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Arthropod and Nematode Parasites, Parasitoids, and Predators of in America North of Mexico

Technical Bulletin No. 1460

Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE ryyscji and Nematode Parasites, Parasitoids, and Predators of Ácrididae in America North of Mexico >,

ai,*' By N. E.|Rp)S^

Technical Bulletin No. 1460

Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE

Washington, D.C. Issued March 1973

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402 — Price: $2.60 domestic postpaid, or $2.25 GPO Bookstore Stock Number 0100-2624

ACKNOWLEDGMENTS

I express my sincere thanks to the following indi- viduals: The coleopterists of the Systematic Entomol- ogy Laboratory, U.S. Department of Agriculture, for checking the nomenclature of the Coleóptera ; George 0. Poinar, Jr., University of California, Berkeley, for his aid with the nematode section ; C. F. W. Muesebeck, U.S. National Museum, Washintgon, D.C., for advice on the Scelio ; A. S. Menke, Systematic Entomology Lab- oratory, U.S. Department of Agriculture, and R. M. Bohart, Entomology Department, University of Cali- fornia, Davis, for their assistance on the Tachytes; H. W. Prescott, Forest Grove, Oreg., for advice on the Nemestrinidae ; J. E. Henry, Investigation Laboratory, U. S. Department of Agriculture, for life information of Goniopsita oohaga ; W. F. Barr, Depart- ment of Entomology, University of Idaho, and R. L. Lavigne, Department of Entomology, University of Wyoming, for their assistance with the . I also give my special appreciation to C. C. Blickenstaff, Grasshopper Investigation Laboratory, U.S. Depart- ment of Agriculture, for his advice and counsel. The maps are reproduced by permission of Denoyer- Geppert©. CONTENTS

Pase Key to orders and families of larval parasites and predators.- 4 Key to orders and families of adult parasites and predators - 6 Grasshopper parasites and parasitoids '^ 9 Diptera Anthomyiidae ^ Calliphoridae ^^ Nemestrinidae ^^ Sarcophagidae ^^ ^^ Hymenoptera 77* ' Scelionidae "" Internal parasites °^ Nematoda |^ Mermithidae ^^ Gordiacea ^^ External parasites ^^ Hymenoptera ^^ Sphecidae ^^ Acariña Hyperparasitoids ^^° Hymenoptera ^^^ Bracomdae ^^"^ Chalcidae ^^^ Perilampidae ^^^ Pteromalidae ^^^ Sphecidae . 1^^ Diptera JJJ Calliphoridae ^^^ Predators ^^^ Coleóptera ^^^ Carabidae ^^^ Cicindelidae 1^2 Cantharidae ^'^^ Meloidae ^'^^ Diptera ^12 Anthomyiidae 212 Asilidae 215 Bombyliidae- 241 Chloropidae 248 Therevidae 250 Hymenoptera 253 Formicidae 253 Araneida 259 References 261 Index 286

IV Arthropod and Nematode Parasites, Parasitoids, and Predators of Acrididae in America North of Mexico

By N. E. REES, entomologist, Grasshopper Research Laboratory, Agricultural Research Service, United States Department of Agriculture, Bozeman, Mont.

This bulletin includes all avail- scriptions in the references listed able information relating to ar- at the end of this bulletin. thropod and nematode parasites The date and page numbers and predators of under the species name in each that occur in the United States section refer to the original de- and Canada. Although the litera- scription. In parentheses preced- ture was the main source of infor- ing the date is the original mation, some data were obtained generic name. Thus from unpublished records of the Blaesoxipha reversa (Aldrich) U.S. Department of Agriculture's (Sarcophaga) 1916: 135-136 Grasshopper Investigations Lab- was described by Aldrich as Sar- oratory at Bozeman, Mont. Parts cophaga reversa on pages 135 and of these records are from near 136 of his 1916 publication. Sacramento, Calif., and are re- Italic numbers in parentheses lated to a period prior to the in the text indicate the source in establishment of the Bozeman the reference list where informa- laboratory. tion is given about the species Each species is treated sepa- under discussion. Thus, for ex- rately. Insofar as possible, ample, (307, p. 233) refers to page information includes accepted 233 of R. W. Smith's 1958 article, nomenclature and synonyms, geo- which contains information on graphical distribution, host spe- Blaesoxipha reversa. This will en- cies, life history and habits, and able the reader to refer only to the descriptions of the egg, larval, pertinent part of an original pupal, and adult stages. The brief article. descriptions of the various stages For brevity, the following are presented only to aid the abbreviations are used: SO = reader in tentative identification. seasonal occurrence (dates given For a more thorough understand- here generally indicate the ing of each species, he is directed earliest appearance of each life to the original or subsequent de- stage in the field but not the dura- TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE tion of the stage unless so stated. menclature for the Diptera, and Several widely separated dates Agriculture Monograph 2, "Hy- often indicate several generations menoptera of America North of per year). LOD = length of de- Mexico," 1951, plus its supple- velopment of each stage (these ments, was the source for the are generally the results of lab- Hymenoptera. Later publications oratory investigations). OWS = and personal correspondence were overwintering stage (and location utilized for the accepted nomen- when known). G/Y = generations clature for the Coleóptera, Mer- per year or per given number of mithidae, mites, and spiders. years. SA = stage attacked (the They are also included in the ref- development stage of the prey or erence list. host that is utilized). N/H = Since the information was ob- number per host (number of tained from many sources, several parasites living at one time within points should be stressed. a single host, or the number of (1) Distributional records are prey utilized per day per pred- often few in number or for scat- ator). MOA — method of attack tered locations. It is therefore (method of parasitizing a host or highly probable that many species obtaining prey). AF — food taken listed here occupy broader ranges by the adult parasite or predator than recorded but with gaps cre- and includes plants frequented ated by sites of unfavorable en- that may or may not be eaten. vironment. Therefore the distri- HAB — habitat of the parasite or butional maps are generalized, predator. indicating the Provinces and The term "unknown" indicates States occupied by each species, that I believe no work has been and are not intended to show that conducted on this particular species distribution ends at Prov- phase. The term "not given" ince or State boundaries. means that although no mention (2) Figures given for incidence of an event was located in the of parasitism and prédation may literature, it is probably known be misleading. These generally by those working with that spe- represent the level of parasitism cies and probably conforms to and prédation at a given time and the general statement for the place and do not indicate the total family. The term "not located" effect on the host population. means that I believe the informa- (3) Seasonal occurrence rec- tion to be in print, but it was not ords are as given for the location located in my search. reported and may deviate as much Agriculture Handbook 276, "A as a month in other locations. Catalog of the Diptera of Ameri- (4) The amount of information can North of Mexico," 1965, was available indicates the amount of the main source of accepted no- work conducted on a species, but ARTHROPOD AND NEMATODE PARASITES IN AMERICA it does not present an accurate ), incomplete descriptions, indication of its effect on grass- and because predators are less hopper populations. An investi- likely to be taken with their prey. gator's interest and location or the Such a key is impractical here. importance of a certain species Generally several references to for other economic reasons is adult descriptions are included at often responsible for where a the end of this bulletin for those study is concentrated. who wish a more complete de- Keys to orders and families are scription. References to articles greatly simplified and designed to that quote original statements are lead the reader to the correct also included, since these may be group. He must then rely on a more easily obtained than the combination of information to ar- original. Even so, not all tax- rive at the correct species. onomic articles have been listed To effectively utilize the para- here because of nonavailability, site larval key, the larvae should superfluous repetition of the sub- be taken from the host. In sim- ject, and, of course, some articles plifying the keys for use by non- may have escaped the literature taxonomists, many details were search. omitted. These details might be In addition to the insects de- needed in separating nonparasitic scribed here, others could be in- species from parasitic ones, but cluded but are only mentioned they can be disregarded if the here. I have observed several reader is certain of the parasitic species of dragonflies taking both nature of the species involved. nymphal and adult grasshoppers. With some species, difficult taxo- At least one hemipteran, probably nomic characters had to be re- Reduviidae, was observed sucking tained and these are diagramed the body juices from an adult at appropriate places in the text. Melanoplus sanguinipes (F.). On In only a few groups are parasite several occasions mosquitoes were larval descriptions sufficiently seen partaking of grasshopper complete to allow keys for the hemolymph through the interseg- larval stage. It may also be neces- mental membranes of the neck sary to use additional available and abdomen while the host aids, such as descriptions, life seemed unaware of the event. histories, and habit information. Other insects may also utilize However, most specimens will grasshoppers as a source of food. have to be reared to the adult Some grasshoppers are canni- stage and the adult keys used. balistic, devouring freshly molted Specific keys were not con- or sick individuals, possibly to ob- structed for the predators because tain needed moisture. One species, of numerous similar species Brachystola magna (Girard), will (many are predatory on other ambush healthy, active hoppers on 4 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE occasion. A few of the larger hoppers or grasshopper eggs. The cricket species and some praying Diptera, Hemiptera, Neuroptera, mantids are also known to feast and Odonata orders contain fam- on weakened grasshoppers. ilies or genera different from, or In foreign countries some spe- in addition to, our reported para- cies of the coleopterous families sitie or predaceous ones. There- Cleridae, Curculionidae, Histeri- fore our list may still be very dae, Scarabaeidae, and Staphylin- incomplete, idae are also predaceous on grass-

KEY TO ORDERS AND FAMILIES OF LARVAL PARASITES AND PREDATORS la. Living within grasshopper egg, nymph, or adult; utilizing but one host during its lifetime 2 lb. Acting as grasshopper ectoparasite or predator 9 2a. Free living within hemocoel of host or within egg; not dependent on outside air source ^ 2b. Connected to outside air supply by tubular or funnel-shaped attachment 3 3a. Connected to body wall or trachéal trunk of host by long, convoluted respiratory tube; posterior spiracular slits arranged radially on circular plate Nemestrinidae 3b. Respiratory attachment short and funnel shaped; posterior spiracles protuberant, spiracle with button Tachinidae (in part) 4a. Wormlike, long, and slender, opaque to colored, without segmentation; oral (mandibular) hooks absent 5 4b. Muscoid or maggotlike in shape, tapering toward anterior end and blunt at posterior; or different but not wormlike 6 5a. Hairlike, very long and slender, with uniform, cylindrical body; anterior end is blunt and rounded, whereas posterior end is swollen and coiled ; many sensory bristles present; two minute eyes on head (horsehair worms) Gordiacea 5b. Body cylindrical, elongate, and often tapers toward both ends; much shorter and broader than horsehair worms; eyes absent Nematoda 6a. Definitely muscoid or maggot shaped; possesses oral (mandibular) hooks; not parasitic in eggs 7 6b. Parasitic in eggs, not found within nymphs or adults; first instar teleaform, second fusiform but lacks respiratory system, third of typical hymenopteran form with respiratory system Scelionidae 7a. Posterior spiracles located in deep cavity, usually without peritreme or button ; posterior spiracular slits not exceeding three ; pair of oral (mandibular) hooks Sarcophagidae 7b. Posterior spiracles not located in deep cavity 8 8a. Posterior spiracles widely spaced and with slits radiating almost in complete circle Anthomyiidae (in part) 8b. Infrabuccal plate present, single median mandibular hook, caudal spir- acles with button, caudal end of body truncate or broadly rounded Tachinidae (in part) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 5

