Digitized by the Internet Archive in 2011 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/morphologytaxono12haye ' / ILLINOIS BIOLOGICAL MONOGRAPHS Volume XII PUBLISHED BY THE UNIVERSITY OF ILLINOIS *, URBANA, ILLINOIS I EDITORIAL COMMITTEE John Theodore Buchholz Fred Wilbur Tanner Charles Zeleny, Chairman S70.S~ XLL '• / IL cop TABLE OF CONTENTS Nos. Pages 1. Morphological Studies of the Genus Cercospora. By Wilhelm Gerhard Solheim 1 2. Morphology, Taxonomy, and Biology of Larval Scarabaeoidea. By William Patrick Hayes 85 3. Sawflies of the Sub-family Dolerinae of America North of Mexico. By Herbert H. Ross 205 4. A Study of Fresh-water Plankton Communities. By Samuel Eddy 321 ■ LIBRARY OF THE UNIVERSITY OF ILLINOIS ILLINOIS BIOLOGICAL MONOGRAPHS Vol. XII April, 1929 No. 2 Editorial Committee Stephen Alfred Forbes Fred Wilbur Tanner Henry Baldwin Ward Published by the University of Illinois under the auspices of the graduate school Distributed June 18. 1930 MORPHOLOGY, TAXONOMY, AND BIOLOGY OF LARVAL SCARABAEOIDEA WITH FIFTEEN PLATES BY WILLIAM PATRICK HAYES Associate Professor of Entomology in the University of Illinois Contribution No. 137 from the Entomological Laboratories of the University of Illinois T U .V- TABLE OF CONTENTS 7 Introduction Q Economic importance Historical review Taxonomic literature 11 Biological and ecological literature 12 Materials and methods 1%i Acknowledgments Morphology ]* The head and its appendages 1 ' Antennae. 18 Clypeus and labrum ™ Epipharyru Mandibles. 22 Epipharynx Maxillae . 37 £ Hypopharynx 00 NO "* N HNN^HlOHlOHlfltON 13 IT) (N| V) N N H CO CS N ") 10 o H Ph CNOnO-^n-hOnOCNn— lOOOOO rtillioaO^^CMNNtJtj. o OMsiOtON'OOl'HOfOiOfO U f^ ,-H ,-H •r-l t-H *H TH T3 NtONONNfO^N | ^*0 cs Ih IT) « 00 O nO nO CO r>4 1 ■<# co o CO th ,-h 00 ■■-h ^h >h fO O co ■tj o»ONOON«ot^Noo^^ I "O <2 00 M NO CS O CO t— 1 C* T" 1 N I'TO cs CS On P4 00 n-h n-H cs or^rO'-toro-^frj'i 1 «o COOOOnOO-^'-iOnOnnO 1 »-» "H NO o n-h cs N-< n-h m U cs -a g On— ItOOOiNON'fOI'lNfO S Jh On nH csOn ifltONN NO CS On P< ONO\OoofO l ooo^~"oiopocs 3 IT) HN*NfNOOOfOOHOf- o U tH* no CO 00 CO N-H CO T3 a a n* 10 to to l 1 h 10 a iJSIa 1 1 cs n-h cs *- T— 1 CO •4 O cs t^ ,-H »H ,— 1 On OO^ON^NOCOI 1 »0 CO JO *g CSOCOt^ 1 |00 l O'TNO o t~- -* cs cs '-' * U ,-H ,-H TJ On On Ed U NO 1 * 1 IflfOIOfOHfOHHO JS 1 w n— 1 es no 10 cs CO • CSOO I TftO'ttJHCOHtot. o O 1 NO cs 00 O 00 CJ cs cs N-" " t3 u OS 00 CN CS 00 m n-H CS 1 INOt^N-H cs ,_, ^ v-H t^ CS n-H | | N-H CO oo Ch On Nt tN fN U5 H H 00 1 1 Oi -H 'H T* o CS T) «H CS •>* i-H 1 1 CO U ^H CS CS ^H T3 M ^H| I ^H CS CO CS N— II I ION On Ed rt 1 1 N^l N-H 1 1 1 CS On Ph j-^l icocsOnOn^hi 1 |p *-H O Jh 1 1 p "■* III*- g '"' 3 P4 NOVO-^nO I 00 N H I ICS'-H NO o CO N-H ■* CS 1 CO ' ' ,— I es ^ CS 00 n-H ■* 1 "* *- II W NO o U N-H CO en a tsi ft a t/1 a hJ 3 4-> 3 2 4- n a X 15 I/O 00 aS M 4) « 5 ■ : pq C c k c bo !/ O « i-5 P- 1 p 1 T 1- K 1 1 1* < a c 1 c ID § i tn H 56 ILLINOIS BIOLOGICAL MONOGRAPHS [140 37 species. The largest number of species belonging to a single genus was the 17 species of Phyllophaga. The other genera distributed among the various subfamilies of Scarabaeidae contained but one, two, three, or four species. In Table VI are shown the habitat preferences of all species collected. These are indicative of the food preferences as a whole but not as related to the individual species, which are shown in Tables VII and VIII. In Table VII it is shown that the most individuals of Phyllophaga were taken in blue grass sod, 234 specimens being found there. The next in preference is corn land, with 140 individuals, which is closely approached by the 126 TABLE VII— FOOD PREFERENCE OF PHYLLOPHAGA GRUBS Species cd 2 is a ■* PQ ed in •4-> cd O a H o U S3 in ed a, m M 60 H o 3 \A CO M 3 09 Pm V 3 d m cd u Eh o 3 in U en en g & » £ pq 5 to o a o U G u". o .3 HH CO eu u S3 en a Ph 3 T3 H o < en CU O o P4 d CU O en O d u en o H Ochrosidia immaculata . Ligyrus gibbosus Anomala binotata Anomala kansana Anomala innuba Anomala undulata Cotalpa lanigera Pelidnota punctata .... Ligyrodes relictus Euphoria inda Euphoria sepulchralis . . Aphodius sp 1787 125 122 235 5 113 2 32 2 1 8 1 143 189 164 2 70 1 479 17 4 8 28 49 2 114 9 21 8 1 79 4 1 1 48 2 3 4 1 10 11 1 120 41 6 53 2 4 3 2 12 141 19 1 6 10 1 138 1 2 1 199 24 9 3 9 1 1 4 122 1 77 3070 392 302 256 144 8 149 118 123 46 9 53 Canthon laevis Trox sp 4 122 Polymoechus brevipes . . Cremastocheilus nitens . Ataen us i nops Trichiotinus piger Stephanucha pilipennis . Polyphylla hammondi . . 12 21 81 8 1 1 Total 2389 44 569 536 204 143 248 17 167 11 142 450 4920 it occurs rather abundantly in corn and oats. The adults apparently show a preference for land that is frequently plowed. Ligyrus gibbosus exhibits the same preference, being found most often in oats and wheat and less often in corn land. The same can be said for Anomala binotata, while 58 ILLINOIS BIOLOGICAL MONOGRAPHS [142 A. kansana occurs more frequently in wheat. Cotalpa lanigera occurs in wheat and corn, usually in the sand-hill area. Davis (1918) has reported this species doing considerable damage to raspberry bushes, strawberries, corn, and grasses. The following species live in fallen logs and stumps: Pelidnota punctata, Polymoechus brevipes, Trichiotinus piger, and Polyphylla hammondi. Table VIII shows 21 individuals of Cremastocheilus nitens taken from logs. These were in ant nests on the surface of the soil under a log on the sand dunes. Several species show a preference for manure and decaying vegetation, such as rotten hay or straw stacks. Among these are Ligyrodes relictus, Euphoria inda, Euphoria sepulchralis, Ataenius inops, and Aphodius sp. The 122 individuals of Trox sp. listed in the miscellaneous column were taken under a dead horse. TABLE IX— COLLECTIONS OF PHYLLOPHAGA AS DETERMINED BY REARING Year of Collection Species 1916 1917 1918 1919 1920 1921 1922 1923 Total P. crassissima 15 36 40 33 196 52 18 6 396 P. rubiginosa 11 7 15 22 10 41 25 4 135 P. rugosa 14 4 9 33 22 24 14 5 125 P. lanceolata 2 16 9 50 1 — 1 — 79 P. submucida — 2 2 — — — 66 9 79 P. implicita 6 4 4 8 20 1 28 5 76 P. hirticula var. comosa — 2 — — 4 18 — — 24 P. praetermissa — — 3 — 8 — — 2 13 P. longitarsa — 1 4 — — — 3 — 8 P. bipartita — — 1 3 — — — 2 6 P. futilis — — — — 3 — — 2 5 P. corrosa — — 2 — — 1 — 2 5 P. glabricula — 2 3 5 P. fusca — — — — — 3 — — 3 P. crenulata — — 1 — — — 1 — 2 P. tristis 1 1 2 P. affabilis — — — — — 1 — — 1 Total 49 74 90 149 264 141 156 41 964 Relative Abundance J. W. McColloch and the writer, in an eight-year study of scarabaeid larvae, made collections of grubs in all kinds of situations and reared many to adulthood. The following discussion of the various collections, for convenience, is divided into two groups. The genus Phyllophaga, with 17 species represented, will be considered as one unit while the remaining 143] LARVAL SCARABAEOIDEA— HAYES 59 16 genera will be considered jointly. Table IX shows the various species as to the year in which they were collected and not the year of maturity. The order of arrangement is based on numbers collected and not on their relationships. As shown in Table IX, 964 individuals representing 17 species were reared from the total of 18,781 grubs of all species collected. Since many individuals of this genus failed to live through the rearing period, a per- centage of those reared based on the number collected would be unfair. However, the percentage reared based on the total species gives an inkling of the relative proportions of Phyllophaga to other species of the family. Of the total (5,884) beetles, only 964, or 16.3 per cent, belonged to this genus. Table IX, as indicated in the column of totals, offers further data on the relative numbers of all the various species found in the locality con- sidered. Of the 964 individuals of all species reared, P. crassissima ranks first, with a total of 396 individuals, or about 41 per cent; P. rubiginosa ranks second, with about 14 per cent; and P. rugosa third, with nearly 13 per cent. If we compare the individual total of each species with the total adults collected during the first seven years (1916-1922) of the eight during TABLE X— RELATIVE ABUNDANCE OF PHYLLOPHAGA ADULTS AND REARED LARVAE Total Adults Comparative Rank Species Larvae Collected Ranking of Reared 1916-1922 Adult Abundance 1. P. crassissima 396 31,996 First 2. P. rubiginosa 135 15,130 Third 3. P. rugosa 125 4,010 Fifth 4. P. lanceolata 79 15,851 Second 5. P. submucida 79 57 Eighteenth 6. P. implicita 76 982 Tenth 7. P. hirticula var. comosa 24 1,696 Ninth 8. P. praetermissa 13 115 Sixteenth 9. P. longitarsa 8 909 Eleventh 10. P. bipartita 6 3,523 Seventh 11. P. futilis 5 10,362 Fourth 12. P. corrosa 5 3,732 Sixth 13. P. glabricula 5 440 Twelfth 14. P. fusca 3 135 Fourteenth 15. P. crenulata 2 127 Fifteenth 16. P. tristis 2 114 Seventeenth 17 P. affabilis 1 44 Nineteenth 18. P. vehemens 2,001 Eighth 19. P. congrua 135 Thirteenth 60 ILLINOIS BIOLOGICAL MONOGRAPHS [144 which this study was in progress (see Table X), some interesting facts are brought out. P. crassissima, the most abundant species among the adults collected, is seen to be also the most numerous among those species collected as grubs. P. lanceolata, while second in number of adults, is fourth in the grub collections. This may be, in part, due to the slightly longer life-cycle of P. lanceolata (Hayes, 1919). P. rubiginosa, while third among the adults, was second among the grubs; and P. rugosa was fifth in the beetle collections and third among the larvae. On the whole, these four important species rank very close, while greater differences are disclosed among the other species. An interesting observed fact is that more indi- viduals of P. submucida were collected as grubs and reared to adults than were taken in the adult collections. The two species, P. vehemens and P. congrua, which ranked eighth and thirteenth, respectively, in the adult collections, were not reared from grubs. TABLE XI— THE RELATIVE ABUNDANCE OF SCARABAEID LARVAE OTHER THAN PHYLLOPHAGA Miscellaneous Species of Scarabaeid Larvae Year of Collection 1916 1917 1918 1919 1920 1921 1922 1923 Total Ochrosidia immacu- lata Ligyrus gibbosus Anomala binotata Anomala kansana Anomala innuba Anomala undulata Cotalpa lanigera Pelidnota punctata Ligyrodes relictus Euphoria inda Euphoria sepulchralis Aphodius sp Canthon laevis Trox sp. Polymoechu s bre vipes Cremastocheilus nitens Ataenius inops Trichiotinus piger Stephanucha pilipennis Polyphylla hammondi 32 5 145 1 13 25 2 17 4 2 155 2 3 9 10 10 1 20 2 155 2 1 45 1 19 723 227 37 154 21 1 1 1 2 315 23 42 4 1 7 74 14 4 4 1 973 116 47 57 15 7 23 5 6 6 2 12 1 81 1 715 27 37 53 1 125 92 2 53 122 3 1 3070 392 302 256 144 8 149 118 123 46 9 53 4 122 12 21 81 8 1 1 Total 221 25 212 223 1167 489 1352 1231 4920 145] LARVAL SCARABAEOIDEA— HAYES 61 In addition to the collections of Phyllophaga, 20 species representing 16 other genera of the family Scarabaeidae are available for comparison with the genus Phyllophaga. These were, for the most part, incidental to the white grub collections, but since all that were found were collected, the data available show the years of abundance as well as the relative numbers. These data are summarized for the various species and the years in which they were collected in Table XL It will be noted that the two most numerous species are Ochrosidia (Cyclocephala) immaculate (Oliv.) and Ligyrus gibbosus (DeG.). The number of Ochrosidia reared (3,070 indi- viduals) far exceeds that of any other species and is more than three times the number of all Phyllophaga combined, whose total figure is 964. It must be borne in mind, however, that the one-year life-cycle of Ochrosidia compared to the two-year and three-year cycles of Phyllophaga makes rearing much easier with less mortality. Space does not permit a dis- cussion of all the facts apparent in Table XI, but it will be noted that the collection contains representatives of the most important subfamilies of the Scarabaeidae. Hibernation Our knowledge of the