton (1949) 713 lb. None 500 II Knowlton (191±9) 6 lb. 500 11 Wettable sulfur 10 lb. I f -46rotenone plus 10 pounds of wettable sulfur per 100 gallons of water were satisfactocy. Stewart (1944) concluded that the rotenone content of the spray should be doubled and that a pint of detergent be added (Table V). U.S.D.A. workers (Ano~vmous, 1944) reported that 7! pounds of 5 percent rotenone per 100 gallons of water gave better kills at 4oo pounds pressure without the addition of sulfur or wetting agent than with one or the other of these materials added. Shull et a1 (1945), ::Bishopp et al (1949), and most other workers are currently recommending this forma.lation. Snipes et al (1948) advocate Wells 1 (1942) formula, stating that it is the most effective of several tried. Tests of Cattle Grub Sp~s in Montana. The purpose of these experiments vas to attempt an evaluation of some of ~he currently recommended spray formulae and application methods, and to improve upon them if possible. Materials and Methods. The animals treated in these tests were range beef stock infested with average numbers of grubs of both species. pressures, from 4oo to 600 pounds were used. Variable !fhe spr~ nozzle was held vertical to and 12 to lS inches from the animals' backs, and produced a pattern about 2! inches in diameter at this distance. During the spra;y operation the animals were restrained in chu.tes or small crowding pens. In one test a special cattle grub teeth vas used. gtlll with two rows of rak&-like :Between the two rows a fan-shaped spray was emitted. In operation, the animal' s back was raked, the spray being forcibly ejected against the back at a nozzle distance of about two inches. -47Nine to 15 ~s after treatment, all grubs were squeezed from the cysts, and mortality was determined. It was found that an interval of 10 days between treatment and examination made mortality determinations relatively easy and sure. Partly healed empty cysts were ignored in the mortality calculations. All living first instar grubs found at the time of examination were presumed to have arrived in the backs after treatment (Bishopp et al, 1926, found that the first stage grubs almost invariablymolt within 10 days after perforating the skin of the back.). Discussion of Results. summarized in Table VI. The data obtained from these experiments are Erratic results were obtained in maey of the experiments, and an eXamination of the data. immediately- suggests that there is need for good experimental work on the use of spra;ys for cattle grub control. Some of the sa.bsequent statements must be considered as preliminary-, pending the collection of more data. Stateoents by Bishopp et al (1926) and Furman and Douglass (1948), suggest that H. bovis is considerably- more resistant to most rotenone formulations than is H. lineatum. Rotenone e:ttracts and y..u-ified rotenone when used at the recommended concentration are not as effective for killing cattle gro.bs as is ground cube root, according to Smith et al (1941), and Furman and Douglass, 1948. These views are definitely- corroborated by the writer in the current experiments. Three wettable powders, impregnated with rotenone extracts also proved to be inferior to the ground cube root. One test made at 600 pounds pressure gave the best control of H. bovis, and suggests the need for more experimental work with higher pressures. All tests on H. bovis indicate that there is much to be desired in r . !l'abla VI. Results o£ Insecticide !rests against Cattle Grubs • For.mu.lation per 100 Gallons o£ Water · Date Sprayed Mar. 3, 1 49 Mar. 3, '49 Mar. 3, 1 Application Spray Species Animals Grubs Percent Technique of (Cube powder contained Pressure Treated Treated Killed PSI• Grubs*"' 5~ rotenone) L 16 208 7~ lb. Sprayed twice Cube powder 450 95 with garden hoe mas sage between sprays - 450 L 15 171 90 7~ lb. Sprayed once Wettable sulfur 10 lb. 450 L 16 264 gg Ou.be powder 7t lb. Sprayed twice 49 Cube powder Mar. 3, 1 49 Cube powder 7t lb. Sprayed once 45o L 15 192 s4 Ma.r. 3, 1 49 Spray-ed once Liquid rotenone extract plus s,ynergists (2.5~ rotenone) S pts. 450 L 13 114 74 Mar. 3, '49 Sprayed once Wettable powder 1mpregnated with rotenono extract plus synergists (2.5~ rotenone) 4 lb. 450 L 14 173 56 Mar. J 1 1 49 Check animals •PSI - Pounds per square inch . L •• L= Hypoderma linea.tum; B= Hypoderma. 1 E2.!!!• ~3 --- - 7 r 3f I ~able Date Sprayed Mar. 1-!a.r. 24, 24, Results of Insecticide Teats. against Cattle Grubs Formulation per Application Gallons of Water Technique (Oube powder contained ~ rotenone) Sprayed once t4g Liquid rotenone ex.tract (2.5% rotenone) 5 pta. 100 Sprayed once t).:.g Liquid rotenone ex- tract none) Mar. VI (Continued). 24, 1 43 (2.5% rote- 10 Spray Species Animals Grubs Percent Pressure of Treated Treated Killed PSI* Grubs** 4oo l.!oo Sprayed once l.!oo 24, May 3, '49 '}t.g Wettable powder im.Sprayed once pregnated with rotenone extract (% rotenone unknown) 7-h lb. Sprayed twice Liquid rotenone extract plus synergists (2.5~ rotenone) 8 pts. B 5 63 32 5 65 23 6 45 44 and L pregnated with rotenone extract plus synergists (2.5% rotenone) 15 lb. Mar. B and L pts. \~ettable powder im- B and L I £' f 4oo B and L 450 B 6 63 lto 19 llS 41 63 19 B Sprayed twice 8 Liquid rotenone ex450 tract plus synergists (~ rotenone unknown) 10 pts. "'"' L= !!YJloderJr!. 