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A STUDY OF SOME WOOD-BORING COLEOPTERA ATTACKING FIRE-KILLED CONIFEROUS TREES IN MICHIGAN by FRANK THOMAS EéRMELEE A THESIS Submitted to the Graduate School of Michigan State College of Agriculture and Applied Science in partial fulfilment of the requirements for the degree of MASTER OF SCIENCE Department of Entomology 1940 THESIS ACKNOWLEDGMENT The writer wishes to express appreciation to Professor Ray Hutson and Associate Professor E. I. McDaniel of the Department of Entomology for their guidance, assistance and sincere in- terest during the course of this study. Thanks are also due other members of the Entomology Department for their helpful suggestions and information. CONTENTS Page Introduction . . . . . . . . . . . . . . 1 Objectives . . . . . . .-. . . . . . . 6 Methods and Equipment . . . . . . . . . 7 Procedure . . . . . . . . . . . . . . . 7 1937 Material . . . . . . . . . . . . . 9 Discussion and Summary of 1937 Results . 13 1938 Material . . . . . . . . . . . . . 14 Discussion and Summary of 1938 Results . 23 1939 Material (Presque Isle Fire) . . . 26 Discussion and Summary of 1939 Results . 31 General Discussion of Species . . . . . 36 General Conclusions and Summary . . . . 53 Literature Cited . . . . . . . . . . . 56 Plates 0 O O C O O O O 0 O O O O O O O 68 INTRODUCTION Kellogg (55) reported the original forest area of the United States as 850,000,000 acres containing 5,200,000,000,000 board feet of sawetimber. Due to clearings, fires, insect depredations, storms, disease and other causes, this vast acre- age had been reduced to 550,000,000 acres containing 2,500,000, 000,000 board feet of saw-timber by 1909. Recent surveys by Garver (27) show the present acreage to be about the same as that of 1909 containing, however, only about l,800,000,000,000 board feet of saw-timber. The above statistics show that our forest resources are not inexhaustible notwithstanding our pres- ent day reforestation and conservation activities. While our total forest area has not decreased alarmingly since the be- ginning of the century, a large part of our choice timber has been removed. INSECT LOSSES T0 FORSSTS AND FORLST PRODUCTS Hopkins (44) quotes Professor Lawrence Bruner, State En- tomologist of Nebraska as saying: "I can agree with Doctor Hopkins that the insects are far more important in destroying our forests than fires." The countless hordes of tree-killing and wood-destroying insects exact an enormous toll, and were ?2- it not for rigorous control measures practiced in various parts of the country, timber losses would be far in excess of present figures. Bark beetles, especially those of the genus Dengnggtggns, and defoliating insects such as the Spruce budworm and larch sawfly are responsible for the major part of our forest loss- es. Hopkins (44) reports that the Black Hills beetle, ‘Qggdrggtgggs pggdeggsae Hopk., destroyed about one billion board feet of coniferous timber over a ten year period, which at $2.50 per thousand board feet, would mean an annual loss of $250,000. This is an example of the destructiveness of but one Species of bark beetle in one national forest. Hopkins (44) bases the insect losses on 10% of the mature trees impossible to utilize. At $2.50 per thousand board feet, he believes $62,500,000 to be a conservative estimate of the annual loss. Hyslop (53) reports forest insect losses to be variously es— timated at around $100,000,000.annua11y. Graham (:54, p. 11) assuming a 10% loss of the annual cut for the years 1925-1929 finds the yearly loss to be 5,913,383,900 board feet. Forest insects such as certain bark beetles and defoliat- ors are classed as primary forest enemies in that they attack living trees (34, p. 202). To this group of insects is attrib- uted the bulk of the forest damage. However, secondary forest insects, that is, those that attack only dead or dying timber, occasionally cause heavy damage. The larvae or "grubs" of these secondary Species, belonging chiefly to the families Cerambycidae, Scolytidae and Buprestidae of the order Coleoptera, excavate cavities in the sapwood and bore tunnels -3- into the heartwood of dead or dying timber either complete- ly destroying or greatly reducing in value a product which otherwise would be highly merchantable. Hopkins (43) states that the aggregate losses from this damage in the coniferous forests of the United States contribute largely to the annual waste of millions of dollars in forest products, which without the injury might easily be utilized. Timber killed by primary insects, fire, storms, disease and other causes are readily attacked and greatly damaged by the depredations of secondary borers. Dead coniferous timber, especially if it is standing, has been found to be in an excell- ent state of preservation for long periods after death, provid- ed no insect damage has occurred. Hopkins (44) found pines, spruces, larch and Douglas fir killed by insects and fire, and which had escaped borer damage, to be entirely suitable for lumber, fuel, pulp and railroad ties from 20 to 30 years after death. In localities where fires, storms or other agencies have killed large areas of timber, sepecially in the Spring or early summer, prompt utilization, barking or submerging in water is often impossible. Wood-boring insects together with fungi which are allowed to reach the heartwood through the borers' tunnels rapidly cause the deterioration of such timber. Losses may be as high as 50% in three months and commonly 100% in three or four years. (18), (42), (4s), (45). Webb, (90) (91), reported damage in excess of $6,000,000 to storm-felled pine in the South by one Species of sawyer, Eggoghamus 5111111522.(F3b-)- Secondary insect damage to felled trees —4- following the New England hurricane in 1938 was also con- siderable (25). ATTRACTION 0F FRESHLY KILLED TREES T0 CERTAIN COLEOPTERA Host trees, especially coniferous, are attractive to wood borers if cut or killed shortly before or during the time in which the adults are flying; such timber is almost certain to become infested. Timber dying late in the summer or fall is not nearly so heavily damaged, especially in the northern for- ests. (9), (45), (87). Graham (32) states that green wood is necessary for the early larval stages of Cerambycidae and Buprestidae. From this it would appear that the period between the death of the tree and dessication of the inner bark and cambium is the most suitable time for successful infestation. Fire-killed or scorched coniferous trees apparently have an attraction for certain insects. The smoke or the odor of the burning bark and needles probably serve as attractants but definite proof as to this is lacking. (49), (74). Deane, Van Dyke, Chamberlin and Burke, (20, p. 18), report that one Buprestid borer, Eglgngphilg gonaputa Lec., cannot even wait for the trees to cool off but flies through the smoke and de- posits its eggs on the still smouldering trunks. This species is said to annoy fire fighters by pinching them on the neck and hands. St. George and Beal (78) found beetles attracted to burned areas within a radius of three or four miles and in Sweden, Tragardh, (88), states that sawyer beetles are attract- ed for miles by fire, ovipositing first on the largest and most severely scorched trees. Trees scorched all around are most ‘35‘ attractive, he says, and only the scorched side of the tree ~ will be attacked. ROBBED-OVER AREAS AND SOURCES OF EPIDEKIC INFESTATIONS Various results have been reported on the possibility of epidemic infestations arising from burns, slash depositions and other favorable breeding places. Miller and Patterson, (62), report fire-killed and fire-injured timber as highly attractive to the western pine beetle, Dendzoctgnus brevicgmis, resulting in heavy concentrations within the burn. Such concentrations, they say, do not develop into epidemics but, on the contrary, return to normal within three or four years. Fires attract beetles but conditions are not favorable for breeding in the burned-over areas. They point out that the sudden interruption and disturbance of the sap flow in burned trees causes an ex- treme moist condition of the inner bark which is unfavorable I for the young bark beetle larvae, causing the death of many of them; in some of these trees, fewer beetles emerge than attack them. Many trees, however, are only moderately injured by the fire and apparently are able to recover. Such trees are readi— ly and successfully attacked by bark