9a. Ectoparasitic on nymphs or adults 10 9b. Predaceous on grasshoppers and/or their eggs during either or both of their larval (nymph) or adult stages 11 10a. Little or no differentiation of céphalothorax and abdomen; first instar with three pairs of legs and called larva; after first molt with four pairs of legs and called nymph; may be found in or on soil or under wings or wing pads of host (mites) Acariña 10b. Legless hymenopterous larva; located in constructed chamber in soil; associated with paralyzed host Sphecidae 11a. With four pairs of legs during all stages of growth; céphalothorax separated from abdomen by narrow stalk; very little metamorphosis (spiders) Araneida lib. Less than four pairs of legs during any stage, legs may be absent 12 12a. Legs completely lacking, no sign of rudimentary structures on thorax ._._ 13 12b. Legs present, may be articulate or merely rudimentary; Coleóptera larvae 18 13a. Legless, completely immobile, whitish; crescent shaped (like crookneck squash) with distinct head on narrow end; 13 distinct segments in addition to head; must be cared for and fed by adult members of colony; predatory only as adults (ants) Formicidae 13b. Not as above, capable of movement 14 14a. Oral (mandibular) hooks present; muscoid in form; antennae absent; head never deeply pigmented ; predaceous on eggs 15 14b. Antennae usually well developed; maxillae present; head and prothorax may be heavily pigmented 16 15a. Body elongate and narrow; posterior spiracular slits oval, posterior spiracles inconspicuous and not pigmented Chloropidae 15b. More muscoid in form; posterior spiracles on conical or domelike protuberances Anthomyiidae (in part) 16a. Posterior spiracles located on third or fourth segment from caudal end ; antennae buttonlike ; setae on thoracic segments shorter than diameter of each segment Therevidae 16b. Posterior spiracles located on second segment from caudal end 17 17a. Body "C"-shaped, never straight; caudal segment without setae; head somewhat hypognathous ; predatory in larval stage on eggs Bombyliidae 17b. Body straight; head and mouth parts usually hook shaped; mandibles usually much shorter than maxillae Asilidae 18a. Thoracic legs segmented and with one or two claws 19 18b. Thoracic legs rudimentary Meloidae (in part) 19a. Pair of hooks on dorsal aspect of fifth abdominal segment; no urogomphi on ninth segment; legs five-segmented and with two claws Cicindelidae 19b. Pair of hooks absent from fifth abdominal segment 20 20a. Legs five-segmented; urogomphi on ninth segment; one or two claws per leg Carabidae 20b. Urogomphi absent 21 21a. May be "C"-shaped larvae; lack dorsal glands on each thoracic and most abdominal segments Meloidae (in part) 21b. Body segments swollen and covered with velvety pubescence with occa- sional setae ; openings to paired dorsal glands present on each thoracic and most abdominal segments Cantharidae 6 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE KEY TO ORDERS AND FAMILIES OF ADULT PARASITES AND PREDATORS la. Legs absent 2 lb. Legs present ^— ^ 2a. Hairlike, very long and slender with uniform, cylindrical body, anterior end is blunt and rounded, whereas posterior end is swollen and coiled ; many sensory bristles present; two minute eyes located on head; generally found in water during adult stage Gordiacea 2b. Body cylindrical, elongate, and often tapering toward both ends; much shorter and broader than Gordiacea; adults generally found in or on grassy soils Nematoda 3a. Wings or rudimentation of wings absent 4 3b. Wings or vestigial wings present; six legs 6 4a. Three distinct body regions; six legs; thorax and abdomen connected by slender petiole on which spine is located; predaceous on very young grasshoppers (winged only at mating flight) Formicidae 4b. Head and thorax combined into céphalothorax; eight legs 5 5a. Little or no differentiation of céphalothorax and abdomen; predaceous on eggs and ectoparasitic on nymphs and adults (mites) Acariña 5b. Céphalothorax separated from abdomen by narrow stalk; predaceous on nymphs and adults (spiders) Araneida 6a. Two wings, thin and transparent 7 6b. Four wings 14 7a. Clothed with heavy pubescence 8 7b. Supporting bristles and/or setae H 8a. Top of head sunken between eyes, eyes bulging; body long and bristly; legs long; predaceous on nymphs and adults which are generally in act of jumping or flying; possess sharp beak which is used to puncture and paralyze host Asilidae 8b. Top of head not sunken between eyes 9 9a. Abdomen long and tapering, beard present, head rounded, legs long; predaceous on nymphs and adults Therevidae 9b. Abdomen stout, not long and tapering 10 10a. Empodia pulvilliform (tarsi with three pads) ; may or may not have long, tapering proboscis pointing downward or posteriorly—Nemestrinidae 10b. Empodia bristlelike or absent; tarsi with not more than two pads; proboscis long and tapering, pointing forward or downward—Bombyliidae 11a. Second antennal segment with longitudinal seam along upper, outer side; thorax with complete transverse suture; lower calypter large 12 lib. Longitudinal seam absent from second antennal segment; thorax with incomplete transverse suture; lower calypter small and rudimentary Chloropidae 12a. Arista generally bare; postcutellum developed; dorsal segments of ab- domen usually with strong bristles and fine hair ; lower edges of dorsal abdominal segments overlapping ventral segments Tachinidae 12b. Arista plumose for more than one-half its length; postscutellum not developed 1^ 13a. Arista plumose for only basal one-half, hypopleura and pteropleura with bristles; sponging mouth parts Sarcophagidae 13b. Arista plumose for most or all of its length 14 ARTHROPOD AND NEMATODE PARASITES IN AMERICA Y

14a. Hypopleura without bristles or if hypopleura are present then pteropleura are absent; posterior cell not narrowed apically; piercing mouth parts Anthomyiidae 14b. Hypopleura and pteropleura with bristles; first posterior cell narrowed or closed apically Calliphoridae 15a. Four thin membranous wings, upper pair larger ; mouth parts chewing— 16 15b. Upper wings hard, not membranous; chewing mouth parts 17 16a. Abdomen attached to thorax by narrow petiole; antennae curled; pro- notum short and collarlike with edges not extending to tegula and with rounded lobes on each side posteriorly; females stock their earthen chambers with paralyzed hosts which become living food for young Sphecidae 16b. Antennae elbowed, 11 to 12 segments or if clubbed only seven to eight visible segments; abdomen broadly attached to thorax; antennae in- serted very close to mouth; when gravid, females may be attached to abdomen of gravid female hosts just prior to their oviposition Scelionidae 17a. Tarsi of forelegs and hindlegs with same number of segments 18 17b. Tarsi of forelegs and midlegs five-segmented, hindtarsi four-segmented ; head wider than first part of thorax ; claws cleft or toothed Meloidae 18a. Antennae filiform or serrated; elytra usually broadly rounded apically and leathery, medial margin is metepisterna, curved; five, five, four tarsal sequence Cantharidae 18b. Elytra not leathery nor as described above 19 19a. Antennae inserted on front of head above base of jaws Cicindelidae 19b. Antennae inserted on side of head between base of jaw and eye Carabidae

GRASSHOPPER PARASITES AND PARASITOIDS

A parasite is an organism that dae, Sarcophagidae, and Tachini- lives at the expense of another, dae of the order Diptera and also either internally or externally, Scelionidae of the order Hymen- and generally utilizes only one optera. The Sphecidae (Hymen- host during its lifetime. A para- optera) and several species of sitoid is a type of parasite and is mites, nematodes, and horsehair defined as an that lives as worms are parasites and not a parasite within an arthropod, parasitoids, since they are either usually another insect, during a parasitic externally or not insects, part of its lifetime and is free The method of parasitism is living during the remainder. All unique for each species but is grasshopper parasites and para- usually very similar between sitoids are parasitic during their species of the same . Eggs larval stage and free living as of some parasites, such as several adults. Grasshopper parasitoids species of the Tachinidae, Sarco- include members from the fam- phagidae, and nematodes, are de- ilies Anthomyiidae, Nemestrini- posited near or on the food of the 8 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE host. When ingested, the eggs jured until after the development hatch and the larvae make their of the parasite is nearly complete. way through the gut wall into Larvae of the Sarcophagidae, the hemocoel where they complete Tachinidae, Anthomyiidae, and the feeding stages. nematodes wander freely through- Eggs of the nemestrinids are out the host's body. Nemestrinid deposited near the host in or on larvae construct a breathing tube wooden posts or in cracks of dry to the outside spiracle of the host vegetation. The larvae hatch and soon after entrance and are bound are blown to the ground among to the outside air supply until they their hosts. Unlike the larvae of mature. Within the grasshopper most , these first-instar larvae eggy scelionid larvae are also free are planidia, posesssing legs and living. capable of locomotion. When the Although a host may be para- host is located, entrance is ap- sitized by several individuals of parently accomplished through the same species, very seldom will the intersegmental membrane. it support more than one species. Once inside, the larvae molt to the Several species of Sarcophagidae second stage and a breathing tube and Nemestrinidae, because of to the outside is constructed. their size, mature singly within When mature, the larvae kill their a host. The strongest of the neme- hosts, often feeding on the re- strinids eliminate all competition maining muscle and nerve tissue. that may be present. When too Some species of Blaesoxipha many sarcophagids are present, (Sarcophagidae) deposit living all apparently suffer; their larvae directly on the flying or growth is retarded and the host jumping host. The larvae quickly often dies prematurely. Either burrow through the intersegmen- female scelionids avoid already tal membrane into the hemocoel. parasitized host eggs, or eggs They pass through three larval parasitized with more than one instars before departing, usually wasp produce only a single para- killing the host. site. Once inside a host, an internal The effectiveness of a grass- parasite begins to develop by hopper parasite or parasitoid is utilizing the available nutrients of generally dependent on its host its host. These generally include specificity, the number and types the hemolymph, fat body, and of alternate hosts, and its adapta- reproductive organs and their bility to a wide variety of ecolog- products. With very few excep- ical conditions. Host-specific para- tions the digestive tract, nervous sites are at the complete mercy of system, and muscles are not in- the fluctuations of the host pop- ARTHROPOD AND NEMATODE PARASITES IN AMERICA