11neatum; B= !Jypoderm .bovis •. •PSI - Pounds per square inch V.a.y 3. ·~ Table VI (Continued). Formulation per Gallons of Water (Cube powder contained . ~ rotenone} 100 Date Sprayed 26 366 89 Sprayed once 500 l3 71 455 68 Sprayed once 4oo l3 52 325 78 lb. Sprayed twice Cube powder Sodium lauryl sul.fate 1 lb. 450 l3 17 96 73 Cube powder 7-ff lb. Sprayed twice anhydrous alkylated aryl poly-ether 1 pt. alcohol 450 l3 23 220 68 Oube powder Apr. 28, •48 Cube powder May 1 May 3. May 3. May 3. Mey 5. 49 1~ Spray Species Animals Grubs Percent Pressure of Treated Treated Killed PSI* Grubs•• l3 29, •4s 3. Application Technique 6oo Apr. 21, tl~g Cube powder Apr. Results of Insecticide Tests against Cattle Grubs. 7! lb. Sprayed once 7! lb. 7i lb. 1ts I 49 Cube powder Wettable sulfur 7i 10 lb. Sprayed twice lb. 450 l3 20 171 65 '49 Oub e powder Wettable sulfur 7! 10 lb. Sprayed twice lb • vi th garden hoe massage between sprays 450 B 2l 238 70 32 101 Sl 1 1 49 Cube powder *PSI - Pounds per square inch 7i B lb. Sprayed twice 450 using special rake-like "grUb scrubber 11 •• L::: Hrnodel'lll!'. lineatum; B= Hrroderma. ~. ~I -49the current control recommendations. ~~s There ver,y little difference in effectiveness between sp~s containing cube alone and those containing in addition to cube, wettable sulfur, sodium la.uryl sulfate, or aneydrous allcy'lated ar,yl poly-ether alcohol. The spray containing sodium lauryl sulfate gave slightlY' better control than the other formulations. Spraying the .animals twice, and massaging their backs vigorousl1 between the two spra.yings with a blunt garden hoe increased kills slightly. Better control of H. bovis was obtained with the special rake-like gun than with any method other than application at 6oo pounds pressure. It is not known whether the better kill should be attributed to the i'akiD& or the more forcible application of the Bpl'S¥ at close range, or to both of these factors. The teeth on this piece of equipment scratched the backs quite seriously-. Number of Treatcents Necessar,y Quite a variable number of treatments are recommended in the literature where both species of grubs are involved. of the u.s.D.A. is to treat ever,y Carpenter and Hewitt gro.b season. (1920) Case, to 20 days (Bishopp et al, 1949). advocated six to eight treatments during the Most authors have preferred to recommend four to five treat- ments at intervals of about 1926; 15 The current recor.amendation 1929; Compton, Dinulesco and Foisoreano, by Stefanski and Obitz 30 ~s throllgh.out the grub season (Anonymous, 1931; 1939). (1935), Anony-mous, 1932; Gibson and Twinn, 1936; Two treatments are considered cufficient Snipes (1942) and Mathysee (1945). La.a.ke -50(1942) suggests that three treatments may be enough to control most of the grubs. Where onlY R. lineatum is present, Mohler (1941) reports that two treatments, properly timed did not give complete kills, but satisfactorr control. re~ted in Smith et al (1941) found that one treatment, applied 30 days after the first grub holes appeared, killed 6o to 100 percent of the entire H. lineatum infestation. Presuming that we do have a treatment for kUling cattle grubs that is effective and practical, the gnccess of a control campaign in a large range-land country like Montana depends directly upon the number of treatments required. A large number of cattle men will not accept a four or five treatment program. They state that by handling the cattle that many times they lose more pounds of beef than would be gained by controlling the grubs in the above way. The writer therefore set out to find what percent of a mixed (H. ~and H. lineatum) infestation could be contacted by two properly timed spray-s. During two seasons individual records were kept on all of the grubs that arrived in the backs of four herds of cattle. In each of the four herds one grub control treatment, properly timed, would have contacted ver,y nearly the entire H. lineatum infestation. In other words, the first normal emergence of H. lineatum from the baCks occurred on or after the date the last individuals of that species perforated the skin of the backs (Table III). This first treatment in all cases would. have contacted a nUmber of the first H. :!!.2!!!. to arrive in the backs. A chart of these four infestations shows that a second treatment applied 60 days after the first would have contacted the following percentages of the -51remaining H. bovis in the four herds: 70, 90, 97, and 100. These high figures were made possible by the fact that in the four herds studied the earliest grohs (of each species) to arrive in the backs remained there an average of three weeks longer than the latest ones to appear (see page 24 and Table III). The four herds examined were small, and the relative number of grubs was likewise small. It remains to be determined whether a similar situation exists generally throughout larger herds and larger areas. Proper Timing of Treatments In order to kill a maximum number of cattle grubs in a two treatment schedule, it is necessary to delay the first treatment until Just before the first grubs are ready to emerge and drop to the ground. cattle owners must determine this date by careful and of their cattle during the early grub season. 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