beetles; in this way only, are they believed to supplement and increase losses initiated by fire. According to Bureau of Entomology reports for 1919 (50) the infestation of beetles within a certain small burned- over area increased up to 1000%, but the numbers in the surround- ing areas decreased and the whole area returned to a normal condition within a comparatively Short period of time. These renovts agree With those of Miller and Patterson in that fire- ~6- injured trees suffer heavily from insect attack. Investiga- tions in this country indicate that neither infested slash nor burned areas become foci for epidemic infestations, (17), (so). (46), (64). In Russia, several investigators have reported fires as becoming a source of infestation affecting healthy trees. Sokanovskii (76) reports migration of bark beetles to healthy Spruce after removal of dead and severely damaged-host trees from a burned-over coniferous area; Stark and Ctaph (79) say development of bark beetles and other insects in large numbers in a coniferous forest fire area is always followed by intens- ive flight to surrounding healthy trees and Porozorov (68) re- ports beetles as multiplying in certain parts of burnt areas of Russian timber followed by migration to healthier trees. was This investigation has been concerned with a study of some of the secondary wood-boring beetles contributing to the deter— ioration of fire-killed coniferous timber in Michigan. Most of the effort has been toward the end of determining what borers are present in such areas and the part each plays in the eventu- al destruction of the wood. From time to time examinations were made of both the fire-injured and uninjured trees within and bor- dering the burned-over areas to determine the effect of such concentrated infestations on the surrounding forest. Observa- tions of insect infestations in slash and storm-felled timber, for purposes of comparison, were thought worth while. While it 97- is obvious that a study of this nature over a period of two or three years is much too short for completeness, it is hoped that this paper will bring out the importance of a group of insects which are commonly associated together in fire-killed timber. METHODS AND EQUIPMENT Infested material was cut from the burned-over areas, brought to the laboratories at East Lansing and placed in screen cages of a mesh size adequate to prevent escape of the smaller insects. For each cage of material left out of doors under nat- ural conditions, a corresponding cage of material from the same tree was placed indoors. Each was given a number so as to in- dicate that the wood came from the same tree. Individual records were kept for all trees brought to the laboratory. Each insect emerging from the wood in the various cages was given its proper number and date of emergence. In addition the smaller branches of at least one tree of each Species was kept separate from the larger limbs and butt portions of the trunk in order to determ- ine the prevalence, if any, of certain insects in different parts of the tree. PROCEDURE' The collection of material for this study was begun in the fall of 1937 and continued through 1938 and 1939, during which time several truck and trailer loads of infested fire-killed ee- coniferous timber was cut in Northern Michigan and brought to ' the laboratories at East Lansing for study. Dead wood from the following trees were cut into suitable lengths, placed in cages and labeled in a manner already described: white pine (Pinus stngbug L.), Norway or red pine (Eipps resingsa Ait.), jack pine or scrub pine (£1np§,bgpksiana uamb.), balsam fir (Apie§_ halsgmga_(L.) Mill.), American larch or tamarack (Lgri;,lazic1pa (Du Roi) Koch.) and black or bog Spruce (Eig§§_mariana (Mill.) B. S. P.). The wood caged indoors was kept in a greenhouse at a tem— perature of between 60 -70 F. The moisture content of the ma- terial was kept as favorable as possible for the immature insects by weekly applications of water. The presence of red ants in the greenhouse caused difficulty, eSpecially during the first year. These voracious insects penetrated deep into the heart- wood and devoured many of the helpless larvae. Further trouble from this source was prevented by a thorough clean—up of hiding places and by placing the cages on long, low, rack—like tables supported by legs immersed in Shallow pans of oil. Tanglefoot was smeared on the supporting rack between cages so as to pre— vent migration of the ants from cage to cage. Very few ants were observed in 1938 and none during 1939-1940. Precautions were also observed in protecting the out-of—door cages against ants but apparently they are not so much of a problem there as indoors. MATERIAL COLLECTED IN égél No fire—killed coniferous wood from Spring fires was available during this year. Material from spring fires is desirable because of its attractiveness to the female beetles for purposes of oviposition during the months in which they are flying, (May, June and July). However, material for study during this year was obtained in the following manner: Sections from the base and larger branches of white pine trees in the vi- cinity of East Lansing, Michigan were scorched by burning a light application of old crank-case oil smeared over the surface. These trees were in a weakened condition due to a heavy infestation of the bark beetles, Lpg galligzgphus (Germ) and Lp§_p1p; (Say) and were felled by the Forestry Department of Michigan State College in the early Spring only because their death appeared imminent. On May 13, 1937, after the superficial burning described above, the logs and limbs were placed in a sunny Spot near a quantity of pine fire-wood which had been infested with wood-boring in- sects during the previous year. A similar amount of recently- felled white pine wood which had not been given the burning treatment was distributed at the same time in the same locality. All wood was collected and caged on September 20, 1937. Portions of both burned and unburned wood was placed in the greenhouse for winter observations with the remainder being left out-of- doorS. At the time of caging, examination proved all pieces of wood to be infested except two fire-treated limbs of about four inches in diameter. e10- By September 20, most of the borer larvae had finished their phloem region feeding as was evidenced by oval-shaped entry holds into the sapwood. In Spite of this protection, red ants destroyed a number of the larvae in the wood of the greenhouse cages. 1937 RESULTS The following two pages are devoted to a record of sec- ondary beetles reared from white pine (Pinus strobus L.), ex- posed to infestation as described under "Material Collected in 1937." - 11 - E. soon 05% 80mm mvmowmm on Hummodmemmuma msos assume .rmm853 BoomamB mmHm. . . I a I .o o d m m . 7m .. my HNMN,no.mhbkhmwlllIIIALrnwrumwLKMBK.mm .Hol .mp , aonmp am up . >mmBmB Bommdsa mmHar o m u P\Hg\.mm «o melN_mm b, wmo m cos a b {we so mxoham Em mm so MERE. .8 Eb} HOdem mp mm >20 mmEOHmm 0m wwwabmm mmbwmb mmo: mmeccw Harm Aemczmm >20 bbwmmm mwbzommm Omommz be wbzuot memIWthEb HHfiwwm. cams swore smms.v mam .. r . 1m in m8 n ma 2 D.o pa m m k8 6r. y m S m +ot. mww ”we k.m 14. ”m r a c We. 0 .b a . . ma “any -Is “Mme WU. (\ (\ 2:33am mousse Home on Umuw msuwmoo ooHHmoaod. p00 mo 00 mm H00 momonm massed. macemmSoo venom. Amonomnnamm.onmmnwwnmmm Amway am am HH we no coo. n. some so Kenna HH. Hmeo. appeasempb m ommppwmm_fieee.v a m H as eon. me. Home ao mow. mm. meo. bmekwmnmpm thhmhm.fimmwv Q find. 0. H090 «0 mod. m0. Hmao. :30- Table VIII shows the extent to which the various woods were infested with the larvae of Mgnochamus scutellagus (Say). A six-foot section was removed from the lower trunk region of the largest trees obtainable in each tree Species. After the average diameter was determined, the larvae were chopped from the wood and counted on October 4, 1939. Table VIII +3 (H 53' 0 mas 0 on”: -P . . <3 0 CUO'U 00 $2100 8440 oIs $94 .axad at, 02> Nae-I IL. 50:40 805 '33 233 ass a I: = Tree Species 4: c: It-I «30‘ white Pine 10 9:5 9.5 Norway (red) Pine 10 44 4.4 Jack Pine 20 176 8.3 Balsam Fir 10 87 8.7 Black Spruce 10 72 7.2 Table 1x shows the number of adults of Mgnochamus pcgpgllgpgg (Say) reared from two entire trees of approx- imately the same Size of four different Species in the Pres- que Isle fire area. These trees were chosen from within an area of about ten acres and averaged about 20 feet in height and about 5 inches in diameter at the base. It was unnecess— -31- ary to cage the smaller branches as they were generally not infested with sawyer larvae. Table IX I o I o I . $40 $40) I 0 0H0 t.m:. acct. 3H m 0660) can.) . 