ulation and usually lag behind may be rapid under favorable during their hosts' population in- conditions. crease. Those parasites that utilize many hosts, some or most of which are of other orders, Díptera generally show no preference for their grasshopper hosts. Since Anthomyîîdae they have such a wide range of This family of flies contain a individuals to use, they seldom large number of species and is affect grasshopper populations. worldwide in distribution. In gen- Most grasshopper parasites and eral appearance the adults are parasitoids, however, utilize at similar to the house except the least three species of hosts, all of mouth parts of most have been which are grasshoppers. Prefer- modified for a predaceous way of ence is usually expressed, and the life. Prey of adult flies usually other hosts are available for use consist of other flies, often of the if preferred species are scarce; same family. The genus Lispa is thus there is less fluctuation in noted as being predaceous on the parasite population. aquatic larvae. Most of the dipterous parasi- Larvae are predators, parasites, toids are not able to survive if plant feeders, or scavengers. Hy- their grasshopper host dies before lemya platura (Meigen) (given in their larval development is com- many publications as Hylemya pleted. Others will die if the host cilicrura Rohd.) is both a serious succumbs before they are able to crop pest destroying corn seed and exit, even though their develop- beneficial as a predator of grass- ment is completed. hopper eggs. Acridomyia canaden- One of the advantages of some sis Snyder is an important para- dipterous parasitoids, such as site of grasshopper nymphs and those of the Blaesoxipha group, is adults in Canada. that they have two to four genera- Eggs of anthomyiid flies are tions per year. In this way they normally oval and white to cream are able to build up their popula- colored. Larvae of the grass- tion rapidly during a year when hopper parasitic species are mus- the host is plentiful. Females of ciform and near white. These de- many of these species produce velop through three instars within from 40 to 80 viable larvae per their hosts before emerging, litter, and a few species produce entering the soil, and pupating. more than one litter in a lifetime. Females of these species are Although the total potential is equipped with a barbed ovipositor, seldom approached, the buildup which insures that the host does 10 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE not escape until oviposition is complete. Taxonomists do not agree as to whether the genus Acridomyia be- longs in Anthomyiidae or in Muscidae. The main differentia- tion between the two families is that the anthomyiids have the first posterior cell open, whereas the cell of the Muscidae is closed or strongly constricted. Since the posterior cell of the two species MAP 1.—Recorded distribution of of Acridomyia listed here is open, Acridomyia canadensis Snyder. I have placed them in the family Anthomyiidae. Incidence of parasitism.—Up Acridomyia canadensis Snyder to 68.8 percent of M. packardii in 1940: 1 Canada and 63.5 percent of M. Synonymy.—None (320, p. oregonensis in Montana. Although 915). many species are heavily parasi- Recorded distribution.—Brit- tized in Canada, M. bivittatus and ish Columbia, Saskatchewan, M. packardii are preferred. (307, Manitoba, Idaho, and Montana. pp. 249-251; 389). (320, p. 915; 38U; 389). (Map Life history and habits.—SO: 1.) Eggs are deposited in July and Host species.—Ageneotettix de- August in Montana. Larvae have orum (Scudder), Aulocara elliotti been found within adult grass- (Thomas), pellucida hoppers in Montana from July 28 (Scudder), Melanoplus angusti- to September 25. Pupae occur pennis (Dodge), M. bivittatus from August to June or July in (Say), M. bruneri (Scudder), M. the soil. Adults first appear in dawsoni (Scudder), M. femurru- the field between mid-July to brum (DeGeer), M. foedus August in Montana and about Scudder, M. infantilis Scudder, M. early June in Canada. LOD : Eggs oregonensis (Thomas), M. pack- unknown. Larvae mature in about ardii Scudder, M. sanguinipes 16 days. Pupae mature in 10 to (Fabricius), Phoetaliotes nebras- 22 months. Adults unknown. censis (Thomas), OWS: Pupae in the soil. G/Y: sp., and sp. (123, pp. Generally one, although some in- ^70-472, 475, 487; 227, pp. 935- dividuals do not emerge until the 936; 306, pp. 28-33; 307, pp. 249- second season. SA: Nymphs and 251; 311, pp. 1-2; 323, p. 264; adults. N/H: 27 to 70 from un- 384; 389). known hosts, 47 in Xanthippus, ARTHROPOD AND NEMATODE PARASITES IN AMERICA 11 and an average of 29.5 from M. Description.—Eggs measure oregonensis in Montana. MOA: 0.5 mm. long by 0.2 mm. wide, Eggs are deposited directly into are cream colored to white, and the hemocoel of the host. AF: are elongate-oval (307, pp. 2i9- Only the females feed on the 251 ; 309, pp. 99,101-102). Larvae hemolymph of the host. Food of (first instar) are 0.6 mm. long by the male is probably nectar and 0.25 mm. wide, are translucent to other plant secretions. HAB : The ivory white, lack noticeable rangelands of Canada, whereas in spines, and have vague segmenta- Montana it can be found in fields tion. The buccopharyngeal arma- of crested wheatgrass on rolling ture is simple and supports a foothills up to 7,200 feet and in single median tooth. First and saddles containing very sparse later instars are separated by the and low-growing grasses. shape of the mouth parts, and the Adults mate shortly after second and third stages differ in eclosión. After a short gestation the form of the anterior and period the female begins stalking posterior spiracles. These are prospective hosts, punctures the widely spaced and almost circu- integument with its rasping lar ; they have radiating papillae. mouth parts, and ingests the re- (309, pp. 99,101-102) Pupae are sulting fluid. Near the completion reddish to dark brown and 3.75 of feeding, the fly brings it ovi- mm. long by 1.5 mm. wide (389). positor forward and inserts it into Adults are minute flies, 4 to 4.5 the feeding wound. Eggs are de- mm. long. Head, thorax, and ab- posited into the hemocoel where domen are black; the last two they mature for several days and areas are covered with a yellow- hatch to larvae. Smith (307) in- ish-gray pruinescence. Parafron- dicates the number of eggs pro- tals, face, and cheeks are grayish, duced per female to be similar to with a brownish-yellow pruines- that produced by A. sacharovi cence. Eight or nine pairs of (Stackelberg), about 200, but I short, fine, parafrontal bristles believe this number is too high. and many fine, pale, hairlike In most of my dissections the bristles run along the ventral eggs never exceeded 100. margin of the cheeks. Antennae There are three free-living lar- are brownish yellow and descend val stages within the host. Pupa- from their insertion opposite the tion occurs about 12 to 48 hours middle of the eyes. Presutural after emergence from the host, thoracic bristles and the four post- and the fly enters diapause, which sutural dorsocentrals are barely is not affected by cold. (227, pp. distinguishable from the sur- 935--936\ 307, pp. 2^9^251; 309, rounding setae. The male pos- pp. 99, 101-102; 311, pp. 1-2; sesses a distinct, wide dorsocen- 389) tral stripe on the abdomen, which 12 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE is lacking in the female. Legs characteristics, such as heat tol- range from yellow in the female erance, habitat, seeking prey, and to tawny yellow in the male; the aggressiveness. apical tarsal segments of all legs Acridomyia fumisquama Snyder are blackish brown. There are only two median posterior bristles 1940: 2 on the midtibia, the foretibia is Synonymy.—None (320, p. bare at the middle, and the 915). bristles of the hindtibia are un- Recorded distribution.—North distinguishable. Wings are a Carolina (311, p. 2; 320, p. 915). milky-white hyaline and the veins (Map 2.) yellowish. Calypters are a pale Host species.—Sweetman (323, brownish yellow and the haltères p. 26i) reported the larvae as be- orange. (311, pp. 1-2) ing parasitic on grasshoppers and Discussion.—Although more gave Smith and Finlayson (309, than one individual of A. cana- p. 101) as the source. However, densis can complete development Greathead (123, pp. U70-U72) in a single host and emerge with- stated this species has not been out killing it, the fecundity of the reared, and neither the articles by host is reduced. Under normal Smith and Finlayson (309, p. 101) conditions, however, many eggs nor Snyder (311, p. 2) support hatch and the larvae mature and Sweetman's statement. I there- depart within a short time of fore included this species with each other, resulting in the death doubt. of the host. If immature larvae Incidence of parasitism.—None are present at this time, they located. either die within the host or exit Life history and habits.—SO: prematurely and rarely survive. Eggs, larvae, and pupae unknown. This parasite prefers the cooler climates, extending southward into the northernmost parts of the United States. The only reported U.S. collections come from Idaho and Montana. It is an effective grasshopper parasite in Canada, but its value in Montana and Idaho has never been ascertained. Its close taxonomic relationship to the Eurasian relative, A. sacharovi, has caused some to re- late the findings of sacharovi to A. canadensis. However, I find MAP 2.—Recorded distribution of them to be different in many Acridomyia fumisquama Snyder. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 13

Adults collected in September in only two notopleural bristles and North Carolina. LOD, OWS, and flesh flies have four. G/Y: Unknown. SA, N/H, and Most blow flies are scavengers ; MOA : Not given. AF and HAB : the larvae live in carrion, excre- Unknown. ment, and decaying flesh. A for- Description.—Eggs, larvae, and eign species, Stomorhina lunata pupae unknown. Adults are very- (F.), departs from the general similar in both size and appear- habit of the family in being pre- ance to A. canadensis except the daceous in the egg capsules of anteroventral and posteroventral Locustana pardalina Walker bristles on the hindfemora are ( ). Estimates from more numerous, longer, and closer South Africa indicate that as together. Legs are dark brown to many as 50 to 90 percent of egg black. Wings are a smoky-brown masses are destroyed by these hyaline. The calypters and hal- maggots and as many as 50 eggs tères are dark brown. (311, p. of the predator can be found in 2) a single egg pod. Discussion.—This species was In the United States only one described by Snyder (311) in the species of blow fly has been re- same article in which he gave his ported as being parasitic on grass- original description of A, cana- hoppers. The eggs of this species densis. Since no original parasite are deposited directly on the records could be located, its status grasshopper and the larvae be- as a grasshopper parasite is come internal parasites. After questionable. completion of development, the larvae exit from the host and Callîphorîdae pupate in the soil. In these re- spects they resemble in habit The calliphoridae, or more com- several species of the Sarcopha- monly the blow flies, constitute gidae. a large number of species and are very similar to the flesh flies Calliphora vicina Robineau- (Sarcophagidae). The two groups Desvoidy are combined by some authorities 1830: 435 into a single family, the Metopi- Synonymy.—Calliphora ery- idae. Blow flies are very often throcephala Meigen, Musca ery- metallic in color, with the arista throcephala Meigen (preoccu- of the antennae plumulose to the pied) C. rufifades Macquart (320, tip, whereas the flesh flies are p. 929). blackish with gray thoracic Recorded distribution.—Can- stripes and have the aristae bare ada and United States including for approximately half their Alaska. Also found in South length. Blow flies generally have America, southern Africa, and 14 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE South Australia (320, p. 929). ulation was diminished by at (Map 3.) least one-tenth to one-eighth. The Host species.—M elanoplus writings of these three men in- spretus (Walsh) (26^, p. 32i). dicate that the attack occurs near Incidence of parasitism.—See molting time and that the larvae MOA. cause the death of the grass- Life history and habits.—SO: hopper within a short time, after Eggs, larvae, and pupae unknown. which the larva emerges from the Adults not given. LOD : Unknown. grasshopper, kills it, and enters OWS and G/Y: Not given. SA: the soil. AF and HAB : Unknown. Nymphs and adults. N/H: Not {26i, p. 32A) given. MOA: Riley (26^, p. 32i) Description.—Eggs, larvae, and included statements by Daniel pupae unknown. Adults: Rob- Parker, C. E. Bessey, and W. L. ineau-Desvoidy {270, p. Jf35) gave Coleman relating to the attacks a short description of C. vicina of this species on grasshoppers. and noted it was very similar to Parker reported that the eggs Calliphora vomitoria (Linnaeus). were deposited at the junction of Discussion.—Although speci- the thorax with the abdomen and mens were apparently collected that the flies were most active at and identiñed, it is interesting to about the second molt of the grass- note that the only references to hopper. Bessey reported attacks this particular species being para- on the grasshopper Melanoplus sitic on grasshoppers occur in the differentialis (Thomas) and noted writings of Riley {26Í.). Descrip- that the grasshoppers were nearly tions of the fly as being blue or completely eaten out. He esti- green and the fact that it was mated that the grasshopper pop- identified indicate that it was not confused with any of the sarco- phagi ds of the flesh fly group. Since it is rather common throughout Canada and the United States, its single reported attack on grasshoppers suggest that it is only an incidental para- site of this insect.