depth to which such insects as white grubs and May beetles penetrate the soil to escape the rigors of cold weather, and to pass their period of hibernation, is limited to a small number of casual ob- servations. Only a few general statements pertaining to the subject are to be found in the literature. While writers assert that such-and-such a species passes the winter "below the frost line" or "below the plow line," no specific attention or careful study has been given to the subject. This is due, in part at least, to the difficulty of studying the habits of insects that live with- in the soil. Another factor involved, and perhaps the most important, is the difficulty of making specific identification of the immature stages of the specimens found in the soil. In most instances they must be reared through their developmental periods to the adult before they can be identified. Furthermore, winter studies of insects in the soil require careful excavation, involving considerable manual labor which must be done during the coldest and most disagreeable part of the year. Criddle (1918) records that in Canada the grubs of certain species of Phyllophaga and allied genera penetrate to a depth of 74 inches, and that the beetles may burrow as deep as 47 inches during the winter. This state- ment indicates at what depth grubs may be found in a more northern climate, but no data are available in regard to the actual depth of the vari- ous species in the more temperate regions of the United States. No doubt the climate has a direct bearing on the subject, and the depth of penetration will vary with the region. In fact, the present study shows that white grubs do not penetrate as deeply in Kansas as Criddle observed in Canada. A discussion of the literature of this subject has been presented by McCol- 62 ILLINOIS BIOLOGICAL MONOGRAPHS [146 loch and Hayes (1923), and the following is taken from a discussion by these writers in collaboration with H. R. Bryson. (1928). The depth at which these insects pass the winter is important in connection with the recommendation of fall, winter, or early-spring plowing as methods of control. In order to make such recommendations intel- ligently, it is essential to have definite information relative to the depth of hibernation of the insects. It was primarily to secure data along this line that the studies reported herein were undertaken. It also seemed desirable to further check the studies of McColloch and Hayes (loc. cit.) in the fall and spring reversals of temperature conditions on the surface and subsurface layers of soil, and the bearing of such changes upon the activities of soil insects in general. An attempt was made to rear to the adult stage all grubs taken in this work. A summary of these rearings is presented in Table XII, which shows the number of grubs of each species identified and the depths at which they were taken. The data on Phyllophaga lanceolata, which are incorporated in this table, were secured from a series of excavations in a wheat field at Goddard, Kansas, March 13, 1919. TABLE XII— SUMMARY OF THE DEPTH OF HIBERNATION OF WHITE GRUBS Depth in Inches Species Total Collected Weighted Maximum Minimum Average All white grubs 1,188 40 3 13.2 Qchrosidia immaculata 101 30 4 13.9 Phyllophaga crassissima 3 17 13 15.0 Phyllophaga rugosa 4 26 10 18.0 Phyllophaga glabricula 3 19 16 17.7 Phyllophaga submucida 1 14 14 14.0 Phyllophaga rubiginosa 1 14 14 14.0 Phyllophaga bipartita 1 15 15 15.0 Phyllophaga corrosa 1 16 16 16.0 Phyllophaga lanceolata 66 20 3 10.3 Anomala innuba 99 15 4 8.9 Anomala ludoviciana 2 30 20 25.0 Diplotaxis sp. 1 15 15 15.0 Bolbocerosoma bruneri 5 15 10 11.6 Table XII brings out the fact that the average depth of hibernation of all species was below the plow line. In fact, grubs of only two species were found above the 6-inch level in the work at Manhattan, while a few grubs of Phyllophaga lanceolata were taken at three to six inches at Goddard. 147] LARVAL SC ARAB AEOIDEA—B AYES 63 While very few determinations were made of the grubs of Phyllophaga collected at Manhattan, it is interesting to note that all were several inches below the plow line. Grubs of Ochrosidia immaculata predominated in practically all collec- tions. Out of a total of 101 grubs of this species, only 12 were found above the plow line. The remaining 89 were found at depths ranging from 7 to 30 inches, with the majority at the 14-inch level. Of Anomala innuba, which ranked second in number of grubs identified, 99 specimens were taken. This species does not burrow downward to any great extent for hibernation. The average depth for it was 8.9 inches, with extremes of 4 and 15 inches. Anomala ludoviciana, on the other hand, apparently burrows deeply into the soil, as evidenced by the two specimens taken at depths of 20 and 30 inches. Pupation. — In a previous paragraph mention was made of the prepara- tion made by the larva in anticipation of the transformation to the pupal stage. This period of inactivity, known as the prepupal stage or semi- pupal stage, is of short duration before the actual molt to the pupal condi- tion occurs. The period is characterized by internal activity, a cessation of feeding, some body shrinkage, and the cleaning out of the alimentary canal. The final molt having been completed, the pupa has now asumed a condi- tion more nearly like the adult form. When freshly transformed, the body is a creamy white, but as development proceeds many of the adult colors are assumed. Hayes and McColloch (1920) have pointed out that during the later stages of development sexual differences of the adult may be dis- tinguished in the antennae and the genitalia, which may be discerned through the cuticula of the pupa. In many cases, these characters are apparent throughout the last half of the pupal stage. Pupation occurs in late summer and autumn in many species (e.g., Phyllophaga), and transformation to the adult occurs shortly thereafter, to enable the insect to pass the winter in the adult stage. Others (e.g., Anomala) pupate in the late spring and early summer, and at the end of the pupal period, after a few days of inactivity, are ready for flight. In the case of those species transforming in the fall it is usually stated in the literature that the winter is passed by the adult at the place of pupation, usually within the exuvia of the pupa. In observations made by McColloch and the writer there is some reason for believing that many adults leave this place of pupation and burrow beneath the frost line. No systematic study has been made of the pupae of the Lamellicornia and no keys are available for their identification. Causal observation indi- cates that there are many differences, and when enough material becomes available it may be possible to describe their recognition characters. It is noteworthy that most pupae of the Melolonthinae are characterized by a pair of pointed caudal appendages, while Anomala and a number of other 64 ILLINOIS BIOLOGICAL MONOGRAPHS 1148 genera have a rounded, blunt caudal end. Without going into details of the morphology of the pupa, it is sufficient to point out that most of the external characters of the adult are apparent (Fig. 195). The Passalidae and Lucanidae pass the pupal period in the decaying wood in which they develop. The coprophagous Scarabaeidae, as a rule, pupate and remain within the ball or mass of manure in which the larvae have developed, while higher Scarabaeidae may merely lie within the molt- ed larval exuviae in the soil. A number of Scarabaeidae, such as Pelidnota punctata and many Cetoniinae (e.g., Euphoria), construct a cocoon in which the pupal period is undergone. In the case of Pelidnota, the cocoon con- sists of fragments of wood, while in the Cetoniinae (e.g., Euphoria) it is made of bits of manure in which the larva grew. Others may use the soil or root-fibers. These cocoons, made of the food material, are oval in form. The outer surface is rough while the inner walls are smooth and polished. They are built by the larva before the transformation of the pupal stage, and the material which holds the structure together is a glandular secretion ejected from the hind intestine. The larva, by its mouthparts and body movements, is able to mold the material into a compact water-tight cocoon. The length of the pupal stage in the many species varies from a week or more to a month and even longer in cooler weather. Emergence is accom- plished by a splitting of the cuticula along the back. Length of Life-Cycle Melolonthinae The length of the life-cycle in the subfamily Melolonthinae is extremely variable. It has long been known that Melolontha melolontha requires three years for development in France and southern Germany and four years in northern Germany. In Mauritius it has been found that Phytalus smithi Ar. has a life-cycle of slightly over one year (De Charmoy, 1912). Although the species of Phyllophaga have been known as important pests for a number of years, only scanty information has been available concerning their life-histories, particularly with reference to the length of the various stages. This is due, in a large measure, to the fact that practi- cally all of their period of development is spent beneath the surface of the soil, where it is difficult to observe their life activities. Chittenden (1899) was the first to report the rearing of a species of this genus. He found, in the case of one individual of P. fervida, that 781 days were required from the date of egg laying to the transformation of the pupa to the adult. This makes a life-cycle of three years for the species if the winter period of adult hibernation is included. Davis (1916), reporting on the length of the life-cycle of 18 species of the genus, notes in his summary that one species, P. tristis, invariably has a two-year period in the region of Lafayette, Indiana, and eleven other species, namely, P. fervida, P.fusca, P. vehemens, 149] LARVAL SCARAB AEOIDEA— HAYES 65 P. rugosa, P. ilicis, P. grandis, P. fraterna, P. hirticula, P. inversa, P. bipartita, and P. congrua, without exception, have a three-year life-cycle. Two species, P. crenulata and P. crassissima, have a three-year cycle that may be extended to four years, and certain other species, as P. futilis, P. ephilida, and P. implicata, ordinarily have a three-year cycle that is often reduced to two years. From this it can be seen that the more important species have, in the latitude of Lafayette, a three-year cycle. Smyth (1917) has found that the life-cycle of P. vandinei occupied approximately one year in Porto Rico. These citations contain practically all our knowledge of the life-cycle of members of the genus, and except for the work of Smyth very little has been learned concerning the length of the immature stages. In a study of the life-history and development of white grubs carried on by the writer in Kansas, seventeen species of the genus Phyllophaga were reared in varying numbers from the egg to the adult state. The devel- opment of one species, P. lanceolata, was described in 1919 and six others in 1920. In 1925, ten others were described and a comparison of their devel- opment made with those previously reported. To generalize, it can be asserted that the results are in accord with those of Davis in showing a decided variation of the length of the life-cycle in most of the species, this variation being found in the length of the larval period. For example, some species have, in the vicinity of Manhattan, Kansas, either a one-, two-, or three-year life-history. With the exception of four species, P. affabilis, P. submucida, P. longi- tarsa, and P. lanceolata, all of the Phyllophaga reared by the writer pupate in the fall and pass the winter as adults. Accordingly, in considering the life-cycle, eight or nine months should be added to the figures presented in the following table to arrive at the total period of life from the time of oviposition to the normal time of death. In the four exceptions noted above, pupation occurs in the spring or early summer and the