00+: . m a), not. 3.x. .--I in. oz was $443 a e s-on-I I» Mmo«- mm0o Sud mun memo mpao FNd mum .gms .gms 30g sgd Tree Species :33?» £3?) 53mg 3:33 White Pine 28 41 69 34.5 Norway Pine 11 16 27 13.5 Jack Pine 31 23 54 27.0 Balsam Fir 17 ‘ 25 40 20.0 DISCUSSION AND SUMMARY OF 1939 RESULTS AS a result of observations for a period of one year it appears that practically all of the damage to timber in the Presque Isle fire region was done by the sawyer Mgngchagus pcgpgllgtgg (Say). No other species of Mgngchgmus has been reared as yet from this locality. The Cerambycids, Agagtngcgggs pusillus (Kby.) and Astylopsis guttata (Say), so commonly associated with.Mgnoch§mu§ in pines (15, pp. 116-119), are almost exclusively bark feeders, rarely entering even the sapwood, and thus cannot be considered of any importance in timber deterioration. _-32- ~ PARASITES AND PREDATORS. The importance of parasites and predators in the Presque Isle burned—over area seemed to be slight. Woodpeckers were quite numerous during the early fall months. They undoubtedly destroyed many larvae especial- ly in the smaller trees where their prey must necessarily be closer to the surface. Woodpecker work was found to be quite common throughout the burned area and might possibly be consid- ered of some importance in the control of wood-boring insects in such situations. (1), (20, p. 25), (50), (55). Hopkins (44), Buttrick (9) and Graham (54, p. 358) consider woodpeck- ers to be a factor in keeping small infestations of forest in- sects from becoming epidemic because of their great powers of mobility enabling them to concentrate on such Spots rapidly, but they do not believe birds can be much of a factor in large areas of dead trees such as the Presque Isle fire area. Strick- land (82) believes that the reproductive rate and territorial limitations of birds prevents their playing an important part in insect control. A number of Clerid larvae were observed under the bark of bark beetle-infested wood, but the Clerids are usually consider— ed to be predators of bark feeding forms such as Qppdppppgppg Spp. or Acapthgcinus Spp. (15, p. 119). Very few parasitic insects were reared from caged wood or observed in the fire locality. Several hymenopterous parasites of the family Braconidae emerged from Norway pine cages. Some of the pupal cases of these insects were found to be in the lar- val tunnels of MQnochamus §cutellatus (Say) indicating that -33- they might be parasitic on the latter Species. (19). Two undetermined Diptera, family Tachinidae, were reared from a cage containing white pine. Considering the habits of this family and the fact that the puparia were found in a heart- wood larval tunnel of Monochamug scutellatus (Say) (Plate XXVIII) it might not be unreasonable to assume the latter insect to be their host, although Tachinid flies do not appear to be adapt- ed for ovipositing in, on, or near protected larvae such as Mogochamus. During late August, 1939 several parasites of the family Ichneumonidae, genus Rhyssa, were captured. These large insects were flying around borer-infested trees, although none were ob- served in the process of egg—laying. Species of Rhyssa are gen- erally considered to be parasitic only on Siricid wood wasps (35). Merrill (61) claims Rhyssa to be primary parasites of Mgngchamus but Rohwer (70, p. 418) believes records of these insects parasitizing Mgpochamus are open to question. While no Siricid larvae or adults were found in the fire area, proof of Rhyssa sp. parasitizing Mgnochamus is lacking as far as this investigation is concerned. MONOCHAMUS SQDTELLATUS (SAY) AND HOST SELECTION Tables VIII and IX Show sawyer larval infestation and numb- er of adults reared from dead wood of various tree species. The counts made indicate a preference by Monochgmug scgfiellatus (Say) for white pine over all other tree Species examined. A definite preference is apparently exhibited by —54- the white-Spotted sawyer for white pine, jack pine, Spruce and fir over Norway (red) pine. If a stand of white and Nor- way pine were to be salvaged after a Spring fire, it would seem advisable to remove the white pine first. Hepkins (43), Graham (31) and others have reported white pine as especially liable to damage by sawyers and other borers. RELATION BETWEEN SECONDARY BORERS AND FUNGI INFECTION OF FIREHKILLED TIMBER. Stickel and Marco (81), Real (2), and others have emphasized the importance of fungi in the deteriora- tion of fire-killed timber. This activity by fungi begins al— most as soon as the tree dies and continues until the wood is reduced to a crumbling mass of humus. Rumbold (71) and Hubert (52) among others have investigat- ed the "blue stain" or "blue rot" caused by ngatostomella Sp. in timber. Rumbold (71) reports two Species of Ceratostom§11a_ to be definitely associated with Qendroctonus and ng bark beetles while Snyder (75), Buttrick (9), Leach, Orr and Christ— enson (57) and others have associated the ambrosia beetles with stained timber. A number of wood Specimens brought from the Presque Isle area showed fungi staining in the sapwood but no attempt was 'made to identify the causal organisms. Boyce (7) says that wood destroying fungi cannot penetrate the bark or living sapwood but must have direct access to the heartwood in order to completely infect a tree. The larval tunnels of Mgngchampg Sp, pave an ideal way for wood-rotting fungi to the heartwood of coniferous timber. Leach, Orr and w35- Christenson (58) found a fair degree of correlation between the number of Mopochamus beetles present in the logs and the amount of heart-wood decay. They found the fungous Peniophora giggnpga to be primarily reSponsible for almost complete decay of Norway pine logs three years after cutting. They believe Buprestid beetles to have little influence in the Spread of wood-rotting fungi because of the closed larval tunnels of these insects. An examination of wood samples from the Presque 1519 fire revealed heartwood decay only in those cases where the tunnels of Mgngchamug scutellatug (Say) had reached this region of the wood. (Plate XIV). While wood-rotting fungi might eventually reach the heartwood regions due to deterioration of the sapwood, sawyer tunnels facilitate and speed up the process. (34, p. 261). Fungi-infested wood was placed in prOper growing conditions in order to obtain reproductive Spores of the fungous. Profess- or Forrest Strong, Forest Pathologist at Michigan State College, examined the cultures and identified the fungous as Peniophora Sp. EFFECT OF THE SEASON OF PRESQUE ISLE FIRE. Buttrick (9) states that the optimum conditions for attack by wood-boring in- sects seems to be following fires which occur early in the grow- ing season. He says that atsuch times, the killing power of the fire is at its maximum. The moist condition of the wood causes fermentation accompanied by a rise of temperature which is favor- able to the rapid development of the larvae, giving them time to become thoroughly established before subsequent dry, hot weather causes seasoning of the wood. Buttrick (9) gives a -35- comparison of the infestation following two fires occurring at different times in South Dakota. "On one large area burn- ed about the middle of June, the infestation by the last of August was so severe that the ground under the trees was white with the dust from the borings, which could be seen drifting to the ground like a light snow. The gnawing of the larvae sounded like the croaking of innumerable frogs. An area close by burned in March was much less severely infested.” The conditions in the Presque Isle fire region was not much unlike those described by Buttrick (9). Piles of boring dust surrounded almost every tree and the ambitious ra5ping of the larvae could be heard as late as October 4th. The pres- ence of sawyer adults soon after occurrence of this fire gave an ominous warning as to what destruction would follow. DISCUSSIQN OF SPECIES OF HOOD—BORING COLEOETEBA REARED FROM DEAD CONIFEROUS TIMBER IN MICHIGAN Doubtful Specimens whose identification could not be accurately'made by comparisons with Species in the Michigan State College Entomological Collection were kindly determined by Doctor Josef Knull, Coleopterist at Ohio State University, Columbus, Ohio. Cerambycidae Mgnochamus scutgllatus (Say) (Mongchamus scutgllatus (Say)) The