Nemestrinidae

"■^ ■ itf. Only two species of Nemestrini- \<" 1- ..•^' dae are known to parasitize grass- hoppers in Canada and the United

MAP 3.—Recorded distribution of States. Both are similar in ap- Calliphora vicina Robineau-Desvoidy. pearance and habits throughout ARTHROPOD AND NEMATODE PARASITES IN AMERICA 15 their life stages but differ in their several days within the trachéal host ranges. Adults are good fliers trunk, possibly for feeding, the but are extremely clumsy on the larva reenters the hemocoel for ground. Although the adult is the remainder of its development. short lived, each female produces During this time it maintains a thousands of eggs, which are tubular conenction with the out- deposited in holes of side air. The location of the at- weathered posts and cracks of tachment is dependent on the dry vegetation. The eggs hatch species. Four feeding larval in- after several weeks, and the small stars are passed within the host mobile ñrst-instar larvae are before the mature larva leaves, blown about by the wind until killing the grasshopper in the they either locate a host or suc- process. cumb after many days of search- The mature larva burrows into ing. In this latter respect they the soil, overwinters, and pupates are extremely hardy. in the spring. The pupa can move The manner of larval entrance and migrate to the soil surface into the hemocoel of the host is prior to emergence of the adult. not yet fully understood. In the Adults of both species have a laboratory Prescott (2i7) ob- yellowish fur and are extremely served Neorhynchocephalus sac- similar in appearance. Eyes of kenii (Williston) larvae entering both are kidney shaped, and the the host through the soft interseg- pitch of their buzzing is different mental membrane of the abdomen from other insects. Females of and Trichopsidae clausa (Osten both species possess a very pli- Sacken) through the thoracic or first abdominal spiracles. Spencer able ovipositor, which first ap- (317) y on the other hand, had pears as a stinger but which will difficulty duplicating this. flex easily when touched. When Upon entrance into the host the these flies are numerous, they larva seeks a trachéal trunk and greatly reduce grasshopper pop- burrows into its center. After ulations.

KEYS TO SPECIES OF NEMESTRINIDAE

FIRST-INSTAR LARVAE Metathoracic filament present AT. sackenii Metathoracic filament vestigial T. clausa

OLDER INSTAR LARVAE Four oval flat-topped projections located on dorsal-caudal slope, which lies in arc between rounded spiracles and end of body T. clausa Four oval flat-topped projections absent N, sackenii 16 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

PUPAE Proboscis long N. sackenii Proboscis very short T. clausa

ADULTS Proboscis long and very obvious N. sackenii Proboscis short and indistinguishable T. clausa

Neorhynchocephalus sackenii 2Jf8, pp. 513-521; 2A9, pp 557- (Williston) 566; 307, pp. 251-253; 315, p. 18; (Rhynchocephalus) 1880: 243 317, pp. 503-509; 323, pp. 109, Synonymy.—Rhynchocephalus 110, 310, U67; 383, pp. 5-10). sackenii Williston, R. subnitens Incidence of parasitism.— Cockerell U,p.219; 320, p. i02). From 20.9 to 41.6 percent of C. Recorded distribution.—From pellucida and 12 to 70 percent of southern British Columbia to M. nevadensis in Canada. High central Manitoba, south into incidence is also noted from areas Georgia, west into California, and of Montana, Idaho, and Arizona, north into British Columbia {307, with the highest rate being 90 pp. 251-253; 320, p. ^02). (Map percent of M. sanguinipes in Mon- 4.) tana and 99 percent of this species Host species.—Aeropedelliis in Oregon. (2^8, pp. 513-521; clavatus (Thomas), Ageneotettix 307, pp. 251-253; 389) deorum (Scudder), Amphitornus Life history and habits.—SO : coloradus (Thomas), Boopedon Eggs are deposited from May nubilum (Say), Camnula pellucida to late July in Canada, mid-June (Scudder), sordi- to mid-July in Idaho and Mon- dus (Scudder), E. sordidus costa- tana, and mid-September to late lis (Scudder), Melanoplus alpinus (Scudder), M. bivittatus ('Say), M. dawsoni (Scudder), M. fem- urrubrum (DeGeer), M. gladstoni Scudder, M. infantilis Scudder, M. sanguinipes (Fabricius), M. spretus (Walsh), neva- densis (Bruner), Morseiella flaviventris (Bruner), and Phoe- taliotes nebrascensis (Thomas) (18, pp. 163, 165, 169-171 ; 19, p. 104; 123, pp. Í61-Í6Í, U75, A79; 126, p. 212 ; 152, pp. U, 37-38 ; 2U7, MAP 4.—Recorded distribution of Ne- pp. 392-393, 396-398, 400-^02; orhynochocephalus sackenii (Williston). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 17

September in Arizona. Larvae to 5,000 small eggs in beetle holes can be found from mid-May to of weathered posts and in cracks early August and through the and crevices of bark, timber, and winter in Canada and late June other forms of vegetation, such to late July and through the as Eriogonum alatum and sage- winter in Idaho and Montana. In brush. Apparently warm tempera- Arizona, larvae are present from tures and the presence of a breeze late September to early October or wind are stimuli for oviposi- and through the winter. Pupae tion. occur in the spring in all locations. After several weeks the eggs Adults first appear in the field in hatch and the small mobile larvae Canada between May and mid- are caught up in the wind and dis- July and between mid-June and persed. They may search many early July in Idaho and Montana. days before either locating a host In Arizona they are present or dying. Very often the survival around mid-September. Adults rate is extremely low. remain alive for only a short Initial penetration into the host time. LOD : Eggs require 8 to 10 apparently occurs through the days. Larvae need about 2 weeks soft intersegmental membrane of in the host. Pupae average 10 to the abdomen. After a short time 32 days. Diapause requires 43 to the parasite establishes contact 110 days. Adults live only 3 to 8 with outside air by means of a days. OWS: As larvae in soil. respirator tube. The origin of the G/Y : One, although some will re- tissue that constructs its respira- main in diapause for an additional tory tube is not certain. The two year or so. SA: AU. N/H: Only most popular concepts suggest one survives. MOA: First-instar that either it is tissue from the planidia seek out the host. AF: parasite or the tube is formed Frequent bigflower groundsmoke from host tissue through the effect (Gayophyturn diffusum T. & G.) of the parasite. Nevertheless as and Achillea millefolium {2U7, pp. the larva ages the respiratory 392-393, 396-398, Í00-Í02; 307, tube becomes longer, allowing the pp. 251-253 ; 37i, p. 71 ; 388 ;389), larva to move about within the HAB: Generally rangeland type, hemocoel of the host. This tube such habitats as fenceposts, cer- is apparently shed during each tain plants. molt with the host's exuvia. The Adult flies can ingest fluid larva first ingests only liquid nourishment. Nevertheless this food. It later attacks the organs stage is short lived, lasting in the of reproduction and their prod- laboratory for 3 to 4 days, some- ucts and finally expands its food times as much as 8. Mating occurs source by destroying and scrap- shortly after emergence, after ing the remaining host tissues vhich the female deposits 4,000 from the body walls. When ma- 18 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE ture it departs through the mem- year after the larva has left its brane between the thorax and host. In some larvae this molt abdomen or thorax and head (fig. may be delayed for several years. 1) and immediately enters the Adults are excellent ñiers but soil. quite clumsy on the ground. When Although a host may be para- in flight, the pitch of their buzz sitized by several nemestrinid varies depending on their func- larvae, only one will survive to tion. Males, for example, posi- complete development. This in- tioned on alfalfa or other vegeta- tense competition is also ex- tion attempting to attract a pressed between the nemestrinid female, have a lower buzz than larvae and any other parasite when just hovering, and the buzz larvae that attack the same host. is more uniform than when the N. sackenii larvae pass through males are skittish or when a fe- four molts. The first occurs im- male is seeking a site for oviposi- mediately on entering the host, tion. Although the eyes are large, followed by three feeding stages sight is apparently extremely within the hemocoel. The last limited. One may approach flying molt to a pupa occurs nearly 1 or stationary flies and gently en-

PN-272B FIGURE 1. Larva of Neorhynchocephalus sackenii (Williston) emerging from male Melanoplus femurmbrum (DeGeer) host. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 19

close them in a box or in the hand as long as no sudden movement is made. Apparently these flies com- municate by the sound of their wings rather than sight. (247, pp. 392-393, 396-398, iOO-402; 2A8, pp. 513-521; 2^9, pp. 557- 566; 307, pp. 251-253; 389)