adults emerge soon after transformation. To further compare the life-cycle of Phyllophaga, it should be noted that Davis (1916) found a two-year cycle in P. tristis and P. lanceolata; a two- and three-year period for P. burmeisteri, P. futilis, and P. implicata; a three-year period for P. arcuata, P. bipartita, P. congrua, P. fraterna, P. fusca, P. grandis, P. hirticula, P. ilicis, P. inversa, P. rugosa, and P. vehemens; and a three-year and possibly a four-year cycle in P. crassissima and P, crenulata. Smyth (1917), in Porto Rico, found for 14 complete records of P. van- dineiSrayth. an average period of 306 days from egg to adult with a maximum of 395 days and a minimum of 212 days, or expressed in months, from seven to thirteen months with an average of about ten months. Except in the case of P. tristis, this is one of the shortest life-cycles reported. It is somewhat shorter, but is probably comparable to the one-year periods of 66 ILLINOIS BIOLOGICAL MONOGRAPHS [150 P. affabilis and P. longitarsa as here recorded. Criddle (1918) reports that P. nitida, P. drakii, P. anxia, and P. rugosa have in Manitoba, Canada, a four-year life-cycle, but gives no definite data on the length of the various stages. Based on actual rearings by the writer, the summaries of the life-cycles of the 17 species under observation are given in Table XIII. TABLE XIII— SUMMARY OF THE LIFE-CYCLE IN THE GENUS PHYLLOPHAGA Species Number of Individuals Reared Number of Days from Egg to Adult Life-cycle in Years Maximum Minimum Average P. affabilis 4 3 19 13 3 4 1 1 48 2 21 29 23 16 81 2 22 389 815 791 734 838 474 461 424 701 811 836 839 723 494 816 789 790 369 495 429 708 459 137 461 424 327 434 445 447 357 457 449 780 442 376.6 703.0 501.8 720.3 591.3 380.0 461.0 424.0 356.4 628.5 735.7 791.5 675.7 469.7 588.3 784.5 470.4 1 P. tripartita 2 and 3 P. corrosa 2 and 3 P. submucida 2 P. vehemens 2 and 3 P. tristis P. f usca 1 and 2 2 P. crenulata 2 P. longitarsa 2 and 3 P. praetermissa 2 and 3 P. rugosa 2 and 3 P. rubiginosa 2 and 3 P. lanceolata 1 and 2 P. futilis 2 P. crassissima 2 and 3 P. hirticula var. comosa . . . P. implicata 3 2 and 3 Other members of the subfamily are little known. The rose chafer, Macrodactylus subspinosus, has been studied and found to have a one-year life-history. Other important genera, such as Polyphylla, Serica, Diplo- taxis, and Dichelonyx, are practically unknown as far as knowledge of their life-history is concerned. Rutelinae It is evident that there are two distinct types of development in the two tribes of the subfamily Rutelinae. In the tribe Anomalini there appears to be invariably a one-year life-cycle, while in the tribe Rutelini it is known that at least two years and often three years are needed to complete the life-history. In the Anomalini, Hadley (1922) has pointed out that Popillia japonic a requires one year to develop. The writer (1918) has shown that Anomala binotata matures in one season. The larvae require, on an average, 8 3 151] LARVAL SCA RABAEOIDEA —HA YES 67 days to develop, and pupation occurs in the fall. On the contrary, Anomala innuba normally matures in the spring, but only requires one year to com- plete growth. Two instances were noted wherein the larvae of A. innuba pupated in December. Anomala kansana also has a life-cycle quite similar to that of A . innuba. It has also been shown (Hayes, 1921) that Strigoderma arboricola Fab., another Anomalini, has a one-year life-cycle, in which development is completed in from 351 to 358 days. In this case the larvae pass the winter and pupate in the spring. In the tribe Rutelini, Pelidnota punctata requires two years to mature, while Cotalpa lanigera needs either two or three years (Hayes, 1925). A summary of the life-cycle in this sub- family, based mostly on rearings by the writer, is given in Table XIV. TABLE XIV— SUMMARY OF THE LIFE-CYCLE IN RUTELINAE Species Number of Individuals Reared Number of Days from Egg of Adult Life-cycle in Years Maximum Minimum Average Anomalini Anomala binotata — 100 119 — 1 year Anomala innuba — 330 356 342.9 1 year Anomala kansana 16 376 339 368.3 1 year Strigoderma arboricola 4 351 358 — 1 year Popillia japonica* — — — — 1 year Rutelini Pelidnota punctata 1 698 698 698 2 and prob- ably 3 years Cotalpa lanigera 21 416 806 604.6 2 and 3 years * Data on maximum and minimum periods not available. Dynastinae The subfamily Dynastinae is remarkable for the fact that it contains some of the largest coleopterous insects. One has only to recall such genera as Dynastes and Strategus to realize this fact. The group contains a num- ber of species whose depredations on crops make them of considerable economic importance, especially in the southern part of the United States and in the West Indies. One species, Ochrosidia (Cyclocephala) immaculata Burm., is very injurious in the larval stage to the roots of staple crops in the central states; and another, Ligyrus gibbosus DeGeer, known as the car- rot-beetle or muck-worm, is destructive in the adult stage to carrots, sunflowers, and other plants. Both of these have been under observa- tion during the course of this study. Ochrosidia immaculata has been reported under the name of Cyclocephala villosa (Hayes, 1918) and the life-cycle was given as one year. Likewise, Ligyrus gibbosus was found (Hayes, 1917) 68 ILLINOIS BIOLOGICAL MONOGRAPHS [152 to have a one-year life-cycle. Ochrosidia differs from Ligyrus in passing the winter as a larva and maturing in the spring. Smyth (1916) has reported on the period of development of five species representing three genera of this subfamily. He found that Strategus titanus Fab., in Porto Rico, required an average of 338 days to reach maturity, Strategus quadrifoveatus Beauv. required slightly over one year, and Ligyrus tumulosus only 77 days. Two other species, Dyscinetus trachypygus and Dyscinetus barbatus, complete their growth in 104 and 144 days, respectively. Phillips and Fox (1917) report the development of Euetheola rugiceps in about 88 days. Smyth (1920) made further reports on the life-cycle in Strategus. The writer has reared three species of this subfamily, Ligyrodes relictus, Ligyrus gibbosus, and Ochrosidia immaculata. A summary of the chief points of interest in their development is given here. Adults of L. gibbosus are present in the soil throughout the winter and early spring. During the latter part of April or the first few days of May, and continuing throughout the summer, they emerge at night and fly to lights, returning to the soil before daybreak. During the summer of 1916, eggs were plentiful from the last of May to late in July. Larvae were present from June throughout the remainder of the summer and early fall, and pupae from the last of July to the last of October. The length of each stage of development is shown in Table XV. Ligyrodes relictus is to be regarded as a beneficial species, living as it does in manure and rotting haystacks and thus materially hastening the processes of decomposition. Smith (1902) reported the beetles injuring the roots of hardy pyrethrums and the roots of sunflowers, but his statement that the species is smaller than the ordinary June-bug and more roughly sculptured, leads to the suspicion that his determination was incorrect. These two characters and the habit of attacking sunflowers would suggest that Ligyrus gibbosus was the species in question. Smith appears to be the only writer who considers Ligyrodes relictus as an injurious species. There are no citations in the literature treating of the life-history of the species, and scarcely any habits are mentioned except that the beetles and grubs live in decaying vegetable matter. The average periods of development of the different stages have been found to be: for the egg stage, 9.3 days; the active larval period, 46 days; the prepupal period, 4.1 days; and the pupal period, 13.1 days. The total of 72.4 days for complete development thus approximates very closely the full period computed from the length of the different stages. Davis (1916) has reported that the species develops in one year, but gives no data on the length of stages. The beetles appear above ground in April or May for the spring flight, returning to the soil each day, where mating occurs. They disappear for a short time in June 153[ LARVAL SC ARAB AEOIDEA— HAYES 69 and July, and the new brood appears in July and August for a second period of flight. The genus Ochrosidia (Cyclocephala) contains some of our common and most injurious white grubs. Forbes (1891, p. 40) reports the grubs of 0. immaculata infesting grass-land, corn on sod, roots of corn, and young oats. Titus (1905, p. 14) found them at the roots of grass and sugar-cane stubble, and Riley (1870, p. 307) recorded them in strawberry beds. Davis (1916, p. 264) states: "Cyclocephala immaculata is frequently found in compost heaps and in cultivated fields, and may obtain its full growth on decaying matter alone or may become a serious field pest, damaging crops similar to those attacked by Lachnostema grubs." To summarize, the life-cycle of O. immaculata is one year. Adults appear at lights in June, July, and early August. Eggs, which are laid in soil, hatch after 9 to 25 days. The larva passes the winter in hibernation. The larval stage was found to average 347 days. The pupal stage varied in length from 8 to 24 days. A comparison of the life-cycles of these three species with others re- ported in the literature is given in Table XV. In this table it is to be noted that, of the species whose life-history is known, the average period of devel- opment ranges from 72 days, or slightly over two months for Ligyrodes, to 430 days, or more than a year, for Strategus quadrifoveatus. TABLE XV— SUMMARY OF THE LIFE-CYCLE IN DYNASTINAE Species Egg Stage Larval Stage Pupal Stage Egg to Adult Aver- age Max. Min. Aver- age Max. Min. Aver- age Max. Min. Average Ligyrodes relictus 9 11 8 50 — — 13 26 9 72 Ligyrus tumulosus 8 13 — — 55 — — 14 — — 77 Ligyrus gibbosus 10 22 7 59 80 43 19 29 11 88 Ochrosidia immaculata IS 25 13 347 384 335 17 21 15 379 Strategus titanus* 17 21 15 344 — — 23 24 — 338 Strategus quadrifoveatus 8 20 — — 385 — — 27 — — 430 Dyscinetus trachypygus" 12 18 10 81 — — 13 16 12 104 Dyscinetus barbatus" 13 — — 106 — — 15 18 13 144- Euetheola rugiceps b 14 — — 60 — — 14 — — 88 a Data from Smyth (1916). b Data from Phillips and Fox (1917). Arrow (1910, p. 259), in his summary of the habits and metamorphoses of this subfamily, points out the following facts. They are mostly confined to the warmer climates and are of somewhat retiring habits, and our know- ledge of their metamorphoses and modes of life is exceedingly scanty. With but few exceptions they are nocturnal or crepuscular and are not easily 70 ILLINOIS BIOLOGICAL MONOGRAPHS [154 found, and in few cases have their early stages been traced. He points out the increase in the size of the eggs, which is discussed on a previous page, as being characteristic of most scarabaeid eggs. Further comment is made that the larvae "do not differ in any marked degrees from those of the Cetoniinae and allied subfamilies," and like the Cetoniinae feed upon decaying vege- table matter, and sometimes upon living roots or woody tissues. A discus- sion of the nest-building habits of Strategus antaeus, as quoted from Manee (1908), shows these larvae to feed first on leaves stored in the nest and then probably on oak roots. Various Dynastinae are known to feed on the roots of grasses, one is known to destroy the roots of sugar cane, and Oryctes nasicornis is found in the refuse-heaps of tanneries, where the larvae feed on the decomposed bark. Cetoniinae Several members of the subfamily Cetoniinae are injurious to vegeta- tion in the United States. Chief among these are the bumble flower-beetle, Euphoria inda, and the green June-beetle, or fig-eater, Cotinus nitida. As a rule, the mandibles of the adults of this subfamily are poorly developed and are fitted only for the eating of such light foods as pollen and sap. There is very little American literature on the life-history of American species of this group. Euphoria inda is commonly assumed, and probably correctly, to have a one-year life-cycle, and the green June-beetle, Cotinus nitida, has been reported by Chittenden and Fink (1922) to have a one-year period of development. In the present study, a number of species have been under observation, such as Euphoria inda, Trichiotinus piger, Osmo- derma eremicola, and Cremastochelius nitens. The life-cycle of none of these has been completely worked out, but two species, Euphoria fulgida and Euphoria sepulchralis, have been carried through to maturity and the re- sults are here summarized (Hayes, 1925). TABLE XVI— SUMMARY OF THE LIFE-CYCLE IN CETONIINAE Species Number of Days from Egg to Adult Length of Life-cycle Maximum Minimum Average Euphoria fulgida Euphoria sepulchralis Euphoria inda Cotinus nitida a Osmoderma eremicola b 434 123 323 74 381.3 92.9 1 year 1 year 1 year 1 year 3 years a Data from Davis and Luginbill, 1921. b Data from Sweetman and Hatch, 1927. 