white-Spotted sawyer, Monochamus pcutellatus (58?) -37- (Plate 1), was found to be the commonest insect breeding in dead coniferous timber in all areas observed in Michigan. In these localities it has been found to be responsible for at least 95% of the damage. Graham (34, p. 260) says that it is so common in the Northeast that practically every log or fresh— ly killed tree left in the woods over summer is almost sure to be infested. DISTRIBUTION: Hopping (47) gives the habitat for this Species as New England, Canada (except British Columbia) and Alaska. Blackman (16) gives the distribution as throughout Canada and the northern part of the United States from coast to coast as far north as the Hudson Bay and Yukon regions and as far south as New Mexico and West Virginia. It is probable that he was including in this range the far western, closely related species M. gzegonensig Lec. Doane, Van Dyke, Chamber— lin and Burke (20, p. 187) refer to M, ogegonensis Lee. as a subSpecieS of M, acutellgtps (Say) HOSTS: Hopping (47) gives the hosts as Pippg §§£OQU§, E, nesinpga, E. banksiana, Pippa cgngdepgig, and Api§§_balsgmea. In 1938, American larch, ngix lgricina, at Whittemore, Michi- gan was found to be quite heavily infested with M, §cut§llatus (Say). Blackman and Stage (3) also reared this Species from Lgrix,1a21cina. Pippa magigna, in the Presque Isle fire area was found to be infested with this sawyer in 1939. in view of the above facts it would seem quite logical to assume that the hosts of M, scutellatus (Say) include the pines, Spruces, firs, and larches throughout its habitat. -38- SEASONAL HISTORY: Under normal Michigan conditions the life cycle requires but one year. This may be lengthened to two, three or even four or more years under adverse moisture or light conditions. (54, p. 261) (5, p. 71). About 95% of the beetles reared in outside cages during 1937 and 1938 com- pleted their deve10pment in one year. On October 4, 1939 three adult exit holes were observed in a jack pine tree kill— ed by the Presque Isle fire on May 6, 1939. A section of the trunk of this tree was removed for larval counts, and in the process, two pupae were found along with a dozen or more lar- vae. The above facts indicate that in some cases the life cycle is completed in less than six months. It seems improba— ble that oviposition had occurred in this tree earlier than May 6, 1939. Conditions in the Presque Isle area were ideal for rapid development due to an almost complete lack of shade and the warm, late autumn season of 1939. Adults of the white-Spotted sawyer are at large from about the middle of May to September and eggs may be deposit- ed at any time between those dates although the bulk of egg- laying probably occurs before the last of July. In depositing her eggs the female commonly chooses a Spot near the juncture of a branch with the trunk or twig with a limb. She cuts a slit (usually transverse) in the bark with her sharp mandibles, turns promptly around and deposits her egg.‘ In observing oviposition in the laboratory, the female, after cutting the Slit in the bark, appeared to pry it up with her ovipositor, after which the egg was inserted as far as possible under the slit and into the green inner bark. -39- In all observed cases Single eggs were laid. Blackman (3) tells of this species constructing "egg pits" in which several eggs are deposited. A thorough examination of bark in which eggs had been laid in the laboratory revealed deposition only in a manner described above. The maximum observed number of eggs laid by a Single female in captivity was five but this is probably less than the usual number deposited. Tragardh (88) states that the minimum number of eggs laid by Monochamus gptgn in Sweden is fifty. Attempts were made by the writer to arrive at the number of eggs normally deposited by sawyers in the following manner. Individual fertilized females were plac- ed in cages together with freshly killed white pine limbs suit— able for oviposition until death of the beetle occurred. Sub— sequent examination of the bark revealed one beetle as deposit— ing five eggs while three others laid only two each. The egg (Plate 111, fig. 2) is snow-white, opaque, elong- ate-oval and about one-eighth inch long. Eggs placed under what was considered optimum conditions required from seven to twelve days to hatch in the laboratory. The larvae when first hatched (Plate III, fig. 2) are of course very small with a head which seems large in proportion to the rest of the body. These legless, white "grubs" equipp- ed with powerful mandibles excavate broad shallow galleries in the phloem region but as they increase in Size penetrate not only the green inner bark but also the surface of the sapwood. Later the larvae enter the sapwood and eventually the heartwood in most cases. (Plates V, VIII). Entrance into the sapwood is always signified by the oval-Shaped holes near -40- the larval feeding excavation. (Plates IV, VII). The Space between the feeding excavations and the outer bark is common- ly crowded with tough, excelsior-like shreds of wood. Larvae hatching in late June up to mid-July usually have entered the sapwood by the first week in September. As the larval tunnels penetrate the sapwood toward the heartwood they increase in size to accommodate the growing larvae. Some of these gaping holes in the sapwood and heartwood measure a half inch in width. (Plate VIII). During this larval feeding period much boring dust or frass is pushed from the tunnels by the activity within. After entering the heartwood, the tunnels run longitudinally for a Short distance and then curves toward the surface making a U-Shaped gallery. The larvae apparently always strive to move toward light. Stark (80) has found that the larvae of Mgnochgmus change the direction of their course if logs are turned so as always to move toward light. Larval activity usually ceases in Michigan by about Octob- er 15, the winter, as a rule, being passed in this stage. Decem- ' bar and January examinations of infested wood showed most of the larvae to be in the heartwood. Feeding activity by the larvae is resumed during April in Michigan; this, of course, depending on prevailing temperatures. Full-grown larvae often measure one and one-half inches or more in length. (Plate VI). Before pUpation, the larval tunnel is extended out of the heartwood toward the surface to within a fraction of an inch from the inner bark. This will later serve as an exit for the adult when it transforms. The larval entrance to the pupal chamber is tightly plugged with excelsior-like frass and the -41- larva retires to the deeper, and more Spacious part of the burrow to pupate. (Plates IX, X). Larvae going through the winter in the most advanced stage are of course the first to pupate in the Spring. Pupation in Michigan may begin as early as the first week in May and continues through the early summ— er until Augustg Observations made in the laboratory indicate that from eight to Sixteen days are required for the pupal peri- od, with the average around twelve days. after transformation, the adult cuts its way to freedom by making use of its Sharp mandibles and emerges through a clean— cut, nearly circular exit hole (Plates XI, fig. 1; XII, XIII). Adults in Michigan have been observed to emerge as early as May 13, but under normal conditions the peak of emergence occurs in June. The adult insects are attractive, somewhat elongate, cylin- drical, black beetles more or less mottled with white. (Plate 1). The scutellum is invariably clothed with a heavy white pubescence, hence the specific name, scutellatus. The antennae are very long and filiform, giving the insect a terrifying appearance to those not familiar with it. The male antennae are twice or more times the length of the body while those of the female are only slight- 1y longer than the body. Size variations are quite pronounced in this Species; some beetles being more than one inch long, while others are less than a half-inch in length. (Plate II, figs. 1 and 2). The color varies from pure black (except the scutellum) in some individuals to those having the wing covers heavily mottled with a white pubescence. (Plate 111, fig. 1). ._42- The adults of the white-Spotted sawyer feed rather voraciously in captivity. Fresh white pine shoots were stripped of bark and needles by their feeding activity when introduced into breeding cages. (Plate XV). Blackman (3) reports feeding on coniferous twigs by adult beetles and Pierson (67) says feeding by Monochamus on underside of coniferous twigs causes them to die, the injury later Show- ing up as red or brown needles. Craighead (15, p. 106) reports feeding by adults on conifer needles to be extensive and that on Spruce trees near logging Operations sufficient branches are sometimes killed as to cause serious defoliation and mal- formation. Adults kept under favorable conditions in the laboratory, lived on the average about two weeks. Blackman (3) reported one beetle as living 31 days but this was probably an excep- tional case. CONTROL MEASURES ADVOCATED: Various methods have been rec- ommended for reducing sawyer injury to merchantable timber. Where possible, prompt utilization of felled or fire-killed timber is the best way to handle the problem. If this is im— possible, other methods may be resorted to, each of which has its disadvantages. Water storage (34, p. 262), (83), (21), (22) is said to prevent damage if ponds or rivers are Situated near enough to the logging region to make this practice conveni- ent. However, Rodd (69) says that larvae in floating trees oft- en survive as long as three months, the only consequence being a delay in their deve10pment. The removal of bark from logs will prevent egg—laying by v-r v43- Sawyers but this would be useless if the larvae had already penetrated into the wood. barking is an expensive process, making such operations on low grade, knotty logs unadvisable. Barking saw logs has one objection in that it causes checking. This is brought about by the more rapid drying and shrinking of the outer portions than the inner parts. Graham (34, p. 123), Dunn (22), and Swaine (83) (86) have found the use of lime sulphur dust on logs to be effective and have also recom— mended the covering of log piles with Spruce or balsam boughs. The use of insecticides and bough coverings does not insure complete protection against sawyers, and Should be resorted to only when it is impossible to employ other methods. nguugyymgg thjJJQtor (Fab.) (Monochamus titillator“ (Fab-D . (page titillator (Fab.)) Hopping (47) considers cgrglinensis Oliv.; mipgg Lec.; and dgntatoz Fab. to be synonyms of titillator. The southern-pine sawyer, Monochamus titillator (Fab.) (Plate XVI) has been reported by Webb (89) (90) (91), St. George and Beal (78), Craighead and Middleton (18), Craighead (15, p. 106) and others as being very destructive to storm-felled timber in the southern states. This species does not seem to be suf- ficiently widespread or numerous in Michigan to be of much econ- omic importance. A number of these beetles were reared from pine slash near Roscommon but it is a Significant fact that they seemed to be completely absent from the Presque Isle fire area. HOSTS: Hopping (46) gives the hosts of this sawyer as -44- Pippg alust 'S, g. stgopus and probable all pines in its habitat as well as Api§§_balsamea. This insect was reared only from pines in Michigan. DISTRIBUTION: The southern pine sawyer is said to be found throughout eastern North America, extending west to Ala- bama in the south and possibly to British Columbia and Alaska. SEASONAL HISTORY: The discussion of the white-spotted sawyer can be applied very closely to the southern pine sawyer in Michigan, there being but little, if any, variation in the habits, life cycle and injury of the two Species. Mgpochgmus ngtatps (Drury) (Mgnochamus confusor (Kby.) (Monochamus confuso; (Kby.) The grey sawyer, Monochamus potatus (Drury) (Plate XVII) did not appear to be of any importance in the localities under observation in Michigan, although Dunn (21), (22) and Swaine (84) consider it as a most important secondary insect in the coniferous forests of eastern Canada, being absent only from .a small section of the north shore of the Ga5pe Peninsula, Quebec. They report this Species along with the white-Spotted sawyer as following forest fires and completely destroying large areas of fire-damaged trees. Herrick (36, p. 417) has found the grey sawyer attacking living trees and Lang (56) reported an individual of this Species as boring through a lead pipe 2% inches thick. HOSTS: Hopping (47) lists the hosts as Pippg strobus, 2. banksiang, P. resinosa, g. pgnderosa, and Piece canadensis. 945- During the course of this study the grey sawyer was reared only from white pine. (g. stgobus). DISTRIBUTION: According to Hopping (47) the range of this insect is the northeastern part of the United Dtates and Canada including British Columbia. SEASONAL HISTORY: From white pine, in which eggs were de- posited during the summer of 1938, two adults emerged January 7, 1940. The wood was brought indoors during November, 1939, after being in outdoor cages Since the time of collection in the fall of 1938. This indicates a two-year life cycle for these individuals at least. Dunn (22) reports a development- al period of one year in eastern Canada if the summer is unus- ually warm but states that two years is the normal life-cycle period. The type of injury and habits of this Species are similar to other sawyers already discussed. Apgpthocinus pusillus (Kby.) (Plate XVII, fig. 1). The synonomy of this Species is quite confusing. Apparently ref— erences to Graphispgus pusillus (Kby.) (38) and Ceratographis pusillus (Kby.). (40) both pertain to the same insect, Acanthgcinus pusillus (Kby.). HOSTS AND DISTRIBUTION: This beetle has been variously reported as feeding on the dead inner bark of pines and Spruces in northeastern United States. Leng (60) lists its distribus tion as Maine and northern New York. In Michigan this borer was reared from jack pine, white pine, Norway (red) pine and black Spruce. AS already stated, the larvae mine only in the inner bark where they are not only harmless but also probably -4g- beneficial in that they seriously compete with bark beetles for this desired phloem-region food. SEASONAL HISTORY: Development is completed in one year, pupation taking place in a peculiar cell constructed just be- neath the bark by building up an elliptical barrier of frass. (Plate XIX). astylogsis guttata (Say) (As-triangle sexes-time (Say)) (Astylopsis commixta (Hald)) (Astylopsis pgpctata (Hald)) (M21125. W Say.) (Leptostylus commixtus Hald.) Astylgpsis guttata (Say) (Plate XVIII, fig. 2) has been reported by Craighead (15, p. 116) as commonly associated with Mongchgmus and Acanthocinus in pines and Spruces throughout eastern United States. Blackman (3) reared it from the tops and limbs of larch (ngig lagicina). Leng (60) gives its dis— tribution as Canada, District of Columbia, Wisconsin and In- diana. Adults were reared in Michigan from white pine in each of the three years of study. Blackman (3) found the develop- ) mental period to extend over two years. All individuals rear~ ed during the course of this study in Michigan required but one year for deveIOpment. The larvae do not appear to penetrate even the sapwood to a very great extent and thus cannot be con— sidered as particularly injurious. All observed beetles of this Species were associated with the white-spotted sawyer, Monochamug scutellatpg (Say). -47- Asemum moestum Hald. Leng (60) records this beetle (Plate XX, fig. 1) as being distributed from Alaska south to Lake Superior and Florida. Craighead (15) reports it as breeding in Picea, Larix and Apipg throughout eastern and Central North America. Packard (65) re— ported this Species as attacking healthy white pine trees, and Hopkins (37) found A, moestum in Spruce and yellow pine. Ac- cording to Blackman (5) it is probable that this borer will be found in a large number of coniferous trees throughout its range. He reared the beetle only in the basal portions of larch trunks and states that A. moestum is one of the primary insects attacking the weakened tree, often attacking trees be- fore they Show any visible Signs of weakness —— sometimes a full year before the entrance of Dendzoctonus Spp. A. mgeptpm_was reared from dead white pine near East Lans- ing, Michigan, adults emerging on May 29 and June 3, 1938. SEASONAL HISTORY: Development of this Species was completed in one year at East Lansing. Blackman (3) reports the life cy— cle as ordinarily requiring two years and Craighead (15, p. 32) says one or.often two years are needed. The larvae were found to have made extensive borings deep into the heartwood of white pine limbs causing injury very sim— ilar to that of Mgnochamus. It was also observed that this Spec— ies was reared only from the moist portions of white pine limbs next to the soil. (32). r48- ngonocherus