Description.—Eggs are very similar in size and appearance to those of Trichopsidae claiisa PN-2726 (Osten Sacken). They are elon- FIGURE 2.—Third-instar larva of Neo- gate, slightly curved, and pointed rhynchocephalus sackenii (Willis- at both ends. They are held to- ton) : Left, dorsal; right, lateral. gether with a weak cement. (247, pp. 392-393, 396-398, 400- 402; 317, pp. 503-509) Larvae bushy yellowish-white hair be- (ñrst-instar planidia) are mobile coming nearly white below. An- and able to stand in an upright tennae are short and reddish position. They are vermiform yellow. Three joints make up the with definite segmentation and style ; the first is short and yellow- setae projecting in pairs on the ish, the second is twice as long as ventral areas of the abdominal the first, and the third is as long segments. There are four poste- as the first two together. The riorly directed filaments, two lat- proboscis reaches to the hind- erally directed filaments, and four coxae. The labium is black and projecting from the ventral sur- the palpi are yellowish. The dorsal face of the caudal segment. Each part of the thorax and the scuttel- ventral filament projects from a lum is brownish black and sup- small conical protrusion. Thoracic ports yellowish hair, whereas the segments bear a pair of filaments pleura and pectus bear long bushy not supported by projections. gray-white hair. The abdomen is (313, pp. 25-27; 314, p. 87; 388; short and broad. Yellowish hair 389) (Fig. 2.) Pupae not given. intermixed with black at the in- (Fig. 3.) cisures covers the basic black. The Adults are 9 mm. long and feet are yellow and the femora are black with light-yellowish hair. white with tufts of hair on the The head is brownish black and underside near the coxae. The broader than the thorax. Vertex, wings are hyaline. (373, p. 243) lower part of the face, cheeks, and Discussion.—On many occa- posterior orbitals of the front all sions this species has been cred- support blackish hair with thick ited with having a major role in 20 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Columbia (320, p. i03). (Map 5.) Host species.— pseudo- nietana (Thomas), Camnula pel- lucida (Scudder), Melanoplus sanguinipes (Fabricius), Metator nevadensis (Bruner), M. paradal- inus (Saussure), áspera Scudder, and T. kiowa (Thomas) (19, p. 10^; 123, pp. Jt61-A6Jt, 4-80 ; 183, pp. 23-24 ; 307, pp. 251-254; 312, pp. 21-24; 313, pp. 25-27; 314, p. 87; 317, pp. 503-509; 323, pp. 109-110, 467; 383, pp. 5-10). Incidence of parasitism.—None PN-2727 located. FIGURE 3.—Pupa of Neorhynchoce- phalus sackenii (Williston) : Left, Life history and habits.—SO: dorsal ; right, ventral. Eggs are deposited in late June to mid-July in Canada. Larvae are present from late July until controlling grasshoppers in spe- spring in Canada. Pupae occur in cific areas. Its presence in an area the spring. Adults appear in late may be enhanced by weathered June to mid-July in Canada. LOD : fenceposts, which are used to Eggs, larvae, and pupae unknown. house the eggs. N. sackenii is Adults live only 3 to 5 days. OWS : extremely vulnerable during the As larvae in soil. G/Y : One. SA : first-instar planidial stage while Fifth instars and adults produce it seeks a host, and unless condi- tions are ideal, the survival rate is extremely low. (317, pp. 503- 509) Trichopsidae clausa (Osten Sacken) (Hirmoneura) 1877: 225 Synonymy.—H irmoneura clausa Osten Sacken (4, p. 218; 320, p. i03). Recorded distribution.—From British Columbia into Ontario, south through Florida, west into MAP 5.—Recorded distribution of California, and north into British Trichopsidae clausa (Osten Sacken). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 21 parasites, although all stages may tral filament projects from a small be attacked. N/H: One. MOA: conical protrusion. Each thoracic Planidia seek out the host. AF: segment bears a pair of filaments Believed not to feed as adults. not supported by projections. In HAB: Rangeland type, generally separating the two species of flat, and with fenceposts for ovi- Nemestrinidae, the metathoracic position. filament is vestigial in T. clausa Mating occurs shortly after and evident on N. sackenii. Later eclosión, and oviposition takes instar stages may be separated by place shortly thereafter. Follow- using the four oval flat-topped ing hatching, the larvae are projections on the dorsal caudal blown about until such time as declivity lying in an arc between they either die or are able to the conspicuous rounded spiracles penetrate the integument of a at the end of the body, which are host. Upon entrance, the life cycle present on T. clausa and absent is similar to that of N. sackenii on N. sackenii. Also the oral hooks except the respiratory tube be- are less pointed on the latter than comes attached to the trachea of in the former. (807, pp. 251-25Í ; the host instead of to the ventral 888) (Fig. 4.) Pupae have the fold of the body wall. The host proboscis sheath much shorter in is killed on emergence of the para- T. clausa than in N. sackenii (807, site. Larvae burrow into the soil, pp. 251-25^). remaining there until pupation in Adults are 9 to 10 mm. long, the spring. Pupae are mobile and basically black, and covered with able to migrate to the soil surface a pale yellowish-gray hair. Both immediately prior to eclosión. the antennae and feet are reddish (307y pp. 251-25Í ; 312, pp. 21-2^) and the eyes are bare. The face is Description.—Eggs are elon- densely covered with a pale yel- gate, slightly curved, and pointed lowish hair as is the thorax. The at both ends. Thus they are very proboscis is present but barely similar to those of N. sackenii distinguishable because it is small. {807, pp. 251-25Jf). Larvae (ñrst- The black abdomen is covered instar planidia) are mobile and with the same pale yellowish hair able to stand in an upright posi- that is found on the rest of the tion. They are vermiform with body. The legs are brownish red definite segmentation and setae and the femur is covered with pale projecting in pairs from the ven- yellowish erect hair. The wings tral areas of the abdominal seg- are hyaline and the haltères red- ments. There are four posteriorly dish. (281, p. 225) (Fig. 5.) directed filaments, two laterally Discussion.—Adults apparently directed filaments, and four pro- are unable to feed and take on jecting from the ventral surface fluids as does N. sackenii and of the caudal segment. Each ven- therefore are believed to survive 22 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE the larval stage; relatively few species are parasitic. Of the latter, only some are parasitic on insects ; the others confine their activities to snails, earthworms, other in- vertebrates, and mammals, even man. Only a few of those affecting insects are parasitic on grass- hoppers (23 species), but they generally are considered the most effective of the parasites that in- fect nymphal and adult grass- hoppers.i These parasites are of two groups, oviparous (egg laying) and ovoviviparous (those hatch- ing eggs within the uterus and depositing living larvae directly on the body of the host). Eggs of the first group may be laid on or near the host or, as with Sarco- •m^K- phaga haemorrhoidalis Fallen, di- rectly on the food of the host. The most important and most PN-2728 FIGURE 4.—Pirst-instar larva of Tri- numerous, however, are the para- chopsidae clausa (Osten Sacken) sites of the second group, which without metathoracic filament. deposit larvae directly on the host in a manner that is characteristic of each species. For example, Blaesoxipha kellyi (Aldrich) and no longer than 3 to 5 days in the B. reversa (Aldrich) prefer a field. moving target, such as a jumping Collection and host records are nymph or a flying adult. The not so numerous as for N. sack- larvae are placed either on the enii. This, coupled with a slightly underside of the wings, usually narrower host range, indicates it near the base, or somewhere on is less important as a grasshopper the abdomen. These larvae quickly parasite. It is nevertheless very find an area of soft or membran- effective under ideal conditions.

Sarcophagidae ' This excludes pathogenic micro-or- ganisms, which are generally associated Most species of sarcophagids, with disease, but which by some defini- or flesh flies, are scavengers in tions are considered as parasites. AR' ^HROPOD AND NEMATODE PARASITES IN AMERICA 23 ous integument, such as at the on the other hand, prefers its tympanum or between segments, victim on the ground and not very and with their sharp mouth hooks active. It stalks its prey and at the (fig. 6) they tear a small hole in proper moment brings its ovi- the host's body and burrow into positor forward and tucks a small the hemocoel. larva within the natural folds of Several species can be enticed the host's body. Blaesoxipha opi- to chase a moving object, such as fera (Coquillett), which also pre- a thrown dirt clod or piece of fers its victim fairly quiet and on wood, even though they are not on the ground, attacks in several mature enough to oviposit or nor- ways. It will sometimes "flip" its mally do not oviposit on a moving ovipositor and throw a sticky host. It is questionable, however, larva at the host, whereas at other even with species requiring a times it pounces on the back of moving host, if this movement is the host, drops a larva, and jumps the stimulus for oviposition. away before the grasshopper has Blaesoxipha hunteri (Hough), time to dislodge it.

PN-2729 FIGURE 5.—Adult of Trichopsidae clausa (Osten Sacken). 24 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

wings

oral hooks

¡nfrahystomal bridge^ ^hypostomal sclerite

FIGURE 6.—^Mouth parts of sarcophagid larva.

Within the body of the host the extensive but includes species of larva develops through three other orders. The successful de- larval instars. It feeds on the he- velopment of a parasite within a molymph or fat body or both of host is apparently dependent its host, but it usually does not on the physiology of the host as injure host tissues, which might expressed by the relationship of cause rapid death of its victim. most dipterous grasshopper para- The reproductive organs and their sites and Melanoplus bivittatus products are either destroyed or (Say). Parasites that complete greatly reduced so that if the host their development before the host survives, it cannot reproduce. becomes an adult usually survive, Usually the host dies immediately but those that do not are killed prior to or shortly after exit of and melanized. the larva. Some die and are Incidence of parasitism is ex- "cleaned out" as the larva scrapes tremely variable, ranging from the digestible tissues from the in- zero to better than 50 percent. tegument before departing. Those This alone, however, is not a cor- that die after the parasite has rect assessment since many spe- departed may do so either from cies have three and four genera- the effects of the parasite or from tions per year and each female secondary infection of the exit produces many larvae. Therefore wound. the overall seasonal effect may be A few of the sarcophagids are much greater. host specific, but usually several Food requirements of the adults species of grasshoppers are in- range from sugars, such as found involved. For several sarcophag- in cut vegetation, to decaying ani- ids, not only is the host range mal and vegetable matter. The ARTHROPOD AND NEMATODE PARASITES IN AMERICA 25 larvae ingest the blood, fat body, chorion but immediately hatches and products of the reproductive and enters the host. systems, such as eggs of the host. Most larval descriptions are of The larvae feed, leave their hosts, the first instar. The larvae of all and are thought not to eat again species are similar in shape, taper- until they become adults. Several ing toward the caudad end from authors, however, indicate that the large and rounded posterior they may feed on grasshopper end. The anal spiracles are found eggs in the soil {818, p. 31). within a hollow Wsterior cavity, Since most of this group are which in many specimens has the ovoviviparous, egg descriptions floor lined with setae. Spines on are generally lacking in the litera- all or most of the segments are ture. Eggs are rather small, range of specific significance. (Fig. 7.) between 30 and 80, and are white At the end of the third instar, to translucent, and elliptical. As larvae exit and burrow into the larvae develop within the eggs, soil. At this stage there is some the oral hooks and buccopharyn- variation in color and opaqueness geal armature can be seen as between species. Colors range black structures within the trans- from white to cream. In some lucent bodies. On completion of species, usually those that exit embryonic development, the larva from already dead and decaying leaves the egg but remains in the grasshoppers, discolored reddish- egg sac. In one species the larva brown to purplish food substances is deposited while within the can be seen within the digestive

anterior posterior cavity

iuccopharyngeal armature oral hooks ^spines

FIGURE 7.—Generalized sarcophagid larva. 26 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE tract through the semitransparent trained taxonomist, and descrip- integument. In other species the tions of most females are lacking larvae are opaque and the diges- in the literature. tive tract or its contents are not The primary characteristic of visible. sarcophagid flies is that the The pupal cases are yellow to arista of the antenna is plumose yellov^ red immediately after pu- to the middle or sometimes just pation and darken with age to beyond and the distal part is bare deep dark red to black. The cases (fig. 8). Other distinguishing are cylindical with horizontal characteristics are the color of the outfoldings laterally on the an- parafrontals and parafacials, the teriormost area. The posterior arrangement of bristles on the cavity and anal opening are vis- parafrontals, the size of the var- ible posteriorly. Pupal size varies ious segments and joints of the considerably both among and antennae with relation to each within species depending on other, and the color and shape of availability of food to the larvae. the proboscis and palpi. The tho- The case appears as a thickened, rax usually contains three to five sclerotized covering and is not black stripes on a gray poUinose smooth, nor is it usually exces- body and also many bristles, sively wrinkled. Segmentation is which are useful for taxonomic rather plain. Outstanding fea- purposes. Two of these bristles, tures of the case are usually the the postsutural dorsocentrals and color, size, and wrinkledness, the anterior acrostichals (which along with the shape of the an- were used by Aldrich (5) in his terior spiracles and the shape and keys), are used here for separa- arrangement of the slits on the tion. posterior spiracles within the Wings are hyaline and may or posterior spiracular cavity. Ad- may not contain costal spines. ditional information is sometimes Legs vary mainly according to the obtained from certain infoldings number and position of bristles or depressions of various struc- and the presence or absence of tures, such as the anal opening. combs or massive numbers of Adult sarcophagid males are hairs. The abdomen has four vis- similar in appearance. Specific ible segments with various ar- differences are in size and in the rangements of bristles on the genitalia. Females differ from distal parts. For our purposes we males in that the eyes are more are including only the bristles of widely separated and slightly the fourth abdominal segment. smaller and, of course, in the Posterior to this is the hypopy- possession of an ovipositor. Spe- gium, which consists of two seg- cific identification of females is ments and which often may be red usually very difficult, even for the to reddish yellow. For positive ARTHROPOD AND NEMATODE PARASITES IN AMERICA 27

antenna^ ,arista

.anterior acrostichals anterior dorsocentrals- ostsutural dorsocentrals costal vein.

fennun tarsus

hypopygium

FIGURE 8.—Generalized sarcophagid adult. identification the most useful portional size of the forceps, characters of the male sarcopha- claspers, and penis. These are gid are the collar shape and pro- illustrated by Aldrich (5).