155] LARVAL SCARABAEOIDEA— HAYES 71 Arrow (1910, p. 24) says of this subfamily, "Of the metamorphosis and habits of the species we know lamentably little." There are, according to him, about 2,500 species in the world. Of those occurring in America but few are of economic significance, and except for the green June-beetle, Cotinis nitida, three species of the genus Euphoria, and one of Osmoderma, nothing has been done toward working out their life-histories. Recently Sweetman and Hatch (1927), in rearing a larva of Osmoderma eremicola for 18 months, concluded that, allowing for outdoor periods of hibernation, the life-cycle would be three years for this species. It is interesting to note that all whose life-cycle has been studied require but one year to develop while Osmoderma with entirely different habits requires a much longer time. The little that is known of the development in this subfamily is summarized in Table XVI. Laparosticti For an interesting account of the biology of the Coprinae and other dung-feeding larvae, the reader is referred to Fabre's The Sacred Beetle and Others (English translation, 1924). Here is to be found this author's study of the development of the sacred beetle (Scarabaeus sacre), the Gymnop- leuri, Copris, Onthophagus, and Geotrupes. Space will not be taken here to quote it. No complete American work has been done on any of our species of the subfamily. Neither has there been any investigation into the length of life of our Aphodinae. The following account of this subfamily is a translation of Schmidt's account in Genera Insectorum, fascicle 110, (1910). The life-history and development of the Aphodinae are little or not well known. It is generally observed that their eggs are laid in dung, from which the developing larvae obtain their nourishment, and that pupation occurs in or under the food material. Many species apparently prefer and seek out the excrement of a definite animal. Certain species develop in the excrement of the sheep or deer, others in that from cattle, some from horses; while others may prefer either that of horses or cattle. Certain European species are known to develop in human wastes. Hubbard has reported A. troglodytes as living in the burrows of the land tortoise, Gopherus polyphemus, and Chapman states that A . porous lays its eggs in the brood chambers of the scarabaeid larvae, Geotrupes, or the material upon which the Geotrupes feed. Another species occurs only in rabbit dung. Psammobius and Rhyssemus inhabit sandy regions. Their habits have not been studied, but it is thought that some species of Rhyssemus live on vegetable materials while others live in dung. Ohaus has noted some species of Ataenius in dung and others under the bark in rotten wood of fallen trees. Some exotic genera of this subfamily (Euparia and Friedenreichi) live with ants, while others (Chaetopisthes, Corytho- derus, and Termitodius) inhabit termite dwellings. 72 ILLINOIS BIOLOGICAL MONOGRAPHS [156 TAXONOMY Early students of American insects confined themselves to a study of the anatomy and classification of adult insects and the working out of their life-histories. This has resulted in descriptions of numerous new genera and species from adult characters and the description of the eggs, larval and pupal stages, and the host plants of numerous species. The number of known life-histories has been greatly increased by the establishment of state and federal experiment stations and the appointment of entomologists for the study of injurious and related species. There has been a great increase in the number of persons interested in the study of immature insects; and there have been published, not only in Europe but also in America, many systematic studies attempting to furnish tables for the identification of immature as well as adult insects. All classification is based on anatomical characters and any attempt at identification must be preceded by morphological studies. The anatomy of immature insects in many cases, while similar in general to that of the adults, is frequently very different, especially when the insects are examined in detail. The larvae contain many structures peculiar to themselves, being in most cases fitted to live a life very different from that of the adults. A study of the anatomy of immature insects is of value from three points of view: ontogeny, morphology, and classification. It has been shown repeatedly that the complete history of the individual — its ontogeny — throws much light upon the relationship of organisms and upon their differentiation, not only as to species and genera but also as to families and orders. Investigators have found that what had been considered as a single species, from a study of the adult alone, proved to be a complex of two or more species when the immature stages were known, and that characters previously believed to be worthless could be used for separating the adults. Ontogenetic and morphological studies should proceed hand in hand. Many structures that are complex and difficult to interpret in the adult are easily understood by a comparison with the conditions found in immature insects. By comparative studies of this sort, the homology and homodynamy of various structural parts are easily determined. While it is impossible, without at least some morphological knowledge, to attempt the identifica- tion of adult specimens, such knowledge is equally pertinent for one under- taking the study of the anatomy of immature insects. The classification and identification of such insects is of the greatest value to the economic 157] LARVAL SCARABAEOIDEA— HAYES 73 worker, because the investigator in this field almost invariably meets with the stages that are doing the damage — the nymphs or larvae — and, unless it is a species with which he is familiar, it would be impossible to identify the pest, if no analytical tables were available, until the adult has been bred. It is hoped that by a careful study of the morphology and classification of immature insects, the labor of identification can be lessened. With these facts in mind, the foregoing morphological studies have been made for the purpose of producing analytical keys to the various groups of white grubs. The author realizes their incompleteness. This is due, in great part, to the fact that many species are yet unknown, that in many instances only one or two specimens of known species are available and no extended consideration of the problem of variation can be made at this time because of the present state of our knowledge of the group. The following keys are therefore submitted as a preliminary step toward the further progress of the work. In the key to genera a few known species have been included. This key appeared in a recent paper by the writer (1928) and is here included with corrections and additions. It is gratifying to note, in view of the previously published statements that larvae of these families could not be separated, that one of Prof. J. W. McColloch's student's reports that this key, as it first appeared, has been helpful in his work with white grubs. The key to species of the genus Phyllophaga contains less than one-third of the known North American species of this genus, but until further progress is made with biological studies of the group little advance can be expected in our acquaintanceship with the many species now un- known in the larval stage. LARVAL KEY TO FAMILIES OF THE SUPER- FAMILY SCARABAEOIDEA 1. Posterior pair of legs small, undeveloped (Fig. 137) ; coxae modified into scraping, stridulat" ing apparatus; antennae three-segmented (Fig. 94); body segments not distinctly divided into annulets and nearly devoid of spines and setae (Fig. 3) . .Passalidae, genus Passalus 1. Posterior pair of legs normally developed, legs may or may not be used for stridulation (Fig. 135, 136) ; antennae four or five-segmented; body segments usually distinctly divided into annulets and more or less covered with spines and setae 2 2. Anal segment not trilobed on cadual aspect, legs not modified for stridulation except in Geotrupes (Figs. 1,12) Scaeabaeidae 2. Anal segment trilobed on caudal aspect (Fig. 151, 154); meso- and metathoracic legs modi- fied for stridation (Fig. 135, 136) 3 3. Labrum usually biemarginate on its distal magin, trilobed (Fig. 33, 36); emargination of peritreme of anterior spiracles on caudal margin as is the case with the remaining spi- racles; anal segment strongly trilobed; larvae feed in wood (Fig. 2) Lucantdae 3. Labrum not biemarginate on its distal margin, more rounded, not trilobed on distal mar- gin (Fig. 25); spiracular peritreme of all segments small, open on dorsal margin, poorly defined (Fig. 123); anal segment feebly trilobed (Fig. 151); larvae feed in carrion (Fig. 7) Trogidae, genus Trox 74 ILLINOIS BIOLOGICAL MONOGRAPHS [158 LARVAL KEY TO THE SUBFAMILIES OF SCARABAEIDAE 1. Galea and lacinia of maxilla not fused (Fig. 98, 100), that is, the mala is deeply bifed; usually coprophagous larvae Laparosticti 2 1. Galea and lacinia of maxilla fused to form the mala (Fig. 97, 104); mostly "leaf chafers" i'LEUROSTICTI 6 2. Tarsi without claws (Fig. 1), a distal seta may be present; abdomen strongly "humped" on the dorsum (Fig. 1) ; labrum trilobed and biemarginate (Fig. 64) 3 2. Tarsi with claws or the tarsus is bilobed on its distal end, abdomen not "humped" on the dorsum 5 3. Antenna four-segmented Coprinae, genus Copris 3. Antenna five-segmented 4 4. Tarsus strongly rounded or blunt on its distal end; tormae of labrum (Fig. 55, t) not meeting on the median line; ental setae of lateral lobes of the epipharynx (Fig. 55, 11) numerous (more than eight) Coprinae, genus Canthon 4. Tarsus terminating in a long seta; tormae of labrum meeting on the median line (Fig. 67. t); ental setae of lateral lobes of the epipharynx scarce (usual one on each lobe) Coprinae, Onthophagus (Fig. 4) 5. Tarsus without claws, bilobed on its distal end; posterior pair of legs considerably shortened (not as much so as in Passalidae) ; second and third pairs of legs modified for stridulation Geotrupinae, genus Geolrupes 5. Tarsus with claws which are longer than trasus; legs not modified for stridulation; third pair of legs not noticeably shortened; body densely setaceous (Fig. 139); anal slit transverse (Fig. 139) Glaphyrinae, genus Amphicoma 6. Mandibles on their caudal aspect without an oval, stridulating area made up of transverse striae (Fig. 84) ; radula usually with two longitudinal rows of mesad pointing spines (not present in Serica) (Fig. 156 to 189); anal slit in the form of an obtuse angle (Fig. 156 to 189) Melolonthinae 6. Mandibles on their caudal aspect, with a distinct oval, stridulating area made up of trans- verse striae (Fig. 81, sa); with or without two longitudinal rows of mesad pointing spines on radula; anal slit not angulate (Fig. 138) and more transverse 7 7. Labrum symmetrical (Stephanucha is not symmetrical but it is a rare species), usually tri- lobed (except in Trichiotinus which is usually recognized by the presence of