mixtus Hald. This interesting little Species (Plate XX, fig. 2) is said to breed in pines and Spruces throughout the eastern and west central United States and Canada. Blackman (3) bred this insect from larch and Leng and Hamilton (59) reported it on pear and willow trees. Adults of this Species emerged in considerable numbers from white pine logs and limbs near East Lansing between May 27 and June 26, 1938. It was not reared from any other locality. SEASONAL HISTORY: The life cycle of all observed individu- als at East Lansing required but one year for completion. ‘Black— man (3) reported the life cycle as extending over a period of two years in larch. ‘ The larvae cut irregular grooves in the sapwood which are surprisingly wide and deep for so small a borer. No instances of this borer extending tunnels into the heartwood were observ- ed, pupation taking place in the sapwood close to the bark sur- lface. Accordingly, P. mixtus cannot be considered as particu- larly damaging in Michigan. OTHER SPECIES Other Species of Cerambycidae reared from dead coniferous timber in Michigan are as follows: AQOngdega (Leptura) canadgnsis (Fab.) (Plate XXI, fig. 1) Anoplodera (heptura) vittata (Oliv.) (Plate XXI, fig. 2) Aguagops proteus (Kby.) (Plate XXVI, fig. 1) Typggegus velutina (Oliv.) -49- Hhagigm lineatum (Oliv.) ‘ Qallidium aaienasium.(wewn.) The above were either reared in such few numbers or were associated with so many others that detailed observations on their injury and habits were impossible. The developmental period, hosts and emergence dates can be obtained from the emergence tables. Buprestidae Chrysobothris scabripennig C. & G. DISTRIBUTION: This metallic wood borer (Plate XXII, fig. 1) is distributed over northeastern United States and Canada west to Lake Superior. (ll, p. l69),(60). HOSTS: Chamberlin (11) lists the hosts as white pine (Pings stzgpus), Spruce (£1§§§_sp.) and hemlock (Tspga,ganadensis). Considerable numbers of this Buprestid were reared from white pine slash at Roscommon, Michigan where it Was commonly associat- ed with Monochamus scutellatus (Say). (Plate XXIII). SEASONAL HISTORY: Two years were required for deveIOpment under Michigan conditions. All the adults seem to mature and emerge at approximately the same time. Seventy—two Specimens of this beetle emerged from indoor cages of white pine limbs be- tween January 26 and February 2, 1940. These cages had been outdoors from the fall of 1938 to November, 1939, at which time they were brought indoors. 1t is reasonable to assume that the beetles would have emerged in the summer of 1940. The flat— headed larvae of this Species (Plate XXII, fig. 2) excavate -50- broad channels entirely in the inner bark and sapwood, rarely penetrating the heartwood at any stage of their deveIOpment. (Plate XXIV). Pupal cells are constructed in the sapwood close to the bark (Plate XXV). The larval entry holes into the sap- wood are very much flattened while the adult exit holes are broadly oval. (Plate XXIII). The larval tunnels, like those of other Buprestids are tightly packed with a fine boring dust or frass. This Species like a closely related form, Chrvsobothris dentipes (Germ.), seems especially well adapted to breeding in thin-barked branches or slash. The larvae of such insects have been found by Graham (51) to be able to withstand the higher temperatures which prevail in the thin-barked portions of trees. Considering the boring habits of Chrysobothris scab i ennis, it must be concluded that this Species has very little economic importance. Two related forms of the same genus which closely resemble Q. scapzipennis are 9. blanchagdii Horn. and Q. dentipes (Germ.). The former was reared only from white pine while the latter was bred from jack pine and Norway (red) pine as well as from white pine. The habits, life cycles, and injuries of these two Spec- ies closely resemble those of Q. scabripennis. -51- Chaleonhora virginiensis Drury. This striking beetle (Plate XXVI, fig. 2), one of the largest of the family Buprestidae, was reared only from the basal portion of one fire-scorched white pine tree near Ros— common. DISTRIBUTION: Chamberlin (ll) records the range of this beetle as Ontario, to Florida, west to Louisiana, Iowa, Illinois, and Michigan. HOSTS: Chamberlin (11) lists the hosts as Pings trobus, 2.21g1da, P. echinata, E, tagda, E. palustris, and Taxodium distigum (Bald cypress). SEASONAL HISTORY: Seven beetles emerged from indoor— caged white pine between January 26 and February 6. 1940,_ which would indicate a life cycle of two years. Buttrick (9), indicates that deveIOpment for these beetles may be completed in a Single year. The mature larvae of this Species are very large, often measuring 40 to 50 mm. in length. Wood attacked by these lar- vae soon becomes completely riddled with their long, flat, winding tunnels (8). Hopkins (39) reported this species to be completely riddling mature white pine timber. According to Buttrick (9), Chalcoghora virginiensis or one of its closely related western forms was almost completely responsible for deterioration of coniferous timber after fires in the Black Hills of South Dakota. This does not seem to be the case in Michigan forest-fire areas, although adults of this Species seem to be quite common in the northern counties, eSpecially around saw mills, during late May and throughout June. GENERAL CONCLUSIONS AND SUHQARY Fire-killed coniferous timber in Michigan does not appear to have any more attraction to wood—boring Coleoptera than timber killed by other agencies. Scorched trees are readily attacked and damaged but emergence records indicate that slash and storm felled trees are even more heavily infested. Fires occurring during the late spring or early summer months render the scorched timber more susceptible to borer attack than fires occurring at other times. Timber killed by any agency is more attractive to wood- boring Coleoptera during the first year after death of the trees. While certain species prefer wood in a more advanced stage of decay, these are cxf little im- portance as far as injury to timber is concerned. Borers such as the sawyers require freshly killed trees and will not as a rule re-infest trees from which they emerge as adults. Examinations of dead timber caused by spring fires re- vealed very heavy damage to merchantable timber by secondary wood borers. The bulk of this injury was confined to the trunks and larger branches; damage to small trees (three-inch diameter and less) as well as the smaller branches of larger trees was negligible. 5. _ 54 - Rather limited examinations of forest trees surround~ ing Michigan slashings and burned-over areas indicated that such areas are not a menace to adjacent healthy timber. The concentration of secondary wood-borers in burned over areas is very high, resulting in the death of some fire- injured trees which otherwise might have recovered, but it does not apnear that borers attacking dead timber can successfully adapt themselves to breeding in healthy trees to any great extent. The White-spotted sawyer, honochamus scutellatus (Say), was responsible for nearly all damage to dead coniferous timber observed in Hichigan. While other species of wood-boring beetles contribute to the ultimate destruc- tion of the wood, they are not present in large enough numbers to be considered of much importance. A study of the biology of the white-spotted sawyer, Monochamus scutellatus (Say), indicates that under normal.conditions in Michigan the developmental period is one year. Individuals developing under adverse moisture or light conditions may require a much longer time o 10. ll. ..55... Natural control of wood-boring beetles by parasites and predators did not seem to be of much significance in the areasinvestigated. Woodpeckers and Hymen— opterous parasites of the family Ichneumonidae appeared to be of the most importance. The amount of fungous infection in the heartwood of coniferous timber one or two years after death of the trees, appeared to be definitely correlated with the extend of heartwood boring by Monochamus scutellatus (Say), The fungous most commonly present was Peniophora g2. and was observed only in the heartwood of logs penetrated by sawyer tunnels. Larval counts and adult emergence records of Michigan fire-killed timber indicate white pine (Pinus strobus) as the favorite host for sawyers as well as other wood boring Cerambycidae and Buprestidae. These records show NorWay pine (Pinus reSinosa) to be much less heavily infested than other coniferous tree species. (l) (2) (3) (4) (5) (6) ('7) (8) _56_ LITERATURE CITED Beal, F. E. 1911. Food of woodpeckers of the United States. U. S. D. A. Biol. Survey Bul. 37. Beal, J. A. 1935. Deterioration of fire-killed Douglas fir. Timberman, vol. 37, no. 2, pp. 13-17. Blackman, M. W. and Stage, H. H. 1918. Notes on insects bred from the bark and wood of the American larch. N. Y. State College Forestry, vol. 18; Tech. publication no. 10; no. 4, pt. 1, 115 pp. Blackman, M. W. 1919. Notes on forest insects. Psyche, vol. 26, no. 4, pp. 85-96. Blackman, M. W. 1931. A revisional study of gnathotrichus. Eich. Jour. Wash. Acad. Sci., vol. 21, pp. 223-236. Blatchley, W. S. 1910. Coleoptera or beetles known to occur in Indiana. The Nature Publishing 00., Indianapolis. 