KEY TO ADULT MALE SARCOPHAGIDS PARASITIC ON GRASSHOPPERS (Based on Aldrich (5) ) la. Three distinct postsutural dorsocentrals 2 lb. Four postsutural dorsocentrals 19 2a. Hindtibiae with villosity 3 2b. Hindtibiae lacking erect villosity 6 3a. Hypopygium entirely black; mid-tibia with patch of yellow hair cov- ering apical third of front side Sarcophaga sinuata Meigen 3b, At least second segment of hypopygium red 4 4a. With almost complete median row of bristles on hindfemur Boettcheria cimbicis (Townsend) 4b. Hindfemora not with long bristles in middle series 5 5a. Anterior acrostichals well developed Blaesoxipha cessator (Aldrich) 5b. Anterior acrostichals not developed . .Sarcophaga sarraceniae Riley 6a. Hypopygium entirely black 7 6b. Hypopygium red on at least second segment 8 28 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

7a. Second segment of hypopygium opaque; palpi yellow; forceps slender with protuberance on attached part, on which is striking tuft of long, straight, black hair Blaesoxipha hunteri (Hough) (in part) 7b. Second segment opaque; palpi black; forceps yellow and strongly curved; no tuft of hair Blaesoxipha atlanis (Aldrich) 8a. First vein of wing hairy; anterior acrostichals hardly differentiated from hairs Helicobia rapax (Walker) 8b. First vein bare 9 9a. Palpi yellow 10 9b. Palpi black H 10a. Forceps with pair of processes on basal attached part, which bear striking tuft of long, straight, black hair Blaesoxipha hunteri (Hough) (in part) 10b. Without above processes or tuft Blaesoxipha opifera (Coquillett) 11a. First segment of hypopygium black; forceps entirely yellow Sarcodexia sternodontis Townsend (in part) lib. Both segments of hypopygium yellow 12 12a. Distal segment of penis in form of long tube that can be seen through in profile beyond lateral processes Blaesoxipha reversa (Aldrich) 12b. Distal part not forming loop that can be seen through 13 13a. Distal segment of penis on its apical half with lateral, thin, expanded margins, which are whitish or transparent, somewhat like a ruffle 14 13b. Penis not with rufflelike expansions 16 14a. Distal segment divided into two subsegments beyond hinge 15 14b. Distal segment not distinctly divided; distal part swollen and whitish near end Blaesoxipha coloradensis (Aldrich) 15a. Division of distal segment near tip, which is divided into two delicate processes attached to fringe Blaesoxipha uncata (Wulp) 15b. Division just beyond middle, apex broadly and deeply cleft into two strong but flat claws, which are not connected with fringe Blaesoxipha falciformis (Aldrich) 16a. Front generally less than one-tenth of head width Blaesoxipha augustifrons (Aldrich) 16b. Front at least one-sixth of head width 17 17a. Stout divergent prongs at tip of penis cover and conceal minute incurved copulatory tube, or at most have V-shaped opening Blaesoxipha aculeata (Aldrich) 17b. Prongs spread apart and are U-shaped so as to show much larger in- curved tube 19 18a. Penultimate lateral processes of penis extend forward and are thick and blacky with spinose outer side - ..Blaesoxipha gavia (Aldrich) 18b. Penultimate lateral processes extend forward only short distance, but extend backward and underneath in form of thin brown margin Blaesoxipha taediosa (Aldrich) 19a. Hindtibiae with villosity 20 19b. Hindtibiae lack villosity 24 20a. Hypopygium entirely black Sarcophaga harpax Pandelle 20b. At least second segment of hypopygium red 21 21a. First segment of hypopygium black on at least apical half - 22 21b. First segment yellow on at least apical half 23 ARTHROPOD AND NEMATODE PARASITES IN AMERICA 29

22a. Prescutellars present; penis with four slender processes at tip Sarcophaga shermani Parker 22b. Prescutellars absent; forceps nearly straight in profile with notch behind near base ..___ Sarcophaga haemorrhoidalis (Fallen) 23a. Anterior acrostichals distinct Blaesoxipha impar (Aldrich) 23b. Anterior'acrostichals not diflferentiated; forceps nearly parallel front and back edges Sarcophaga sarracenioides Aldrich 24a. Anterior acrostichals either undifferentiated or but little larger than adjacent hairs Sarcodexia sternodontis Townsend (in part) 24b. With one or two pairs of erect anterior acrostichals, several times as large as adjacent hairs; anterior dorsocentrals also large and erect Blaesoxipha kellyi (Aldrich)

Blaesoxipha aculeata (Aldrich) high as 25 percent (307, pp. 219- {Sarcophaga) 1916: 143-144 221). Synonymy.—Acridiophaga ac- Life history and habits.—SO: uleata (Aldrich), Sarcophage ac- Eggs, larvae, and pupae unknown. uleata Aldrich (320, p. 9H). Adults first appear in Canada Recorded distribution.—South- about late June to early July and ern Canada and the United States remain until late September. (5, p. lU; 307, pp. 219-221 ; 320, LOD : Gestation requires about 9 p.9U;389). (Map 6.) days. Eggs unknown. Larvae Host species.—Ageneotettix de- average 12 days. Pupae mature in orum (Scudder), Amphitornus 13 days. Adults unknown. OWS: coloradus (Thomas), Aulocara As larvae in the soil. G/Y: Two elliotti Thomas, Camnula pellu- to three. SA: Adults and fifth- cida (Scudder), Melanoplus bivit- instar nymphs. N/H: Not given. tatus (Say), M. femurrubrum MOA: Live larvae are deposited (DeGeer), M. packardii Scudder, M. sanguinipes (Fabricius), M. spretus (Walsh), anaSchistocerca americana (Drury) (5, p. 14.4.; 38, p. 6; 123, p. 48i; 12A, pp. U, 13; 152, pp. 37-38; 300, pp. 58, 60, 61; 306, pp. 28-33; 307, pp. 219-221; 308, pp. 187-199; 318, pp. 31-3A; 325, p. 14; 326, p. 22.) Also the species E. opaca (listed by Greene (124, pp. U, 13)), but the order and genus are not given. Incidence of parasitism.—In