ocelli): epipharynx with a conspicuous, curved row of small spines in the region of the distal sensory area (Fig. 57, st) ; dorsum of abdomen behind the last spiracle-bearing segment not divided transversely by an impressed line thus appearing as one segment (Fig. 11) ; some species are "black crawlers," others found in soil, wood, and manure Cetoniinae 7. Labrum asymmetrical, not trilobed; epipharynx without a conspicuous, curved row of small spines in the region of the distal sensory area (Fig. 42) ; dorsum of abdomen behind the last spiracle-bearing segment divided transversely by an impressed line making it appear as two segments (Fig. 10) ; species of varied habits ■ 8 8. Ental aspect of the labrum with a series of transverse striae on the lateral margins at bases of lateral setae (Fig. 40, st) Tribe Anomalini Rutelinae 8. Ental aspect of the labrum without a series of transverse striae on the lateral margins at bases of lateral setae (Fig. 50) 9 9. Stridulating teeth of maxillae (Fig. 106, ms) sharply pointed and curved, apices directed distally (Fig. 1 14) ; distal segment of maxillary palpus usually without a distinct, setaceous sensory area Tribe Rutelini Rutelinae 9. Stridulating teeth of maxilla not pointed but strongly truncate being as broad as long, not curved, and not directed distally (Fig. 124) ; distal tooth of the series of stridulating maxil- lary teeth twice as wide as the others in the series; distal segment of the maxillary palpus usually ending in a setaceous sensory area Dynastinae 159] LARVAL SCARABAEOIDEA— HAYES 75 LARVAL KEY TO GENERA OF SUBFAMILY MELOLONTHINAE 1. Anal slit obtusely angulate, not trilobed (Fig. 156); tarsal claws of posterior legs less than half as long as claws of the other legs; distal sensory area of epipharynx usually with seven to eight strong spines (Fig. 25). A few species of Phyllophaga, e.g., lanceolata (Fig. 26), have less than seven spines but can readily be distinguished by the lateral striae of the epipharynx 2 1. Anal slit more acutely angulate, faintly trilobed (Fig. 140); tarsal claws of posterior legs more nearly equal in length to the claws of the other legs, never less than half as long; distal sensory area of epipharynx never with more than four strong spines (Fig. 31, sp) . .3 2. Longitudinal double row of spines of radula short, scarcely more than eight spines to a row (Fig. 141); head dark brown in color; dorsum of abdominal segments very densely spinose; striations of lateral margins of epipharynx indistinct (Fig. 25) ; epipharynx never with a submarginal, distal row of striae Polyphylla 2. Longitudinal double row of spines of radula longer, usually more than ten spines to a row (Fig. 156); head light yellow in color; dorsum of abdominal segments less densely spinose; striations of lateral margins of epipharynx distinct (Fig. 26, si), epipharynx with a sub- marginal, distal row of striae, sometimes difficult to observe in some species (Fig. 26, sms) Phyllophaga 3. Epipharynx with three strong spines in distal sensory area (Fig. 31); radula with a con- spicuous transverse row of spines (Fig. 140) Serica 3. Epipharynx with four strong spines in distal sensory area (Fig. 34) 4 4. Setae of radula not hooked at the tip; presence or absence of double longitudinal rows of spines on radula questionable; 1 Distal end of abdomen sparsely clothed with shorter, stronger setae; claws of posterior tarsi less than half as long as those of other tarsi. . . . Diplotaxis 4. Setae of radula hooked at the tip (Fig. 142) ; radula with a short, double row of longitudinal spines; distal end of abdomen densely clothed with long delicate setae; anal opening sharply acute; claws of posterior tarsi equal in length to claws of other tarsi Macrodactylus LARVAL KEY TO GENERA OF SUBFAMILY RUTELINAE 1. Ental aspect of the labrum with a series of transverse striae on the lateral margins near bases of lateral setae (Fig. 41, st.) Tribe Anomalini 2 1. Labrum without such transverse striae on its ental lateral margins (Fig. 44) Tribe Rutelini 4 2. With two of the four spines of the distal sensory area of the epipharynx strongly fused at the base making a large spine with its distal end bifed (Fig. 52, sp) ; longitudinal, double row of spines of radula parallel, not divergent, with eight spines in the left row and nine in the right row Strigoderma arboricola 2. Without two spines of the distal sensory area of the epipharynx fused at base to form a bifed spine; longitudinal double row of spines of the radula either parallel or diverging posteriorly 3 3. Radula with longitudinal rows of spines short, about seven spines in each row; rows strongly divergent posteriorly; sensilhae of distal sensory area of epipharynx at bases of distal spines about equal in size and arranged nearly semicircularly (Fig. 41) Popillia japonica 3. Radula with longitudinal rows of spines usually longer, with twelve to thirteen spines in each row (Fig. 143); rows more or less parallel and frequently converging posteriorly; 1 The only specimen of "Diplotaxis" available does not show the radula distinctly. A specimen in the Illinois Natural History Survey Collection labeled "Diplotaxis" is apparently a species of Serica. 76 ILLINOIS BIOLOGICAL MONOGRAPHS [160 sensilliae of distal sensory area of epipharynx at bases of distal spines unequal in size and not arranged as definitely in a semicircle (Fig. 40, 43, 46) Anomala 4. Larvae found in rotten logs or stumps, sometimes under dried manure, Labrum wider than long; -with or without (Fig. 144) two longitudinal rows of spines on radula; setae of lateral margins of labrum of various lengths but not strongly curved (Fig. 44, 50) 5 4. Larvae usually formed in sandy soils, labrum (Fig. 50) as long as wide, strongly rounded but asymmetrical; without longitudinal rows of spines on radula; lateral margins of labrum with strongly curved, flattened setae which increase in length distally Cotalpa lanigera 5. Larvae found in rotten logs or stumps, without two longitudinal rows of spines on radula (Fig. 144) ; distal sensory area not produced into a single large, chitinous tubercle, without semicircular ring at base (Fig. 44) Pelidnota punctata 5. Larvae under rotten logs or stumps, or under dried manure on sandy soils; with two longitudinal rows of spines on radula; distal sensory area of epipharynx produced into a single large chitinous tubercle, having between its base and the distal margin a narrow chitinous semicircle Polymoechus brevipes LARVAL KEY TO GENERA OF SUBFAMILY DYNASTINAE 1. With a single ocellar spot at base of antenna 2 1. Without an ocellar spot at base of antenna 4 2. Radula with a longitudinal cleared area surrounded by recumbent spines similar to other spines of radula, spines not strongly differentiated as in other subfamilies having the double longitudinal row of spines; usually found in manure. .Ligyrodes {Ligyrus) relidus 2. Radula without such a longitudinal, non-setose area formed by the absence of spines or setae 3 3. Distal sensory area of epipharynx produced into a long, proximal pointing, chitinous process which is curved at its apex, at its base are numerous large setae (Fig. 48, dsa); head brownish-tan in color; usually found in burrows near manure Strategus 3. Distal sensory area of epipharynx not produced into a long, proximal pointing, chitinous process nearly devoid of setae in the region of the distal sensory area (Fig. 151); head nearly black in color, densely punctate; usually found on soil among dead leaves in woods Xyloryctes 4. Epipharynx with the chitinous portion of the distal sensory area produced to form a single broad tubercle (fig. 45) ; labrum strongly asymmetrical; sides of labrum strongly rounded; setae of radula very short, not hooked (Fig. 145) Ligyrus gibbosus 4. Epipharynx with the chitinous portion of the distal sensory area produced to form two tubercles or spines (Fig. 53, 54), labrum more nearly symmetrical; sides of labrum less rounded 5 5. Distal sensory area of the epipharynx produced in the form of two broad tubercles close to the distal margin of the labrum (Fig. 54) ; head and mandibles almost black in color; head strongly punctured; prothorax with a large, brown more heavily chitinized area on the sides which is deeply bipunctate ; usually extremely large grubs Dynastes tityrus 5. Distal sensory area of the epipharynx produced in the form of two small spines and more remote from the distal margin of the labrum (Fig. 53) ; head yellow in color; sparsely and finely punctate; prothorax without brown, chitinous areas on sides; never extremely large grubs Ochrosidia (Cydocephala) immaculata* 5 Dyscinetus has not been studied. Davis (1916) states that D. Irachypygus has "a dark brown head which is inconspicuously reticulate and covered with irregularly placed fine punctures, in this respect differing from all species (which he mentions) except Strategus, the head of which is much more coarsely punctate and the species is much larger. The ventral surface of the anal segment bears a patch of hooked spines and the upper surface of the same segment is covered, excepting along the median line, with fine hairs, those at the tip being shorter, stouter and more spine-like." 161] LARVAL SCARAB AEOIDEA— HAYES 77 LARVAL KEY TO GENERA OF SUBFAMILY CETONIINAE 1. Labrum almost as long as wide, not distinctly trilobed (Fig. 66); emarginations of labrum relatively shallow; a distinct ocellar spot at base of each antenna; distal sensory area of epipharynx produced into a conspicuous chitinous tubercle; larvae live in wood Trichiotinus 1. Labrum considerably wider than long, distinctly trilobed, relatively deeply emarginate (Fig. 62) ; no ocellar spot at base of antenna; distal sensory area not produced into a chit- inous tubercle; larvae live in various situations, one genus, (Osmoderma,) in wood. . .2 2. Epipharynx with a chitinous, semicircular carina near distal margin of the median lobe (Fig. 65) ; proximad of this ridge is a semicircle of about sixteen sensory pores. 3 On the side of the epipharynx additional pores apparently make the semicircle continuous almost to the clypeo-labral suture; tarsal claws normal, curved and sharply pointed; larvae live in ant nests Cremaslocheilus 2. Epipharynx without a chitinous, semicircular carina on distal margin of median lobe; the median lobe is provided with a conspicuous semicircle of spines (Fig. 68) ; tarsal claws usually modified into blunt, cylindrical, setaceous tubercles 4 3 3. Epipharynx with about ten spines in a semicircular row in distal sensory area; placed somewhat obliquely (Fig. 68); no chitinous sensory tubercle in proximal sensory area near clypeo-labral suture mesad of tormae; larva live in sandy soil (rare) . .Stephanucha 3. Epipharynx usually with more than ten spines (about 15-17) in a semi-circular row in distal sensory area; placed almost transversely; with a well-defined sensory tubercle in proximal sensory area near cylpeo-labral suture (Fig. 57, 59); larvae crawl on their backs 4 4. Radula of last abdominal segment without a longitudinal, double row of mesad pointing spines; larvae five in wood Osmoderma eremicola 4. Radula of last abdominal segment with a longitudinal double row of mesad pointing spines; larvae five in manure or soil that is rich in decaying organic matter 5 5. Radula of last abdominal segment (Fig. 150) with spines of the longitudinal rows short, separated from each other by a distance nearly equal to the length of the spines; apices of opposing spines distant from each other by less than the length of an individual spine; spines about twice as long as their width at base; antepenultimate antennal segment longer than the terminal segment; usually found in rich, sandy or loam soil Cotinis 5. Radula of last abdominal segment (Fig. 148, 149) with spines of longitudinal rows longer, separated from each other by a distance much less than the length of the spines; apices of opposing spines separated from each other by a distance greater than the length of an individual spine; spines considerably more than twice as long as width at base; antepenul- timate antennal segment not longer than the terminal segment, larvae usually found in manure 6 6. Radula of last abdominal segment with spines of longitudinal rows diverging posteriorly (Fig. 149) Euphoria sepulchralis 6. Radula of last abdominal segment with spines of longitudinal rows converging posteriorly (Fig. 148) Euphoria inda and Euphoria fulgida LARVAL KEY TO GENERA OF FAMILY LUCANIDAE 1. Antennae five-segmented 2 1. Antennae four-segmented 3 2. Dorsal lobe of the three anal lobes acutely pointed on its ventral margin, lateral lobes with * In the only specimen available for study the epipharynx is almost devoid of setae but in their place numerous pores are present which may be trichopores whose setae have been lost, or they may, in fact, be sensillia. 