1385 pp. _ Boyce, J. S. 1921. Fire scars and decay. Timberman, vol. 22, no. 7, p. 37. Burke, H. E. 1909. Injuries to forest trees by flat-headed borers. U. 8. Dept. Agr. Yearbook, pp. 399-415. (9) (10) (ll) (12) (15) (14) (15) (16) -57... Buttrick, P. L. 1912. Notes on insect destruction of fire-killed timber in the Black Hills of South Dakota. Jour. Econ. Ent., vol. 5, pp. 456-464. Casey, T. L. 1909. Studies in the American BupreStidae. Proc. Wash. Acad. 1 Science, vol. 11, pp. 47-178. Chamberlain, W. J. 1926. Catalogue of the Buprestidae of North America north of Mexico. 289 pp. - Craighead, F. C. 1920. Direct sunlight as a factor in forest insect control. Proc. Ent. Soc. Wash., vol. 22, no. 5 pp. — Craighead, F. C. 1921. Hopkins host- selection principle as related to certain Cerambycid beetles. Jour. Agr. Research, vol. 22, pp. 189-220. Craighead, F. C. 1922. EXperiments with spray solutions for preventing insect injury to green logs. U. S. Dept. Agr. Bul. no. 1079. 11 pp. Craighead, F. C. 1923. North American Cerambycid Larvae. Dom. of Canada Dept. of Agr. Tech. Bul. no. 27, ns., 239 pp. Craighead, F. C. 1925. The Dendroctonus problem. Jour. Forestry, vol. 23, no. 4, pp. 340-354. (17) (18) (19) (20) (21) (22) (25) (24) - 58 — Craighead, F. C. 1927. Relation of insects to slash disposal. U. 8. Dept. Agr. Dept. Circ. 411. Craighead, F. C. and Middleton, W. 1930. An annotated list of the important North American forest insects. U. S. Dept. Agr. Misc. Pub. no. 74. Cushman, Rd A. 1926. Some types of parasitism among the ichneumonidae. Proc. Ent. Soc. Wash., vol. 28, pp. 29—51. Doane, R. w., Van Dyke, E. c., Chamberlain, W. J., Burke, H. W. 1936. Forest insects. McGraw-Hill Book Co., Inc. New York. 463 pp. Dunn, M. B. 1931. Sawyer beetles in pine and balsam fir. Canada Dept. Agr. Div. For. Insects, sp. circular. Dunn, M. B. 1931. An investigation of Monchamus beetles and their control. 22nd. Ann. Rep. Quebec Soc. Prot. Plants, pp. 86-88. Dunn, M. B. 1936. The function of wood-boring insects in the development of the forest. Rep. Ent. Soc. 0nt., vol. 66, pp. 8-11. Felt, E. P. 1906. Insects affecting park and .woodland trees. N. Y. State Museum Memoir 8. ..59.. (25) Felt, E. P. and Bromley, S. W. 1938. Insect damage to trees as consequence of the New England hurricane. Science, vol. 88, Sup. for Nov. 4, 1938, p. 12. (26) Fiske, W. F. 1907. Notes on insect enemies of wood-boring Coleoptera. Proc. Ent. Soc. Wash., vol. 9, pp. 23-27. (27) Carver, R. D. 1939. The national timber stand. Jour. Forestry, vol. 37, pp. 92-96. (28) Graham, S. A. 1921. Controlling insects in logs by exposure to direct sunlight. Jour. Forestry, vol. 19, pp. 512-514. (29) Graham, S. A. 1922. Effect of physical factors in ecology of certain insects in logs. 19th. Rep. Minn. State Ent., pp. 22-40. (30) Graham, S. A. 1922. Some entomological aspects , of the slash disposal problem. Jour. Forestry, vol. 20, pp. 437-447. (31). Graham, 8. A. 1924. Temperature as a limiting factor in the life of subcortical insects. Jour. Econ. Ent., vol. 17, pp. 377-383. (32) Graham, S. A. 1925. The felled tree trunk as an ecological unit. Ecology, vol. 6, pp. 397-411. (33) Graham, 8. A. 1930. Ornithology and forest entomology. Mich. Acad. 801., Arts and Letters, vol. 11, pp. 389-397. (54) (36) (57) (59) (40) (41) -60... Graham, S. A. 1939. Principles of forest entomology. HoGraw-Hill Book Co., Inc. 410 pp. Hanson, H. S. 1939. Ecological notes on the Sirex wood wasps and their parasites. Bul. Ent. Res., vol. 30, pt. 1, pp. 26-27. Herrick, G. W. 1935. Insect enemies of shade trees. Comstock Publishing Co., Ithaca, New York, 417 pp. Hopkins, A. D. 1889. Report on investigations to determine the cause of unhealthy conditions of the spruce and pine. W. Va. Agr. Exp. Sta. Bul. no. 56, pp. 197-461. Hopkins, A. D. 1893. Catalogue of West Virginia forest and shade tree insects. W. Va. Agr. Exp. Sta. Bul. no. 32. 82 pp. Hopkins, A. D. 1898. Buprestid larvae destructive to living white pine timber. U. S. Dept. Agr. Div. of Ent. Bul. no. 17, pp. 47-48. Hopkins, A. D. 1904. Catalogue of exhibits of insect enemies of forests and forest products. U. S. Dept. Agr. Div. of Ent. Bul. no. 48. 55 pp. Hopkins, A. D. 1904. Insect injuries to forest products. Yearbook, 1904, pp. 381-398. (44) (45) (46) (47) (48) - 61 - I Hopkins, A. D. 1909. Insect depredations in North American forests and practical methods of prevention pnd control. U. S. Dept. Agr. Bur. of Ent. Bul. 58, part 5. Hopkins, A. D. 1910. Insect injuries to the wood of dying and dead trees. U. S. Dept. Agr. Bur. of Ent. Cir. no. 127. Hopkins, A. D. 1910. Insects in their relation to the reduction of future supplies of timber and general principles of control. U. S. Dept. Agr. Bur. of Ent. Circ. 129. Hopkins, A. D. 1912. Damage to the wood of fire-killed Douglas fir and methods of preventing losses in western Washington and Oregon. U. S. Dept. Agr. Bur. of Ent. Circ. 159, 4 pp. Hopping, R. 1915. The entomological aspects of slash disposal. Soc. Am. Forestry Proc., vol. 10, pp. 183-185. Hopping, R. 1921. A review of the genus Monochamus Serv. (Cerambycidae, ColeOptera.) Can. Ent., vol. 53, pp. 252-258. Horn, G. H. 1886. A monograph of the Species of Chrysobothris inhabiting the United States. Trans. Am. Ent. Soc., vol. 13, pp. 65-144. (49) (50) (51) (54) (55) (56) (57) Houser, J. S. 1931. Damage to lumber caused by insects. Ohio Agr. Exp. Sta. Bul. no. 478, pp. 41-46. Howard, L. 0. 1919. Report of Entomologist. Hubbard, H. G. 1897. The Ambrosia beetles of the United States. U. S. Dept. Agr. Bur. of Ehto BUl. (DS.), n0. 7, pp. 9-300 Hubert, E. E. 1921. Notes on sap stain fungi. Phytopathology, vol. 11, pp. 214-224. Hyslop, J. A. 1938. Losses occasioned by insects, mites and ticks in United States. Bureau of Ent. Publication. Jaenicke, A. J. 1921. Relation between fires and insect damage. Timberman, vol. 22, no. 3, pp. 113-116. Kellogg, R. S. 1909. Timber supply of United States. U. 3. Dept. Agr. For. Service Circ. no. 166, p. 6. Lang, F. 1919. Insects damaging lead. Ent. Monthly Hag., 3rd. series, nos. 60 and 61. pp. 278-279. Leach, J. G., Orr, L. W., and Christensen, C. 1934. Interrelationships of bark-beetles and blue-staining fungi in felled Norway pine timber. Jour. Agr. Research, vol. 49, pp. 315-341. (58) (59) (60) (61) (62) (65) (64) - 55 _ Leach, J. G., Orr, L. W., and Christensen, C. 1937. Further studies on the interrelationship of insects and fungi in the deterioration of felled Norway pine logs. Jour. Agr. Res., vol. 55, no. 2, pp. 129-140. Leng, C. W. and Hamilton, J. 1896. The Lamiinae of North America. Trans. Am. Ent. Soc., vol. 23, pp. 101-178. Leng, C. W. 1920. Catalogue of the Coleoptera of America north of Nexico. Pub. by John D. Sherman, Mt. Vernon, N. Y. 470 pp. Merrill, J. H. 1915. On some genera of the pimpline Ichneumonidae. Trans. Am. Ent. Soc., vol. 41, pp. 109-154. Miller, J. M. and Patterson, J. E. 1927. Preliminary studies on the relation of fire injury to bark beetle attack in western yellow pine. Jour. Agr. Res., vol. 34, pp. 597-613. Mudinger, F. G. 1924. Apreliminary list of the Buprestidae and Cerambycidae of Cranberry Lake Regions, New York. N. Y. State College For. Tech. Pub. no. 17, part 4, pp. 313-320. Patterson, J. E. 1927. The relation of highway slash to infestations by the western pine beetles in standing timber. U. S. Dept. Agr. Tech. Bul. no. 3. (65) Packard, A. S. 1890. U. 5. Ent. Com. 5th. Rep., pp. 697-699. (66) Pierson, H. B. 1923. Insects attacking forest and shade trees. Maine For. Ser. Bul. no. 1, p. 28. (67) Pierson, H. B. 1927. Manual of forest insects. Maine For. Ser. Bul. no. 5, pp. 105-106. (68) Prozorov, S. S. 1929. (Burnt areas in pine forests as foci of infection) Trud. Sib. Inst. Sel. - Khoz Lesovod, vol. 12, pt. 3, reprint, 54 pp. 17 figs., 6 graphs, 28 refs. (In Russian). (69) Rodd, E. 1914. (Multiplication of Monochamus on pines in woods of Altai district). For. Jour., vol. 44, nos. 6-7, pp. 1048-1064. (70) Rohwer, S. A. 1921. The North American Ichneumon flies of the tribes Labenini, Rhyssini, Xoridini, Odontomerini and Phytodietini. Proc. U. S. Natl. Museum, vol. 57, pp. 405-474. (71) Rumbold, Caroline. 