Canada it averages 2 to 3 percent MAP 6.—Recorded distribution of with spotty locations reaching as Blaesoxipha aculeata (Aldrich). 30 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE on moving or flying hosts. AF and segments 5 to 11. The dorsal part HAB: Unknown. of the bands on segments 5 to 8 Mating may occur any time is separated from the ventral part after emergence, followed by an by a small gap. Segments 10 and 8- to 11-day gestation period. 11 are without dorsal spines. Each female produces on an Hairlike bristles are on the diskal average 44 larvae per litter. areas below the midlateral line of Smith (307) indicated that fe- segments 2 to 11. Six short spines males may produce more than one surround the anal opening and a litter. First-instar larvae are able scattering of spines is on the floor to remain healthy in the uterus of the posterior cavity. Four for up to 3 weeks after hatching. fleshy lobes are on the posterior The adult fly attacks moving or ventral surface of the last ab- flying grasshoppers and deposits dominal segment. Two stout oral from one to two larvae at a time hooks curve slightly downward per host, which immediately bur- and outward at the tips from the row into the hemocoel through the buccopharyngeal armature. The softer parts of the integument. infrahypostomal bridge is fused These develop through the usual with the hypostomal sclerite. three larval stages, which require Within the posterior cavity are about 9 days, and then emerge, the usual pair of two-lobed, sharp- killing the host. After burrowing tipped spiracles and a pair of into the soil, the larvae either clawlike processes with apical pupate within several days or spines. {309, p. 91) enter diapause. Nondiapausing Pupae are large, measuring 8.5 adults emerge 14 to 16 days later. mm. long to 3.0 mm. wide, and Those in diapause require a tem- are a dull yellowish red to dark perature of 8° C. for 67 days red. Tubercles on the edge of the before they can continue develop- posterior cavity are fairly indis- ment. (S07, pp. 219-221; 319, tinct. Anterior spiracles located pp. 300-301 ; 389) near the apex of the puparium Description.—Eggs are gen- contain five yellow lobes on a dull- erally opaque white and taper to- reddish basal part. The posterior ward both ends. Larvae (first cavity is small, elliptical, and instar) are 1.5 mm. long by 0.3 shallow. Within are two deep-red mm. wide, translucent white, spiracular plates, each containing cylindrical, and tapering. The three large yellow slits, which are first eight segments are encircled pointed at the posterior end. The with spines and the ninth has a two middle spiracular slits are dorsolateral patch of spines. A nearly parallel with each other. A transversely divided band of small anal opening is darkened spines is on the ventral half of and faintly depressed. It is located ARTHROPOD AND NEMATODE PARASITES IN AMERICA 31 between the upper and lower hor- He therefore speculated that hosts izontal depressions. {12.Í, pp. U, other than grasshoppers are 13) probably involved, although none Adult flies are 6 to 9 mm. long. were observed. He indicated that Antennae are dark brown to this parasite was of little eco- blackish. The third joints are nomic importance. nearly double those of the second. Aristae are plumose for slightly Blaesoxipha angustifrons more than half their length. Palpi (Aldrich) are long and the proboscis is short (Sarcophaga) 1916: 142-143 and fleshy. Outer vertical bristles Synonymy.—Acridophaga ca- are absent. Thorax is a gray pol- ridei (Brethes), Sarcophaga ca- linose with the usual three to five ridei Brethes, S. caridei Aldrich, stripes. Anterior acrostichals are S. angustifrons Aldrich {320, p. well developed, but there are only 9U). three postsutural dorsocentrals. Recorded distribution.—Brit- The abdomen is gray pollinose ish Columbia, Kansas, Texas, New and tessellated. It has a rather York, North Carolina, and Geor- distinct median blackish line. The gia (320, p. 9U). (Map 7.) fourth abdominal segment is en- Host species.—Schistocerca tirely black. From it rises a row cancellatta Serveille, S. paranen- of more than a dozen bristles. sis Burmeister, Dichroplus macul- The hypopygium of the male is ipennis (Blanch) Liebermand, D. entirely yellow. The first segment pratet ñs Bruner, and D. elonga- contains a row of bristles across tus G. Tos (6, p. 590 ; 203, pp. 216, the hind margin, and the second 221-228). segment supports numerous erect, Incidence of parasitism.—0.5 coarse black hairs. Forceps are yellowish brown toward the apex and concave. Legs are black. The middle femur possesses a short comb on the posterior edge. Two bristles are on the outer side of each middle tibia, whereas the hindtibiae lack villosity. Wings are hyaline and have small costal spines. (5, pp. 143-14.^) Discussion.—Smith (307, p. 221) observed that although fe- male flies in the field were gravid and surrounded by ample hosts, MAP 7.—Recorded distribution of the rate of parasitism was low. Blaesoxipha angustifrons (Aldrich). 32 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE percent of D. macuUpennis in posterior. There is a slight sep- 1949 in South America (6, p. 590). aration between the dorsolateral Life history and habits.—SO: parts of the encircling spines on Eggs, larvae, and pupae unknown. segments 5 to 7 from the venter Adults not given. LOD : Gestation half. Near the anal opening are requires 7 to 10 days (caridei). several short spines and on the Eggs unknown. Larvae mature in floor of the posterior cavity is a 5 to 7 days (caridei). Pupae take patch of the same. The two fairly 13 to 16 days (caridei). Adult stout oral hooks curve downward (caridei) females in the labora- and outward to sharp tips. tory take 30 to 69 days to mature Within the posterior cavity are a from the larval to adult stage. pair of two-lobed, sharp-tipped OWS: Not given. G/Y: Believed spiracles, the outer lobes of which to be three (caridei). SA: Not are longer than the inner. (309, given. N/H: Normally only one. pp. 90-91) Pupae unknown. MOA: Deposit first-instar larvae Adult males are 8 to 8V2 T^^- on the host in flight or jumping. long. Antennae are dark brown to AF and HAB : Unknown. blackish. The third joints are A. caridei females produce up nearly twice the size of the sec- to 104 larvae. These are deposited ond. Aristae are plumose for on the abdomen, one or two at a slightly more than half their time. They immediately enter the length. Palpi are long, whereas hemocoel of the host through in- the proboscis is short and fleshy. tersegmental membranes where There is a pale beard below and they feed on the internal organs. the usual three rows of black When mature they emerge at any hairs lining the back of the head. point in the abdomen. (203, pp. Outer vertical bristles are absent. 216, 221-228-, 316, pp. 500-501) The thorax is gray pollinose and Description.—Eggs unknown. supports the usual three to five Larvae (as described by Smith black stripes. Anterior acrosti- and Finlayson (309) for A. cari- chals are well developed, although dei) : First-instar larvae are ap- there are only three postsutural proximately 1.3 mm. long by 0.3 dorsocentrals. The gray pollinose mm. wide, translucent white, and abdomen is tessellated and con- cylindrical with a forward taper- tains a distinct median blackish ing. Posterior end is blunt. The line. More than a dozen bristles in first seven segments are encircled a row are on the entirely black with spines anteriorly. On the fourth segment. The hypopygium ventral half of segments 4 to 11 is yellow and shiny. The first are dense transverse bands of segment is medium sized and sup- thornlike spines, which are di- ports a row of bristles along the vided on the venter so that the hind margin, and the second seg- anterior part is wider than the ment is globose and has several ARTHROPOD AND NEMATODE PARASITES IN AMERICA 33 erect, coarse black hairs. Forceps The S. caridei of Argentina and are reddish brown and almost Uruguay is an aggressive parasite straight on the front edge. of the grasshoppers Schistocerca All legs are black; the hind- cancellata Serveille and S. para^ tibiae lack villosity. A short comb nensis Burmeister, but it appears is located on the posterior edge only to be of minor importance. and below on the middle femur. It was introduced into Canada On the outer front side of the but apparently was never re- middle tibia are two bristles. The covered. Although specimens of female is very similar to the male the Canadian S. caridei (or B. except the genital segment is red, angustifrons) were collected in the tergal section supports long different areas, I was unable to marginal bristles, and the sternite find any mention of hosts associ- is developed into a short blunt ated with them. Therefore I in- point, which does not bend for- clude B. angustifrons only as a ward and barely protrudes from possible grasshopper parasite in the abdomen in repose. Also there Canada or the United States. is a row of three bristles on the outer front side of the middle Blaesoxipha atlanis (Aldrich) tibia. (5, pp. H2-H3) {Sarcophaga) 1916: 100 Discussion.—There is some Synonymy.—Sarcophaga at- question as to whether or not B. lantis Aldrich, Tephromyiella angustifrons is parasitic on grass- frankliniana Townsend {320, p. hoppers as the result of confused 9^5). synonjmiy. Spencer and Bucknell Recorded distribution.—South- {318, p. 31) reported that the ern Canada and in the United species S, caridei, which was in- States north of a line from south- troduced into Canada from South ern California through Virginia America, is the same as their local {320, p. 9i5). (Map 8.) S. caridei and that their S, caridei Host species.—Aeropedellus is the same as B. angustifrons. clavatus (Thomas), Ageneotettix Parish and Gushing (23^), also deorum (Scudder), Camnula pel- reported caridei from Texas, lucida {Scudder ), Chorthippus whereas Aldrich (6, p. 590) and longicornis (Latreille), Dendro- Lloyd {203, pp. 216, 221-228) re- tettix quercu^ Packard, Melano- ported it from South America. plus angustipennis (Dodge), M. Stone et al. {320, p. 9U), how- bivittatus (Say), M. confusus ever, indicated that the S. caridei Scudder, M. dawsoni (Scudder), of South America is not the same M. femurruhrum (DeGeer), M. as the S. caridei of Canada and foedus Scudder, M. packardii presented the North American Scudder, M. sanguinipes (Fabri- species as B. angustifrons. Other cius), and Spharagemon collare records support either view. (Scudder). The status of A. de- 34 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE 17. MOA: Unknown in the field. In laboratory investigations the female approaches active or rest- ing hosts, extends the tip of the abdomen forward between its ?%■■•& ■... legs, and places one or more lar- vae on the host. AF and HAB: Unknown. (307, pp. 2Í1-2U) Description.—Eggs: No de- scription was located because the .^^■"'■^ eggs are hatched in the uterus. \ Larvae (first instar) are 1.2 mm. MAP 8.—Recorded distribution of long by 0.25 mm. wide, translu- Blaesoxipha atlanis (Aldrich). cent white, rounded posteriorly, and tapering anteriorly. Segments orum as a successful host is 1 to 7 are each nearly completely questionable since Smith (307) encircled on the anterior margin reported that all observed para- with posteriorly directed spines, sitic larvae had died before ma- whereas segments 9 and 10 are turing within this species. (38, encircled with anteriorly directed p. 6; 123, pp. U76, k8h; 12A, PP. i, spines. Spines on segments 3 and 10-11; 258, p. 88; 300, pp. 58, 4 are stout, well pigmented, cone 60-61; 306, pp. 28-33; 307, p. shaped, and closely located in 2i3 ; 318, p. 30 ; 325, p. H ; 326, p. bands several rows wide. Fusi- 23) form bands of spines are on the Incidence of parasitism.—The venter of segments 5 to 10, five highest reported rates of parasit- to six rows wide, with the larger ism were 54.5 percent of M. spines in the anterior row. Spines angustipennis, about 15 percent of are lacking on the floor of the M. sanguinipes, and 50 percent of posterior cavity, whereas several C. pellucida in Canada (307, p. surround the anal opening. Oral 2 A3). hooks are sicklelike and relatively Life history and habits.—SO: straight when viewed from the Eggs, larvae, and pupae unknown. ventral profile. The infrahyposto- Adults are found anytime after mal bridge and the hypostomal the first of June in Canada with sclerite are not fused. Within the parasitism only in August and posterior cavity is a pair of two- September. LOD: Gestation re- lobed, sharp-tipped spiracles quires 9 days. Eggs unknown. with the usual spiracular slits. Larvae 10 days. Pupae 12 days. (309, pp. 86, 95) Adults unknown. OWS : As larvae Pupae are 6.0 mm. long by 2.0 in soil. G/Y: Three in Canada. mm. wide and are a dull yellowish SA: Mostly adults. N/H: Up to red. The posterior end of the un- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 35 wrinkled case is more broadly ceps are yellow with blackened truncate than the anterior, and tips, slender, though not diver- the last two segments are distinct. gent, and strongly hooked for- Anal tubercles are lacking. An- ward. terior spiracles are small and each Legs are black and the hind- contains seven lobes on a dull tibiae lack villosity. A short dark-red base. Within the small well-developed comb is located on and shallow posterior cavity are the hind part of the middle femur, two red spiracular plates, each and two bristles are on the outer with three short, narrow yellow front side of the middle tibia. The slits ; the middle slits are parallel third joint of the female antennae with each other. Near the lower is less slender than that of the edge of each spiracular plate is male and it is more grayish. Outer a wrinkled area. The anal open- vertical bristles are present. The ing is small, dark, and inconspic- abdomen is distinctly reddish, and uous. {12Jf, pp. Uy 10-11) the genital segments are a dull-red Adult length ranges between opaque and enclose a well-devel- 51/^ and 8 mm. Male antennae are oped short larvipositor similar to black. The third joints are slender that of J5. hunteri (Hough). The and twice as long as the second. hindfemur of the female lacks a Aristae are plumose as usual. comb. (5, pp. 100-102) Both palpi and proboscis are black Discussion.—S m i t h {307) and ordinary. Three rows of abun- stated that although the incidence dant black hairs are located be- of parasitism is generally below hind the eyes on the back of the 15 percent for M. sanguinipes, it head and the outer vertical is nevertheless a persistent para- bristles are absent. The thorax is site. In Canada it can be found gray pollinose and displays the associated with B. hunteri. In usual three to five stripes. There Montana it has been collected only are only three postsutural dor- rarely since 1939. socentrals and two pairs of large anterior acrostichals. Abdomen is Blaesoxipha cessator (Aldrich) gray pollinose with three stripes ; (Sarcophaga) 1916: 84 the lateral ones are sometimes Synonymy.—Sarcophaga cessa- changeable. A row of about 16 tor Aldrich (320y p. 9Í6). lateral bristles is located on the Recorded distribution.—Ore- fourth segment. The first segment gon, Idaho, California, Arizona, of the hypopygium is small, and New Mexico {320, p. 9JÍ6). brown, and pollinose and has a (Map 9.) row of six to eight small bristles Host species.—Melanoplus dif- near the apex. The small reddish- fer entialis (Thomas) {386). brown second segment lacks con- Incidence of parasitism.—None spicuous hairs or bristles. For- located. 36 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE on the hind margin. The second ^ segment is globose, red, and not bristly. Forceps are slender and - tapering, gently curving forward at the apex. Legs are black. A short comb of only six blunt * spines is located behind and be- low on the midfemur. There are two bristles on the midtibia on the ' outer front side near the middle and a moderate villosity spread- ing both ways on the hindtibia. ■ MAP 9.—Recorded distribution of Both the pulvilli and the claws Blaesoxipha cessator (Aldrich). are elongated. Wings are hyaline. (5, pp. 84-85) ^ Life history and habits.—SO: Discussion.—About nine speci- Eggs unknown. Larvae are active mens were reared from M. differ- around August 1 in California. entialis collected in the fall of Pupae unknown. Adults not given. 1927 at Patterson, Calif., and LOD, OWS, G/Y, SA, N/H, MOA, identified by J. M. Aldrich. The AF, and HAB : Unknown. (389) lack of additional host records Description.—Eggs, larvae, and indicates this to be a parasite of pupae unknown. Adults are 9 minor importance. to 11 mm. long. Antennae are Blaesoxipha coloradensis dark brown and reach three- (Aldrich) fourths the distance to the vibris- sae. Aristae are of medium length (Sarcophaga) 1916: 139 and plumose to the middle. There Synonymy.—Sarcophaga col- are three rows of black hairs be- oradensis Aldrich, Blaesoxipho- hind the eye, and the beard is theca coloradensis (Aldrich), B. abundant and white. Five to seven caudata Townsend {320, p. 947). - stripes may be seen on the dorsal Recorded distribution.—Brit- part of the thorax when viewed ish Columbia, Alberta, Saskatche- from behind. There are three wan, Manitoba, California, Colo- postsutural dorsocentrals and rado, Texas, and Massachusetts three pairs of anterior acrosti- {320, p. 947). (Map 10.) chals. The abdomen is black and Host species.—Aeropedellus tessellated with gray pollen with clavatus (Thomas), Camnula pel- an indication of three dark lucida (Scudder), Melanoplus bi- stripes. Hypopygium is large. The vittatus (Say), M. dawsoni first segment is reddish brown (Scudder), M. femurrubrum (De- and somewhat pruinose. It sup- Geer), M. gladstoni Scudder, M. ports a row of small stout bristles infantilis Scudder, M. packardii ARTHROPOD AND NEMATODE PARASITES IN AMERICA 37