4 Claws of Stephanucha have not been examined. 78 ILLINOIS BIOLOGICAL MONOGRAPHS [162 a concentric oval line (Fig. 154) Dorcus 2. Dorsal lobe of the three anal lobes with its ventral margin strongly rounded; lateral lobes without a concentric oval line Lucanus 3. Caudal region of radula of the last ventral segment with spinose setae; anal opening on each side Platycerus 5 3. Caudal region of radula without spinose setae (Fig. 153) 4 4. Lateral lobes of anus large, subtriangular, not emarginate at their ventral point of union; superior lobe large Ceruchus 4. Lateral lobes of anus plainly eliptical, emarginate at their ventral point of union ; superior lobe small (Fig. 153) Sinodendron ARTIFICIAL KEY TO SOME KNOWN THIRD INSTAR LARVAE OF THE GENUS PHYLLOPHAGA 1. Longitudinal rows of radula with less than five spines (usually four in right and three in left row) ; the tips of opposing spines greatly overlapping, extending almost to the bases of the opposing spines (Southwestern species) (Fig. 157) cribrosa (Lee.) 1. Longitudinal rows of radula with more than five spines in each row; the tips of opposing spines never greatly overlapping (excepting longitarsa which has 8 to 12 spines) 2 2. Each longitudinal row of radular spines composed of a series of spines varying from two rows at the anterior end to three and four rows posteriorly, these compound rows and space between them being rather strongly divergent posteriorly; cephalic spines usually shorter than the caudal spines (Southwestern species) (Fig. 158) farcta (Lee.) 2. Each longitudinal row of radular spines composed of a single series of spines, not composed of several rows as in far da, may or may not diverge posteriorly ; spines of various lengths . 3 3. Longitudinal rows with never more than 16 spines in each row, usually less but not fewer than five 4 3. Longitudinal rows with more than 16 spines in a row 10 4. Two rows of longitudinal spines arranged in the form of a distinct oval; with 12-13 spines in each row; spines short, scarcely as long as the distance separating the bases of ad- joining spines (Fig. 188) tristis (Fab) 4. Two rows of longitudinal spines arranged in nearly parallel rows; with from 5-16 spines in each row; spines variable in length and distance apart 5 5. Majority of spines in each row separated from each other at base by a distance that is less than the length of the individual spines; each row of spines strongly irregular; with 8 to 16 spines in each row 6 5. Majority of spines in each row separated from each other at base by a distance equal to, or greater than the length of the individual spines; each row of spines more nearly regular or parallel; number of spines in each row never more than 14 7 6. Spines 14 to 16 in a row; apices of opposing spines separated by a distance equal to or slightly less than the length of the individual spines; the majority of spines directed cephalo-mesad (Fig. 163) gracilis (Burm.) 6. Spines 8-12 in a row; apices of opposing spines extending beyond the meson and over- lapping or crossing each other; the majority of spines directed caudo-mesad (Fig. 162) longitarsa (Say) 7. Most of the apices of opposing spines separated from each other by a distance equal to or greater than the length of the individual spines; 11 to 13 spines in each row (Fig. 180) implicita (Horn) 7. Most of the apices of opposing spines separated from each other by a distance less than the length of the individual spines 8 6 Genus not seen. The characters given above are from Perris (1877) after Mulsant. 163] LARVAL SCARAB AEOIDEA— HAYES 79 8. Most of the apices of opposing spines reaching the meson and very narrowly separated from each other; 12-13 spines in each row (Fig. 171) vehemens (Horn) 8. Most of the apices of opposing spines not reaching the meson with the opposing apices more widely separated from each other; 13-14 spines in each row 9 9. Spines shorter, much less than half the length of the distance separating the majority of adjacent spines at their base; 13 spines in each row (Fig. 156) lanceolata (Say) 9. Spines longer, usually greater than half the length of the distance separating the majority of adjacent spines at their base; 14 spines in each row (Fig. 175) drakei (Kby.) 10. Spines of radular rows not stout; individual spines usually equal in length or shorter than the intervening distance between the bases of adjacent spines; the bases of the spines much narrower than the interval between the spinep 11 10. Spines of radular rows usually stout; individual spines in most instances much longer than the intervening distance between the bases of adjacent spines; in many species the bases of the spines are wider than the interval between the adjacent spines 18 11. Apices of most of the opposing spines meeting, or slightly overlapping each other, on the meson; 19-22 spines in each row; a conspicuous transverse row of setae between the radular spines and the anal opening (Fig. 161) ephilida (Say) 11. Apices of most of the opposing spines not meeting on the meson, the tips of opposing spines being separated from each other by a distance equal to or greater than the length of the individual spines; 19-29 spines in each row; no conspicuous transverse row of setae between the radular spines and the anal opening 12 12. Spines of radula arranged in nearly parallel rows; 29 spines in left row, 25 in right row; spines short, scarcely longer than their width at base (Fig. 173) horni (Smith) 12. Spines of radular rows arranged in more irregularly parallel rows, no rows with more than 27 spines; most spines considerably longer than their width at base 13 13. Longitudinal rows with from 25 to 27 spines 14 13. Longitudinal rows with from 17 to 21 spines 15 14. Rows of spines very irregularly parallel: with 25 spines in right row and 27 in left row; spines about equal in length to the distance between adjacent bases (Fig. 189) inversa (Horn) 14. Rows of spines more regularly parallel; with 26 spines in each row; spines usually shorter than the distance between adjacent bases (Fig. 160) latifrons (Lee) 15. Radular rows of spines diverging anteriorly 16 15. Radular rows of spines not diverging anteriorly 17 16. Radular rows of spines approximate at caudal ends (distance between opposing apices of caudal spines less than the length of the spines) ; 19 to 20 spines in each row (Fig. 183) delata (Horn) 16. Radular rows of spines less approximate at caudal ends of rows (distance between opposing apices of caudal spines at least equal to or greater than the length of the spines); 17 to 19 spines in each row (Fig. 172) fusca (Froel.) 17. With 19 to 21 spines in each longitudinal row; the two rows constricted at middle 6 (Fig. 176) marginalis (Lee.) 17. With 18 spines in each longitudinal row; the two rows not constricted at middle (Fig. 174) fervida (Fab.) 18. Spines of radular rows short, scarcely longer than the intervening spaces at their bases; the rows of spines strongly diverging caudally then converging near their caudal ends. . 19 18. Spines of radular rows considerably longer than the intervening spaces at their bases; not strongly divergent but in a few species becoming gradually curved near caudal ends 20 9 In the single third instar of this species studied this constriction appears but this may be an abnormality since several second instar grubs available do not show this constriction. 80 ILLINOIS BIOLOGICAL MONOGRAPHS [164 19. Spines of radular rows strongly divergent before the middle; rows not meeting at their anterior and posterior ends; with 23 spines in each row (Fig. 186) vetula (Horn) 19. Spines of radular rows becoming suddenly strongly divergent at or behind middle; rows meeting at anterior end; left row with a conspicuous interval devoid of spines; with 27 and 28 spines in the rows (Fig. 167) calceata (Lee.) 20. Rows nearly parallel; apices of most opposing spines separated by a distance less than the length of the spines (Fig. 164) futilis (Lee.) 20. Rows more or less parallel or gradually curving; apices of the majority of opposing spines separated by a distance equal to or greater than the length of the spines 21 21. Rows short, occupying less than half the distance between the anal opening and the anterior margin of the segment; spines regularly placed and rows slightly curving near caudal end; 20 to 22 spines per row (Fig. 187) affabilis (Horn) 21. Rows longer, occupying one-half or more of the distance between the anal opening and the anterior margin of the segment; spines of varying arrangement; usually, but not always, with more than 22 spines per row 22 22. Spines short, not more than twice as long as width at base 23 22. Majority of spines considerably more than twice as long as width at base 25 23. Spines of uniform size and length, rows more regularly spaced and gradually curving out- ward at center and approaching one another at each end; with 24 spines in one row and 29 in the other (Fig. 177) fraterna Harris 23. Spines not of uniform size and length, rows more irregularly spaced and not gradually curving outward; with not less than 27 spines in any row 24 24. Rows of spines nearly parallel, but with rather jagged rows not noticeably approaching each other at the ends; with 27 in one row and 29 in another (Fig. 165) . .prunina (Lee.) 24. Rows of spines not parallel, slightly curving and approaching each other at caudal end; with 33 spines in one row and 32 in another (Fig. 184) ilicis (Knoch) 25. Rows strongly curved anteriorly to form a rounded cephalic end composed of smaller spines, one row with 24 spines, the other with 28 spines leaving two conspicuous spines at caudal end with no spines opposing them in the other row (Fig. 185) crenulata (Froel.) 25. Rows not conspicuously rounded at anterior end; without unopposed spines at caudal end of one row 26 26. Rows of spines longer, with 32 spines in one row and 31 in the other (Fig. 179) profunda (Blanch.) 26. Rows of spines shorter, with less than 30 spines in any row 27 27. Spines irregularly placed, forming jagged rows 28 27. Spines regularly placed, forming more even rows 30 28. Three caudal spines of each row not more than half as long as the majority of the spines cephalad of them: with 25 spines in one row and 24 in the other (Fig. 181).. balia (Say) 28. Three caudal spines not differing greatly in length from the cephalic ones 29 29. With 27 spines in each row (Fig. 166) congrua (Lee.) 29. With 25 spines in one row and 26 in the other (Fig. 168) crassissima (Blanch.) 30. With both rows of spines bent or constricted mesally near the middle of the rows, 29 spines in each row (Fig. 182) hirticula (Knoch.) 