1931. Two blue-staining fungi associated with bark-beetle infestation of pine. Jour. Agr. Res., vol. 43, pp. 847-873. (72) Salman, K. A. 1934. Entomological factors affect salvaging of fire injured trees. Jour. of Forestry, vol. 32, no. 9, pp. 1016-1017. (75) (74) (75) (76) (77) (78) (79) (80) Schedl, K. E. 1931. Morphology of the bark- beetles of the genus Gnathotrichus. Smithsonian Misc. Coll., vol. 82, no. 10. Show, S. B. and Kotak, E. I. 1925. Fire and the forest. U. 8. Dept. Agr. Dept. Ciro. 358., pp. 6—7. Snyder, T. E. 1927. Defects in timber caused by insects. U. S. Dept. Agr. Bul. no. 1490. 46 pp., 45 figs., 51 refs. Sokanovskii, B. 1929. (On effect of insects in burned over areas). Defenses des Plantes, vol. 5, no. 5-6., pp. 667-668. (In Russian). St. George, R. A. 1929. Protection of log cabins, rustic work and unseasoned wood from injurious insects. U. S. Dept. Agr., Farmers Bul. no. 1582, 19 pp. St. George, R. A. and Beal, J. A. 1929. The southern pine beetle, a serious enemy of pines in the south. U. S. Dept. Agr., Farmers Bul. no. 1586, 18 pp., 17 figs., 2 refs. Stark, V. N. and Ctaph, B. H. 1925. (Importance of forest fires as creating foci of bark-beetles in Briansk government). Defense des Plants, vol. 2, no. 4—5, pp. 205-212. (In Russian). Stark, N. K. 1926. (Some observations on the life of Longicorns). Defense des Plantes, vol. 3, no. 4-5, pp. 343-349. (In Russian). (81) (82) (85) (84) (85) (86) (87) -66... Stickel, P. W. and Marco, H. R. Forest fire damage studies in the northeast. III, Relation between fire injury and fungal infection. Jour. Forestry, vol. 34, no. 4. pp. 420-423. Strickland, E. H. 1928. Can birds hold injurious insects in check? Sci. honthly, vol. 26, pp. 48-56. Swaine, J. m. 1916. Some features of interest in connection with our studies of forest and shade tree insects. Ont. Ent. Soc. Report, no. 47, pp. 96-97. Swaine, J. M. 1917. The large pine sawyer, Monochamus confusor. Ont. Ent. Soc. Report 471, pp. 96-97. Swaine, J. M. 1925. The factors determining the distribution of North American bark-beetles. Can. Ent., vol. 58, pp. 261-266. Swaine, J. M. 1928. Forest entomology and its development in Canada. Pamph. Dept. Agr. Canada, no. 97, 20 pp. Tothill, J. D. 1923. Injury to fire-killed lumber in New Brunswick by the softwood borer. 63rd. Annual Report Crown Land Dept., New Brunswick, pp. 86-87. (88) (89) (90) (91) Tragardh, I. 1929. (On the injury caused by the pine sawyer, Monochamus sutor and its prevention). Medd. Skogsforsoksanst, no. 25, pp. 171-288. Sweden. Webb, J. L. 1909. The southern pine sawyer, Monochamus titillator. U. S. Dept. Agr. Bur. of Ent. Bul. no. 58, pt. 4. Webb, J. L. 1911. Injuries to forests and forest products by round-headed borers. U. S. Dept. Agr. Yearbook, 1910, pp. 341-358. Webb, J. L. 1911. Monochamus titillator probes lumber business. Am. Lumberman, July 15, p. 48. -68- PLATES I‘Ln‘l‘ll I . Adult male and female of the white- Spotted sawyer, Monochamus scutellatus (Say). Male on right. K 2. I‘LL 1‘77. I . Fig. 1. Size variation in males of Honochamus scutellatus (Say). _‘._ I Fig. 2. Size variation in females of Honochamus scutellatus (Say). 1142? III. Big. 1. Color and size veriation in females or fionochamus scutellatus (Say). Fig. 2. Egg and newly hatched larva of Monochamus Scutellatus (Say). .X 4. ILLTE IV. 5'. 4:5:L-t Sawyer work in sapwood of jack pine. Note nearly circular exit holes of adults and the more flattened entry holes of larvae. PLATE V. O -- . -~ . ......~--«.~- .‘ ‘ ' ' a. ““31" 71-min ’ “may... in . , ‘ v ‘ .A'W-ku -. . .. w“ “'1 -MJu‘~ "-1 ~' ‘1!!! 7 _: an --—$-.,...";-;.Lar' sawyer work in heartwood of jack pine. PLATE VI. 3'“ ""‘. ‘ “cur—K #9:, ‘7 ‘.. >-. - ' . . 3‘“: v. .. ' wwd" ,JLO‘J '9 '5‘“ 4 V ' *W‘“ --‘m warm" “5': ~ Longitudinal section of jack pine showing mature larvae of Monochamus so utellatus (Say). Note pupal chambers and adult exit tunnels in upper part of picture. X 1/5. EIATE VII. Section of white pine limb showing sawyer and bark beetle association. PLATE VIII. _ O-— s 11.. ‘4lwullwfluxfl! ”if... .. i .3“ ._._ .53.. E. ._.:.. 55...... .2: . O. .l . .5 . 1 . .. fJAW. fim‘fifl‘ga’r :1: V’.« U pig} .- ..- .3... .I.. 3.55%.). . ....5%. .xx inn-'57.. u 2%.»: | I.‘- : .o..89 h gain. o "be .9.......-. xvqn, 5“ £2.3’- 33.2.2 4......1 fie .. . a... a II. J. 33:14.! .arsumwmumana I”.I {in}! Sawyer tunnels in heartwood of Jack pine. (Slightly reduced). ‘5 , 11‘ ‘P .LL.14L-J 1‘5. .. ‘. p a . a. ’2.“ ‘ ' ~ In ‘ci:::.: : *‘~+-z~w¥ .. . .. ’ v ' '~ ~ ' ,.- i -. ,_ . g _ Hs‘. arc-«16:44 M _' .‘“ m 53S: I‘M-u- .. - g,’ V" ~ . .— WW ‘ M _- 4.4;: (“I palm” . I—cgaol'ihl Q51‘ “5‘.” «2.123% Section of white pine log showing larva, pupa and newly transformed adult of Monochamus scutellatus (say). X 2/3. . _.. ....._.tnu..”~..¢wm....fl. . ...x...ac.«...a.1 sari... . , ml... 3...: knit... Q. ...l..1..._.. ..111...11- 1?... Q. o 0'. s PLATE X. .\.vs. i... 2.... .r. u» t .._.c. .W..{Fm “MM”... . . .& ._.‘.I.‘1‘ .p.i:1..;c .4... p-_ Q X 1%. k pine log showing pupal chamber Jac and pupa of Monochamus scutellatus (Say). Section of '1) f LL4LL-- 4'». o -- >‘___ -_ . . . c - v A“ Fig. 1. Adult exit holes of Monoohamus scutellatus (Say). (Natural size). Fig. 2; Mites on head and thorax of white-spotted suw'er. X &%. 1“.” ‘.'.1I,‘ .L'ulLlu ;{I I o White-spotted sawyer adult emerging from white pine limb. {Slightly enlarged). XIII. H L ‘J ELK -_.‘ - ' I -.—_-—- - - Exit hole of adult sawyer in ( Slightly reduced ). white pine limb. ILATE XIV. Section of balsam fir limb showing fungous growth in sawyer tunnels. PLATE XV. Injury to white pine twigs by adult feeding of monochamus scutellatus (Say). ’ ~ 1" 1r??? .LL;‘..L.J ‘LJJ... Adult male and female of the southern pine sawyer, Monochamus titillator (Fab.). Male on left. (Slightly enlarged). Adult male of the grey sawyer, nonochamus notatus (Drury). X 1%. 1214:1132 -.V I I I o Fig. 1. Adult male and female of Acanthocinus pusillus (airby). Hale on right. i 2. --¢ . . - - Fig. 2. Astzlopsis guttata (Say). X 5. ILATE XIX. .. --._.._————-__..__ -_. o..-- - _.. . - .. Section from black spruce limb with bark removed showing pupal cell of Acanthocinus pusillus (Lirby). (Natural size). ’7‘ "'3" V‘.’ J. .14..) aux. Fig. 1. Adults of lsemum moestum Bald. .51 4.3;. Fig. 2. Adults of Pogonocherus mixtus Hald. X 2%. Rig. 1. enoplodera canadensis (Eab.). Fig. 2. Anoplodera vittata (Oliver). X2. lIATE LXII. ‘ 1' vs Fig. 1. Adult beetles of flat-headed borer, Chrysobothris scabripennis C. & G. X 1%. Fig. 2. Larva of flat-headed borer, Chrysobothris scabripennis C. & G. X 2. Section of white pine limb showing sawyer and Buprestid feedinj excavations in the sapwood. l, wor; or the Buprestid, Chiysobethris scabripennis C. a }.; 3, work of :Larxmx,2"onochamus Sp.; C, larval (v entry holes of honochamus sp.; 3, larVal entry holes of C. scahrllenhis; 7, adult exit holes of C. sclbripennis. ILAIT XXIV. Cross section of white pine limb showing larval tunnels of fionochamus sp. in heartwood and Chrysobothris scabripennis in the sapwood. ILaTE AXV. Section Of white pine limb showing mature larva of Chrysobothris scabripennis U. & G. just prior to pupation. -X 1%. Fig. l. Fig. 2. PLAIE XXVI. Adults of Acmanps Lroteus (Lirby). Adults of Chalc0phora virgihiensis Drury. X 1%. ILIIE XXVII. Fig. 1. Adults of sawyer parasite, Ichneumon (Ephialtes) mesocentrus (Grav.). Fig. 2. Section of white pine limb showing cocoon of Ichneumon mesocentrus in larval tunnel of Monochamus 3p. PLATE XXVIII. Section from heartwood of white pine limb showing puparia of a Tachinid fly in Monoohamus tunnel. . '.'.J #:3’,“ prxm ~. .. ..-“* .. ‘ - fag-#2 ' 3' . L .I ‘1’ it" 11" 2i Q ’l\ ' 'v'.“ '1' a ‘ . 6; . _%x. K. 'o" . 31' ‘ H}. "r’; - - .23}: .' 1‘; -' iii.“ 1\ ' ‘t5‘1‘7 .1. ,1 .. . ‘3"’.";I' 1' ‘3.)- h- -. 7w .~ 3‘1 1.3% (A .. .' 14 " .3. *z‘ ‘1. ’5‘ ' "1‘ 7".0' 1" 1 1". ‘ {1‘ 1, .15- o... ,‘ '41::- 3" 3'1 .6; A. . .I. v... - ' . 1.1;, . if} ‘\i I i" » ' .1. . .41‘iL‘fizl ‘ .4 t 7 IO U \ ' i." . ' o 1‘ I. .\ MICHIGAN STATE UNIVERSITY LIBRA 293)" (III III lllllllllll