30. With not more than one, or none, of the rows constricted near the middle of the row; not more than 26 spines in each row 31 31. Apices of most of the opposing spines widely separated, being nearly twice the length of the spines apart 32 31. Apices of most of the opposing spines not as widely separated, being considerably less than twice the length of the spines apart 53 32. With the right row of spines constricted or bent near its middle; 28 spines in one row and 24 in the other (Fig. 178) corrosa (Lee.) 165] LARVAL SCARABAEOIDEA— HAYES 81 32. With neither row of spines constricted or bent near the middle; 23 spines in one row and 20 in the other (Fig. 159) torta (lee.) 33. Rows of spines rather regularly curved throughout; with 22 spines in one row and 25 in the other (Fig. 170) tnicans (Knoch) 33. Rows of spines somewhat bulging behind the center; with 24 spines in one row and 26 in the other (Fig. 169) tripartita (Horn) SUMMARY The foregoing study of the larvae of North American Lamellicornia, including the now recognized families — Scarabaeidae, Lucanidae, Trogidae, and Passalidae — attempts to bring together our knowledge of their biol- ogy, including the writer's life-history studies, and presents keys for their identification based on morphological studies. No comparative studies of the structural characters of these insects have hitherto been attempted, and it is hoped that this work, though far from being complete, will afford a stepping-stone to further progress in our knowledge of the group. For taxonomic purposes the characters of the mouthparts and the last abdominal segment have proved the most useful. The analytical keys can be considered only preliminary, inasmuch as a great many of our species are still unknown in the larval stage. The long life-cycle in many species makes rearing very difficult. In the discussion given to biology, there have been brought together, in a comparative way, the more general facts concerning postembryonic development. Some consideration is given to the late embryonic processes, and larval development is considered in a general way, as is also pupal development. This is followed by more-detailed life-history studies in the sub-families Melolonthinae, Rutelinae, Dynastinae, Cetoniinae, and the coprophagous species of the family Scarabaeidae. 82 ILLINOIS BIOLOGICAL MONOGRAPHS [166 BIBLIOGRAPHY Arrow, G. G. 1910. Coleoptera Lamellicornia (Cetoniinae and Dynastinae). The Fauna of British India including Ceylon and Burma. 322 pp., 2 pi. Taylor and Francis, London. Boas, J. E. V. 1893. Ueber die Stigmen der Melolonthalarve. Zool. Anz. Jahrg., 16 (No. 431) : 389-391. Boving, A. G. 1921. The larva of Popilliajaponica Newman and a closely related undetermined ruteline larva. A systematic and morphological study. Proc. Ent. Soc. Wash., 23: 51-62. 2 pi. BtTRMEISTER, H. C. C. 1842. Handbuch der Entomology, 3: 550-68. Berlin, Reimer, 1832-1855. Chapuis, M. F., and Candeze, M. E. 1855. Catalogue des larves des coleopteres connues jusqua ce jour avec la description de plusieurs especes nouvelles. Mem. Soc. Scien., Liege, 8: 341-653 [also ap- peared as a separate] Carpenter, G. H. 1912. Mouthparts of some beetle larvae. Quart. Jour. Micr. Soc, 57: 373-376. Chittenden, F. H. 1899. Some insects injurious to garden and orchard crops. U. S. D. A. Bur. Ent. Bui. 19: 1-77. Chittenden, F. H., and Fink, D. E. 1922. The green June beetle. U. S. D. A. Bui. 891: 1-52. 7 fig., 10 pi. Criddle, N. 1918. The habits and control of white grubs in Manitoba. Can. Agr. Gaz., 5: 449-454. Davis, J. J. 1913. The life-cycle of Lachnosterna tristis Fabr. Jour. Econ. Ent., 6: 276-278. 1915. Cages and methods of studying underground insects. Jour. Econ. Ent., 8: 135-139. 1916. A progress report on white grub investigations. Jour. Econ. Ent., 9: 261-281. 1918. Common white grubs. U. S. D. A. Farm. Bui. 940: 1-28. 21 figi Davis, J. J., and Luginbill, P. 1921. The green June beetle or fig eater. N. C. Agr. Exp. Sta. Bui. 242: 1-35. 9 fig. De Charmoy, D. C. 1912. Rapport sur Phytalis smilhi (Arrow) et autres scarabies s'attacquant a la canne a sucre a Maurice, 355 pp., 10 pi. Port Louis, [not seen] DeHaan, W. 1836. Memoires sur les metamorphoses des coleopteres. Nouv. Ann. Mus. Nat. Hist., 4: 125 [also appeared as a separate] Erichson, W. F. 1848. Naturgeschichte der Insecten Deutschlands, Erste Abt., Coleop., Vol. 3. 968 pp. Nicolai, Berlin. Fabre, J. H. 1924. The sacred beetles and others. Translated by A. T. de Mattos. 425 pp. Dodd, Mead, and Co., New York. Forbes, S. A. 1891. On the common white grubs (Lachnosterna and Cyclocephala) . 111. Ent. Rep., 17: 30-53. 1894. A monograph of insect injuries to Indian corn. 111. Ent. Rep., 18: 1-171. 15 pi. 167] LARVAL SCARABAEOIDEA— HAYES 83 Grandi, G. 1925. Contribute alia conoscenza biologica e morphologica di alcuni Lamellicorni fillo- fagi. Boll. Lab. Zool. Gen'l. e Agraria, Portici, 18: 159-224. Gravely, F. H. 1915. Notes on the habits of Indian insects, myriapods and arachnids. Ind. Museum Record, 11: 483-539. 1916. Some lignicolous beetle-larvae from India and Borneo. Ind. Museum Record, 12: 137-175. Hadley, C. H. 1922. The Japanese beetle. N. J. Dept. Agr., Bur. Stat, and Insp., Circ. 45: 1-20. Hayes, W. P. 1917. Studies on the life history of Ligyrus gibbosus DeG. (Coleoptera) . Jour. Econ. Ent., 10: 253-261. 1918. Studies on the life history of two Kansas Scarabaeidae. Jour. Econ. Ent., 11: 136-144. 1919. The life-cycle of Lachnosterna lanceolata Say. Jour. Econ. Ent., 12: 109-117. 1920. The life histories of some Kansas Lachnosterna. Jour. Econ. Ent., 13: 303-318. 1921. Strigoderma arboricola Fab. — Its life-cycle (Scarab., Coleop.). Can. Ent., 53: 121-124. 1924. The biology of Anomala kansana. Jour. Econ. Ent., 17: 589-594. 1925. A comparative study of the history of certain phytophagous scarabaeid beetles. Kans. Agr. Exp. Sta., Tech. Bui. No. 16. 146 pp., 10 pi. 1927. The immature stages and larval anatomy of Anomala kansana H. and McC. (Scarab., Coleop.). Ann. Ent. Soc. Amer., 20: 193-203. 3 pi. 1928. The epipharynx of lamellicorn larvae (Coleop.) with a key to common genera. Ann. Ent. Soc. Amer., 21: 282-306. 3 pi. Hayes, W. P., and McColloch, J. W. 1920. Some observations on the genitalia of Lachnosterna. Ann. Ent. Soc. Amer., 13: 75-80. 1 pi. 1928. Ecological studies of Kansas scarabaeid larvae (Coleop.). Jour. Econ. Ent., 21: 249-260. Lamark, J. B. P. De. 1817. Histoire naturelle des animaux san vertebres. Tome IV. 587 pp. Verdiere, Paris. Latreille, P. A. 1817. (Insects). Cuvier, Regne Animal 1, 782 pp. Paris. Leng, C. W. 1920. Catalogue of Coleoptera of America North of Mexico. 470 pp. J. D. Sherman, Mount Vernon, N. Y. Leng, W. C, and Mctchler, A. J. 1927. Supplement to the Leng Catalogue of Coleoptera of America North of Mexico. 80 pp. John D. Sherman Jr., Mt. Vernon, N. Y. MacGiixivray, A. D. 1923. External insect anatomy. 388 pp. Scarab Pub. Co., Urbana, Illinois. Manee, A. H. 1908. Some observations at Southern Pines, N. Carolina. Ent. News, 19: 286-289. McColloch, J. W. 1917. A method for the study of underground insects. Jour. Econ. Ent., 10: 183-187. , McColloch, J. W., and Hayes, W. P. 1923. Soil temperature and its influence on white grub activities. Ecology, 4: 29-36. McColloch, J. W, Hayes, W. P., and Bryson, H. L. 1928. The hibernation of certain scarabaeids and their Tiphia parasites. Ecology, 9: 34^2. 84 ILLINOIS BIOLOGICAL MONOGRAPHS [168 MtTLSANT, E. 1842. Histoire naturelle des coleopteres de France, Part II. Lyon (et Paris), Maison. (later Paris, Magnen and Blanchard Co.) Part 2, Lamellicornes. 624 pp., 3 pi. Osten-Sacken, R. 1862. Descriptions of some larvae of North American Coleoptera. Proc. Ent. Soc. Phila., 1: 105-130. 1 pi. 1874. Description of the larva of Pleocoma. Trans. Amer. Ent. Soc, 5: 84-87. Perris, E. 1877. Larves des coleopteres. 590 pp., 14 pi. (Lamellicorns, pp. 91-122, pi. 4, 5.) Dey- rolle, naturaliste, Paris. Phillips, W. J., and Fox, H. 1917. The rough-headed corn stalk-beetle in the Southern states and its control. U. S. D. A. Farm. Bui. 875: 1-10. 8 fig. Riley, C. V. 1870. White grubs in strawberry beds. Amer. Ent. and Bot., 2: 307. 1870. Insects injurious to the grape vine. Amer. Ent. and Bot., 2: 295. RlTTERSCHAUS, K. 1925. Eine neue Art von Eisprengern bei Lamellicornien-larven (Phyllopertha horticola L. und Anomala aenea DeG.). Zool. Anz., 62: 31-33. 1927. Studien zur morphologie und biologie von Phyllopertha horticola L. und Anomala aenea Geer (Coleop.). Zeits. Wiss. Biol., Abt. A, Zeits. Morph. and Okol. Tiere, 8: 271-408. SCHIODTE, I. C. 1874. De metamorphosi Euleutheratorum observations. Naturalist. Tiddskrift (series 3) 9: 227-376, pi. VIII-XIX. 1874. Note sur les organes de stridulation chez les larves des coleopteres lamellicornes. Ann. Soc. Ent. France, 5th series, 4: 39-41. Schmidt, A. 1910. Coleoptera Lamellicornia Fam. Aphodiidae. Genera Insectomm. Fasc. 110. 155 pp., 3 pi. Bruxelles. Sharp, D. 1918. Insects. Cambridge Natural History, Vol. 6. 626 pp. Macmillan & Co., London. Scopoli, J. A. 1763. Entomologia carniolica .... Vindobonas, 420 pp. Smith, J. B. 1902. An injurious root chafer. N. J. Agr. Exp. Sta. Ent. Rep. for 1901. Reference, p. 490. Steinke, G. 1919. Die Stigmen der Kaferlarven. Archiv. fur Naturges.,85 :(Abt. A) 1-58. 3pl. Smith, L. B., and Hadley, C. H. 1923. The Japanese beetle. U. S. D. A. Dept. Circ. 363: 1-66. Smyth, E. G. 1916. Report of the South Coast Laboratory. Porto Rico Dept. Agr. Rep. pp. 45-50. 1917. The white grubs injuring sugar cane in Porto Rico. II. The Rhinoceros beetles. Porto Rico Dept. Agr. Jour. 4 (No. 2) : 3-29. SWEETMAN, H. L., AND HATCH, M. H. 1927. Biological notes on Osmoderma with a new species of Ptiliidae from its pupal case (Coleoptera). Bui. Brook. Ent. Soc, 22: 264-266. Titus, E. S. G. 1905. Some miscellaneous results of the work of the Bureau of Entomology. The sugar- cane beetle (Ligyrus rugiceps Lee). U. S. D. A. Bui. 54: 7-18. 169] LARVAL SC ARAB AEOIDEA— HAYES 85 PLATE I 86 ILLINOIS BIOLOGICAL MONOGRAPHS [170 EXPLANATION OF PLATEJ Lateral Aspects of the Larvae oe Scarabaeoidea Fig. 1. Pinotus Carolina. Fig. 2. Sinodendron rugosum. Fig. 3. Passalus cornutus. Fig. 4. Onthophagus vaca (after Mulsant). Fig. 5. Amphicoma sp. Fig. 6. Aphodius sp. (probably fimetarius). ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 6 APHODIUS SP . HAYES LARVAL SCARABAEOIDEA PLATE I flpPftflY Of THE li^^smr of t^m 171] LARVAL SC ARAB AEOIDEA— HAYES 87 PLATE II 88 ILLINOIS BIOLOGICAL MONOGRAPHS [172 EXPLANATION OF PLATE II Lateral Aspects of the Larvae of Scarabaeoidea Fig. 7. Trox sp. Fig. 8. Anomala kansana. Fig. 9. Ochrosidia immaculata. Fig. 10. Ligyrus gibbosus. Fig. 11. Euphoria inda. Fig. 12. Phyllophaga crassissima. ABBREVIATIONS EMPLOYED ant antenna PC preclypeus as anal slit psc postclypeus es epicranial suture r radula f front I prothorax I labium II mesothorax mi mandible III metathorax mp maxillary palpus 1-10 abdominal segments ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII HAYES LARVAL SCARABAEOIDEA PLATE II UNMESSITY OF OiNOIS 173] LARVAL SC ARAB AEOIDEA— HAYES 89 PLATE III 90 ILLINOIS BIOLOGICAL MONOGRAPHS [174 EXPLANATION OF PLATE in Cephalic Aspect of the Larval Heads Fig. 13. Canthon laevis. Fig. 14. Aphodius sp. Fig. 15. Amphicoma sp. Fig. 16. Phyllophaga crassissima. Fig. 17. Serica sp. Fig. 18. Anomala kansana. Fig. 19. Ligyrus gibbosus. Fig. 20. Cotalpa lanigera. Fig. 21. Euphoria inda. Fig. 22. Troxsp. Fig. 23. Sinodendron rugosum. Fig. 24. Passalus cornutus. ABBREVIATIONS EMPLOYED ant antenna tnd mandible ea epicranial arm pc preclypeus es epicranial suture Pel precoila f front Psc postclypeus fcs fronto-clypeal suture v vertex I or lab labrum ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 13 CANTHON LAEVIS 14 APHODIUS SP 15 AMPHICOMA SP 19 UGYRUS GIBBOSUS 20 COTALPA LAMIGERA 21 EUPHORIA 1NDA SmODEWDgQN PUGOSUM y 24 PAS5ALUS COPNUTUS HAYES LARVAL SCARABAEOIDEA PLATE III up* OF . ' mmm r m mm 175] LARVAL SCARABAEOIDEA— HAYES 91 PLATE IV 92 ILLINOIS BIOLOGICAL MONOGRAPHS [176 EXPLANATION OF PLATE IV Epipharynx Fig. 25. Polyphylla decemlineata. Fig. 26. Phyllopkaga lanceolata. Fig. 27. Phyllophaga tristis. Fig. 28. Macrodactylus subspinosus. Fig. 29. Phyllophaga fusca. • Fig. 30. Dorcus sp. Fig. 31. Serica sp. Fig. 32. Phyllopkaga rugosa. Fig. 33. Sinodendron rugosum. Fig. 34. Diplotaxis sp. Fig. 35. Phyllophaga cribrosa. Fig. 36. Passalus comtUus. Fig. 37. Phyllophaga futilis. Fig. 38. Phyllophaga corrosa. Fig. 39. Trox sp. ABBREVIATIONS EMPLOYED cp chitinous plate Psa proximal sensory area els clypeal sensillia so sensillia cist clypeo-labral suture sc sense cone dsa distal sensory area sms submarginal striae 11 lateral lobe sp spines Is lateral striae st setae ml median lobe t torma ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII C P i S c v psa 25 PaYPHYUA PECEMLIHEATA dsa 26 PHYLLOPHAGA IANCEOLATA 27 PtlYLLDPtlAGA TR15TIS -psa 31 SERICA SP. 32 PHYOOPHACA RUGOSA cp--"'|---sc 33 SINODENDRON RUCOSUM 3< PHYLLOPttAGA FUTIL1S Cp' , v- x T sc psa^ck 38 PHYLLOPHACA CORROSA els 39 TROX SP. HAYES LARVAL SCARABAEOIDEA PLATE IV tmwt OF THE UNivcRsiit m usmm 177] LARVAL SCARABAEOIDEA—HAYES 93 PLATE V 94 ILLINOIS BIOLOGICAL MONOGRAPHS [178 EXPLANATION OF PLATE V Epipharynx Fig. 40. Anomala orientates. Fig. 41. Popillia japonica. Fig. 42. Ligyrodes relictus. Fig. 43. Anomala kansana. Fig. 44. Pelidnota punctata. Fig. 45. Ligyrus gibbosus. Fig. 46. Anomala innuba. Fig. 47. Polymoechus brevipes. Fig. 48. Strategus antaeus. Fig. 49. Anomala binotata. Fig. 50. Cotalpa lanigera. Fig. 51. Xyloryctes satyr us. Fig. 52. Strigoderma arboricola. Fig. 53. Ochrosidia immaculata. Fig. 54. Dynasles lityrus. cp els cist dsa 11 Is ml ABBREVIATIONS EMPLOYED chitinous plate Psa proximal sensory area clypeal sensillia so sensillia clypeo-labral suture sc sense-cone distal sensory area SUSS sub marginal striae lateral lobe sp spines lateral striae St setae median lobe torma ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII dsa dsa 40 ANOMALA ORIENTALS .dsa sp 41 POPlLLiA 4AP0NICA Sa ~\J/ /A^^^K- ^ Sa sp 42 LI&YRJDDFS RELICTU5 dsa cp sc "cts 43 ANOMALA KANSANA -t sc >-"■£_ _ 4_ - - els 44 PELIDNOTA PUNCTATA dsa 45 LIGYRU5 GIBBOSUS dsa 'V psa "sc 46 ANOMALA 1NNUBA dsa psa 47 POLYMOECHUS BREV1PES dsa SS. 1- st 48 5tR*ATEGU5 ANTAEUS sa 49 ANOMALY B1N0TATA dsa 50 COTALPA LANl&ERA dsa 51 XYL0RYGTE5 5ATYRJJ5 dsa ~~ cls\ 52 STRJGODERMA ARBO^ICOIA 53 OCHORDSIDIA IMMACULATA 54 DYNASTE5 TITYRU5 HAYES LARVAL SCARABAEOIDEA PLATE V ir y 01 THE 179] LARVAL SCARAB AEOIDEA— HAYES 95 PLATE VI 96 ILLINOIS BIOLOGICAL MONOGRAPHS [180 EXPLANATION OF PLATE VI Epipharynx Fig. 55. Canthon laevis. Fig. 56. Aphodius sp. Fig. 57. Euphoria fulgida. Fig. 58. Geotrupes stercorarius. (after Schiodte) Fig. 59. Cotinis nitida. Fig. 60. Euphoria sepulchralis. Fig. 61. Amphicoma sp. Fig. 62. Osmoderma eremicola. Fig. 63. Euphoria inda. Fig. 64. Copris tullius. Fig. 65. Cremastocheilus sp. Fig. 66. Trichiotinus piger. Fig. 67. Onthophagus sp. Fig. 68. Stephanucha pilipennis. Fig. 69. Phyllophaga futilis. sense cone U ABBREVIATIONS EMPLOYED chitinous plate psa proximal sensory area clypeal sensillia sa sensillia clypeo-labral suture sc sense-cone distal sensory area sms submarginal striae lateral lobe sp spines lateral striae St seta median lobe torma ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII ■psa 61 AMPttlCQMA 5P. ml 62 OSMODERJiA EREMICOLA dsa ml ! 11 63 EUPHORIA INDA. dsa sp 65 CREMAST0CHEILU5 SP. dsa St st- 66 TRJCHIOTINUS PIGER. 't psa 67 ONTHOPHAGUS SP. 68 STEPHANUCHA P1LIPENNIS 69 PHYLLOPHA&A FUTILIS HAYES LARVAL SCARABAEOIDEA PLATE VI 01 THE university m umm 181] LARVAL SC ARAB AEOIDEA- HAYES 97 PLATE VII 98 ILLINOIS BIOLOGICAL MONOGRAPHS [182 EXPLANATION OF PLATE VII Right and Left Mandibles. — Cephalic Aspect Fig. 70. Anomala kansana. Fig. 71. Aphodius fimetarius. Fig. 72. Stephanucha pilipennis. Fig. 73. Euphoria inda. Fig. 74. Cotalpa lanigera. Fig. 75. Ligyrus gibbosus. Fig. 76. Phyllophaga crassissima. Fig. 77. Mandibular articulation. Fig. 78. Canthon laevis. Fig. 79. Passalus cornutus. Fig. 80. Sinodendron rugosum. ABBREVIATIONS EMPLOYED ac acia b brustia dl dentes el extensotendon mo molar area pa preartis pel precoila rt rectotendon sb scrobe sc scissorial area ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 79 PASSALUS CORMUTUS 80 5IM0DEMD£0N KUGOSUM HAYES LARVAL SCARABAEOIDEA PLATE VII i — t 01 THE UNIYEPS'T OF 1UHQI3 183] LARVAL SCARABAEOIDEA—EA YES 99 PLATE VIII 100 ILLINOIS BIOLOGICAL MONOGRAPHS [184 Fig- 81. Fig. 82. Fig. 83. Fig. 84. Fig. 85. Fig. 86. Fig. 87. Fig. 88. Fig. 89. Fig. 90. Fig. 91. ac el dt el he mo EXPLANATION OF PLATE VIII Right and Left Mandibles — Caudal Aspect and Epipharynx Anomala kansana. Euphoria inda. Cotalpa lanigera. Phyllophaga crassissima. Canthon laevis. Amphicoma sp. Stephanucha pilipennis. Ligyrus gibbosus. Passalus cornutus. Pinotus Carolina. Epipharynx. Anomala kansana. Showing connection of epipharynx and hypopharynx. ABBREVIATIONS EMPLOYED acia PC clypeus pla dentes rt extensotendon sa hypopharyngeal chitinization sc molar area I precoila postartis rectotendon stridulating area scissorial area torma ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII sa-- -sa ac r „ -/ pta P ta '' rt --A J&~et 81 ANOMALA KAttSANA 82 EUPHORIA INDA 83 COTALPA LANLGEPA 84 p CPASSISSIMA 85 CAMTHOM LAEVIS 86 AMPtllCOMA SP 87 STEPHATiUCHA PILIPENNIS 88 LIGYKUS GIBB05US 90 PINOTUS 89 PASSALUS_ CAROLINA 91 ANOMALA KATiSATiA CORttUTUS HAYES LARVAL SCARABAEOIDEA PLATE VIII OF THE uwvBfcmr of klunois 185] LARVAL SCARABAEOIDEA— HAYES 101 PLATE IX 102 ILLINOIS BIOLOGICAL MONOGRAPHS [186 EXPLANATION OF PLATE IX Caudal and Lateral Aspects of Head; Antennae, Maxillae and Parts, Spiracles, Legs Fig. 92. Phyllophaga crassissi?na. Caudal aspect of the head. Fig. 93. Canlhon laevis. Caudal aspect of the head. Fig. 94. Passalus cornutus. Antenna. Fig. 95. Stephanucha pilipennis. Antenna. Fig. 96. Anomala kansana. Antenna. Fig. 97. Ligyrus gibbosus. Cephalic aspect of right maxilla. Fig. 98. Canthon laevis. Cephalic aspect of right maxilla. Fig. 99. Amphicoma sp. Cephalic aspect of right maxilla. Fig. 100. Aphodius sp. Cephalic aspect of right maxilla. Fig. 101. Anomala kansana. Prothoracic leg. Fig. 102. Anomala kansana. Metathoracic leg. Fig. 103. Passalus cornutus. Cephalic aspect of right maxilla. Fig. 104. Stephanucha pilipennis. Cephalic aspect of right maxilla. Fig. 105. Anomala kansana. Caudal aspect of right maxilla. Fig. 106. Anomala kansana. Cephalic aspect of right maxilla. Fig. 107. Anomala kansana. Articulation of maxilla and postgena. Fig. 108. Anomala kansana. Lateral aspect of the head. Fig. 109. Articulation of mandible and maxillae with head capsule. Fig. 110. Anomala kansana. Labrum. Fig. 111. Amphicoma sp. Stridulating teeth of right maxilla. Fig. 112. Anomala kansana. Stridulating tooth of left maxilla. Fig. 113. Anomala kansana. Stridulating teeth of right maxilla. Fig. 114. Cotalpa lanigera. Stridulating teeth of right maxilla. Fig. 115. Ligyrus gibbosus. Mesal aspect of fused galea and lacinia. Fig. 116. Cotalpa lanigera. Mesal aspect of fused galea and lacinia. Fig. 117. Canthon laevis. Mesal aspect of divided galea and lacinia. Fig. 118. Phyllophaga crassissima. Stridulating teeth of right maxilla. Fig. 119. Canthon laevis. Stridulating teeth of right maxilla. Fig. 120. Euphoria inda. Mesal aspect of fused galea and lacinia. Fig. 121. Phyllophaga crassissima. Mesal aspect of fused galea and lacinia. Fig. 122. Anomala kansana. Left prothoracic spiracle. Fig. 123. Trox sp. Left prothoracic spiracle. Fig. 124. Ligyrus gibbosus. Stridulating teeth of right maxilla. Fig. 125. Euphoria inda. Stridulating teeth of right maxilla. ABBREVIATIONS EMPLOYED ac alacardo of occipital foramen ant antenna opg occipito-postgenal suture cd cardo pa preartis d claw PC preclypeus ex coxa pel precoila es epicranial suture Pf palpifer / front pg postgena fnt femur pra parartis g gena pre paracoila I labrum Ps parastipes or subgalea Ic labacoria Psc postclypeus lp labial palpus pla. postartis m mala pie postcoila ntd mandible s stipes mla maxillaria or maxillary pleurite sc subcardo mp maxillary palpus sm submentum ms maxillary scraper si stridulating tooth mt mentum th terbio-tarsus mx maxilla tr trochanter oc occiput V vertex ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 94 PASSALUS 95 STEFHAHUCHA 114C0TALPA ^^ 1 1 8PHYLL0PHA&/T* 1 1 9 CANTHON 120 121 m$r 123 V_^ 124 LIGYRUS ^ 125 EUPHORIA 122 TROX . HAYES LARVAL SCARABAEOIDEA PLATE IX 1 7 0! THE IWEBSflT OF iLUNOIS 187] LARVAL SCARAB AEOIDEA- HAYES 103 PLATE X 104 ILLINOIS BIOLOGICAL MONOGRAPHS [188 EXPLANATION OF PLATE X Cephalic Aspect of Head; Hypopharynx, Stridulating Legs Fig. 126. Ochrosidia (Cyclocephala) immaculata. Cephalic aspect of the head Fig. 127. Pinotus {Copris) Carolina. Cephalic aspect of the head. Fig. 128. Cotalpa lanigera. Hypopharynx. Fig. 129. Euphoria inda. Hypopharynx. Fig. 130. Canthon laevis. Hypopharynx. Fig. 131. Phyllophaga crassissima. Hypopharynx. Fig. 132. Ligyrus gibbosus. Hypopharynx. Fig. 133. Anomala kansana. Hypopharynx and mandibles showing their relation to each other. Fig. 134. Anomala kansana. Hypopharynx with pharynx attached. Fig. 135. Ceruchus piceus. Metathoracic leg showing stridulating surface on the trochanter. Fig. 136. Ceruchus piceus. Mesothoracic leg showing stridulating surface on the coxa. Fig. 137. Passalus cornutus. Meso- and metathoracic legs showing the stridulating modifi- cation. ex fm he mi msc ABBREVIATIONS EMPLOYED coxa mtl metathoracic leg femur ph pharynx hypopharynx pta postcoila mandible tt tibio-tarsus mesothoracic leg tr trochanter ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 128 COTALPA LAHIGERA PIM0TU5 CAROLINA 126 OCHPJDSIDIA IMNACULATA 129 EUPHORIA INDA 132 LIGYR.US GIBB0S5US 130 CANTHON LAEVI5 131 PHYLLOPHAGA C-RA55I5S1MA. 133 IRPOPHAHYNX: 134 HYPOPHAPYNX AND MANDIBLES AND PHARYNX 135 ^M^ 136 CERUCHUS PICEUS 137 PA2.S.ALUS CORNUTUS CERUCHUS PICEUS _ . HAYES LARVAL SCARABAEOIDEA PLATE X UBRARV OF THE 1891 LARVAL SC ARAB AEOIDEA- HAYES 105 PLATE XI 106 ILLINOIS BIOLOGICAL MONOGRAPHS [190 EXPLANATION OF PLATE XI Radula of Last Ventral Abdominal Segment Fig. 138. Aphodius sp. Fig. 139. Amphicoma sp. Fig. 140. Serica sp. Fig. 141. Polyphylla variolosa. Fig. 142. Macrodactylus subspinosus. Fig. 143. Anomala kansana. Fig. 144. Pelidnota punctata. Fig. 145. Ligyrus gibbosus. Fig. 146. Strategus anteaus. Fig. 147. Dynastes tityrus. Fig. 148. Euphoria inda. Fig. 149. Euphoria sepulchralis. Fig. 150. Cotinis nitida. Fig. 151. Trox sp. Fig. 152. Passalus cornutus. Fig. 153. Sinodendron rugosum. Fig. 154. Dorcus sp. (Dorsal). Fig. 155. Dorcus sp. (Ventral). ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 141 1«' POUTHYLIA VARIOLOSA MACPODACTYLUS SUBSPINOSUS 143 ANOMALA KANSANA 146 144 '^ 145 N ___^ ^TOATrriK A1 smj ftTT o DYNASTES T1TYRUS PELIDNOTA PUNCTATA LIGYRUS GIBBOSU5 STRATTGUS AKTEAUS 148 EUPHOPJA IWDAeuph^ SEPUL cHPALI$ 150 7 |YI V 151 TRPX 3P connis nrriDA 152 PASSALUS CORTiUTUS 153 ^^ 154 _.-_.„ ._ 155 SINODENDPOtt PJJGOSUM D0 lgEL SP HAYES LARVAL SCARABAEOIDEA PLATE XI or the UMiVERsnry or hunois 191] LARVAL SCARAB AEOIDEA- HAYES 107 PLATE XII 108 ILLINOIS BIOLOGICAL MONOGRAPHS [192 EXPLANATION OF PLATE XII Radula or Last Ventral Abdominal Segment in Phyllophaga Fig. 156. Phyllophaga lanceolata. Fig. 157. Phyllophaga cribrosa. Fig. 158. Phyllophaga farcta. Fig. 159. Phyllophaga torta. Fig. 160. Phyllophaga latifrons. Fig. 161. Phyllophaga ephilida. Fig. 162. Phyllophaga longitarsa. Fig. 163. Phyllophaga gracilis. Fig. 164. Phyllophaga futilis. Fig. 165. Phyllophaga prunina. Fig. 166. Phyllophaga congrua. Fig. 167. Phyllophaga calceata. ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 156 PLAMCEOLATA 157 T? CRIBROSA 158 PFARCTA 159 PTORTA 162 PLOX1G1TARSA 165 PPRUH1NA 160 PLATIFROttS 161 PEPHILIDA 163 p GRACILIS 164 "PFUTIL1S 166 PCOHGRUA. 167 P CAIXEATA HAYES LARVAL SCARABAEOIDEA PLATE XII APIARY UNIVERSmr m ILLINOIS 193] LARVAL SC ARAB AEOIDEA-H AYES 109 PLATE XIII 110 ILLINOIS BIOLOGICAL MONOGRAPHS [194 EXPLANATION OF PLATE XIII Radula or Last Ventral Abdominal Segment in Phyllophaga Fig. 168. Phyllophaga crassissima. Fig. 169. Phyllophaga bipartita. Fig. 170. Phyllophaga micans. Fig. 171. Phyllophaga vehemens. Fig. 172. Phyllophaga fusca. Fig. 173. Phyllophaga horni. Fig. 174. Phyllophaga fervida. Fig. 175. Phyllophaga drakei. Fig. 176. Phyllophaga marginalis. Fig. 177. Phyllophaga fraterna. Fig. 178. Phyllophaga corrosa. Fig. 179. Phyllophaga profunda. ILLINOIS BIOLOGICAL MONOGRAPHS VOLUME XII 168 PCRAS5ISSMA 169 PBIPAPTITA 170 PMICAMS 171 PVEHEMEN5 172 PFU5CA 173 PHOPNl 174 PFEPVIDA 177 P FPATEPJ1A 175 PDPAKEI 178 P C0PJ2OSA 176 PMAR&IMAL1S 179 T? PROFUNDA HAYES LARVAL SCARABAEOIDEA PLATE XIII UP