the dying host and into the soil where they either pupate within several days or enter diapause re- quiring about 46 days at 8° C. to break. Description.—Eggs unknown. Larvae (first instar) are about 1.6 mm. long by 0.45 mm. wide and are ivory white. Clumps of long spines several rows wide are lo- cated on the posterior lateral parts of segments 4 to 10; the MAP 10.—Recorded distribution of longer are located anteriorly in Blaesoxipha coloradensis (Aldrich). the group. Each of the first four segments are completely encircled Scudder, and M. sanguinipes with spines on the anterior mar- (Fabricius) (38, p. 6 ; 123, p. i8i ; gins, whereas the dorsal parts of 306, pp. 29-33; 307, p. 223; 318, segments 5 to 11 are free of pp. 30, 32). spines. There are 15 to 20 fine Incidence of parasitism.— spines on the floor of the posterior Smith (307, p. 223) reported that cavity and five coarse ones at the parasitism rarely exceeds 15 per- tip of each clawlike process in the cent, although the highest per- lateral part of the cavity. Sur- centage recorded was 48.3 in M. rounding the anal opening are sanguinipes in Canada. also several spines. Two sickel- Life history and habits.—SO: shaped oral hooks curve down- Eggs, larvae, and pupae unknown. ward and outward toward the Adults in Canada first appear tips, and infrahypostomal bridge early in June. LOD: Unknown. articulates with the hypostomal OWS : As larvae in the soil. G/Y : sclerite. Anal spiracles are sharp Two in Canada. SA: All stages, tipped and two lobed; the outer although instars 2 to 5 are most lobes are nearly twice as long as often attacked. N/H: Unknown. the inner. (309, pp. 92-9^) MOA : Unknown ; young deposited Pupae unknown. as larvae. AF and HAB : Un- Adults: Antennae are dark known. brown or blackish with the third Smith (307, pp. 222-223) noted joint about 11^ times that of the that a female may contain up to second. Palpi are rather long and 50 larvae, which are deposited on black, and the proboscis is short the host. The larvae burrow into and fleshy. The thorax is gray the hemocoel of the host and pass pollinose and displays the usual through the usual three instar three stripes. Anterior acrosti- stages. They then burrow out of chals are well developed and the 38 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

dorsosutural centrals number wan, Manitobia, and most of three. The abdomen is gray poUi- United States north of a line from nose and tessellated; it has a southern California, through rather distinct blackish line. Hy- northern Texas, and through popygium of the male is entirely northern South Carolina (5, pp. yellow and shiny; the first seg- 137-138; 38, p. 6; 221, pp. 724- ment is medium size and the sec- 728; 320, p. 9A7). (Map 11.) ond is globose with numerous Host species.—Camnula pellu- coarse black hairs. Forceps are cida (Scudder), Melanoplus dev- reddish yellow with black at the astator Scudder, M. sanguinipes apex and sickle shaped. Legs are (F.), and Oedaleonotus enigma black and the hindtibiae lack vil- (Scudder) (5, pp. 137-138; 38, p. losity. On the outer front side of 6; 123, pp. Jf6A-Jt8i; 126, p. 239; the middle femur is a row of four 168, p. 148;221, pp. 72U-728; 307, or five bristles, and two bristles pp. 231-232; 318, pp. 30-32; 325, are on the outer front side of the p. 13; 326, p. 25). middle tibia. The genital segment Incidence of parasitism.—Ee- of the female is red and bears corded parasitism in California a heavy row of bristles. A slen- has reached 20 to 26 percent, der, round, sharp larvipositor is which wiped out a grasshopper grooved behind and curved some- population of 20 per square yard what forward. Legs are bristly in 2 months, whereas Canada has and the femora are uncommonly recorded a 4-percent level (221, stout. (5, pp. 139-li.O) pp. 724-728). Discussion.—Smith (307, p. Life history and habits:—SO: 223) indicated that although the Eggs unknown. Larvae were incidence of parasitism in Canada reared from Canadian grass- is low, this is a relatively per- hoppers between mid-July and sistent species and is therefore an important early-season parasite. Since mortality within its hosts is high, the populations cannot in- crease as rapidly as some other grasshopper parasites.

Blaesoxipha falciformis (Aldrich) (Sarcaphaga) 1916: 137-138 Synonymy.—Sarcophaga falci- formis Aldrich, Acridiophaga fal- ciformis (Aldrich) (320, p. 9i7). Recorded distribution.—Brit- MAP 11.—Recorded distribution of ish Columbia, Alberta, Saskatche- Blaesoxipha falciformis (Aldrich). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 39 mid-September. Pupae unknown. cated, Middlekauff {221, pp. 72^- Adults in Canada occur from 728) presents excellent drawings. early June to early September. Pupae unknown. Total life cycle normally requires Adult flies are 51^ to 8 mm. 30 to 60 days. LOD : Eggs hatch long. Males are distinguished by within the uterus. Larvae 9 to 10 dark-brown to blackish antennae days. Pupae 8 to 10 days. Adults and the third joints are slightly unknown. OWS : Not given. G/Y : less than twice as long as the Two to three in California. SA: second. Aristae are plumose for Generally the adult with some slightly more than half their fifth-instar njrmphs. N/H: Not length. Palpi are rather long and given, MOA: Host is attacked the proboscis is short and fleshy. both in the air and on the ground. On the back of the head are three Larvae are inserted through the rows of black hairs. Vertical tough integument of the femur bristles are absent. The thorax is into the muscle tissue. AF: Un- gray pollinose with the usual known. Middlekauff {221) was three to five stripes. Anterior able to keep adults alive 8 days acrostichals are well developed, on honey and yeast in the labora- and there are three postsutural tory. HAB : Can be seen on grass dorsocentrals. The abdomen is a and dirt clods just prior to attack. gray pollinose and tessellated; it Middlekauff {221, pp. 72^-728) has a distinct median blackish found from seven to 47 eggs or line. On the black fourth segment larvae in each of 10 dissected wild is a row of more than a dozen females (average 22.1). He was bristles. Hypopygium of the male unable to determine how many is entirely yellow and shiny. The had already been deposited. Fe- first segment is medium size and male flies attack grasshoppers on has a row of bristles across the the ground generally by inserting hind margin, whereas the second their sharp larvipositors directly segment is globose and has nu- into the muscles of the femur of merous erect, coarse black hairs. the host. Here the larvae remain Sickle-shaped forceps are yellow for several days before migrating at the base and grade into black to the hemocoel for 5 or 6 days at the tip. Legs are black and the while they complete development. hindtibiae lack villosity. There is They then exit from the host, a short comb on the posterior edge usually through the neck mem- of the middle femur, and two brane, burrow into the ground, bristles stand on the outer front and pupate several days later. side of the middle tibia. Wings {307, pp. 231-232) are hyaline and have small costal Description.—Eggs hatch in spines. (5, pp. 137-138) the uterus. Larvae: Although no Discussion.—I know of no other formal larval description was lo- grasshopper parasite that inserts 40 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE a larva into the muscles of the Eggs, larvae, and pupae unknown. femur. Since this larva spends Adults not given. LOD, OWS, =■ about two-sevenths of its life in G/Y, SA, N/H, MOA, AF, and ^ the femur of the host, it can be HAB : Unknown. overlooked by this percentage in Description.—Eggs, larvae, and * hemocoel examination and thus pupae unknown. Adults: Aldrich ^ lower the observed incidence of (5) described this species as a parasitism. The observation by variety of B. (=S.) aculeata and Middlekauff (221) indicates that stated that the only difference be- , in limited areas this insect can be tween the two is in the male an effective parasite. genitalia. In B. gavia, "the lateral -^ lobes of the distal segment of the . Blaesoxipha gavia (Aldrich) penis are much larger, heavily (Sarcophaga) 1916: 145 chitinized, and spinose on the Synonymy.—Sarcophaga acu- outerside; while the apical proc- : leata var. gavia Aldrich (320, p. esses are stout and much more 9U). strongly divergent at their base ^ Recorded distribution.—Brit- than in aculeata, so that they form ish Columbia, New Mexico, Kan- a U rather than a V." Palpi are sas, and New Jersey (5, p. 1A5; black. On the thorax are three 320, p. 9U). (Map 12.) distinct postsutural dorsocentrals ' Host species.—Camnula pellu- and well-developed anterior acro- cida (Scudder) and Melanoplus stichals. Hindtibiae lack erect sanguinipes (Fabricius) (38, p. 6 ; villosity. The first vein of the 318, pp. 31-32). wing is bare. (5, p. US) Incidence of parasitism.—None Discussion.—It appears that located. the main reason for the sporadic Life history and habits.—SO: distributional records, low num- . ber of host species, and lack of incidence and figures is that this '■ species is very similar to B. . aculeata and many records may unknowingly be attributed to B. aculeata that belong to B. gavia. Blaesoxipha hunteri (Hough) (Sarcophaga) 1898: 207 Synonymy.—Sarcophaga hun- teri Hough (in Hunter (ISA)), Protodexia synthetica Townsend, S. aenigma Reinhard, P. hunteri (Hough) (A, p. 572; 320, p. H5). MAP 12.—Recorded distribution of Blaesoxipha gavia (Aldrich). Recorded distribution.—South- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 41 em Canada and the United States {320, p. H5). (Map 13.) Host species.—Aeropedellus clavatus (Thomas), Ageneotettix deorum (Scudder), Amphitomus coloradus (Thomas), Aulocara el- liotti Thomas, Camnula pellucida (Scudder), Chorotphaga viridi- fasciata (DeGeer), Cordillacris occipitalis (Thomas), Melanoplus angustipennis (Dodge) M. bivat- tatus (Say), M. bruneri Scudder, MAP 13.—Recorded distribution of M. confusus Scudder, M. dawsoni Blaesoxipha hunteri (Hough). (Scudder), M. differentialis (Thomas), M. femurrubrum (De- preferred hosts, although many Geer), M. foedus Scudder, M. in- other species are often heavily fantilis Scudder, M. marginatus attacked. Many larvae infecting (Scudder), M. packardii Scudder, nymphs of M. bivittatus survive, M. sanguinipes (Fabricius), Met- whereas nearly all that infect ator pardalinus (Saussure), adults die within the host early Pklibostroma quadrimaculatum in life. (5, pp. 102-105; 183, p. (Thomas), Psoloessa delicatula 9) (Scudder), and Trackyrhachys Life history and habits.—SO: kiowa (Thomas) of the order Eggs, larvae, and pupae spring to Orthoptera and several species of fall.Adults first appear in Canada Lepidoptera (5, pp. 102-105; 38, about early June and in Montana p. 6; 123, pp. U76, Jk85; 12U, pp. and North Dakota several weeks U, 10; 16Jf, p. Ul; 168, pp. U7- later. LOD: Gestation requires U8; 183, p. 9; 22A, p. 25; 281, p. about 9 days. Eggs hatch in the 83 ; 306, pp. 28-33 ; 307, pp. 229- uterus. Larvae mature in about 231; 309, pp. 86, 96-97; 322, p. 10 days. Pupae 14 to 15 days. 313; 323, p. 118; 325, pp. 5, 7, 8, Adults unknown. OWS: As larvae H; 326, p. 26; 336, pp. 31, 33; in soil. G/Y: Two to three in 380; 389). Canada. SA: Adults and fifth- Incidence of parasitism.—The instar nymphs mostly, with rare highest parasitism recorded in attacks on the fourth instars. Canada was 36.1 percent in M. N/H : Generally one, although two packardii. In Montana it has are possible. MOA : Approach the reached slightly over 40 percent host on the ground, deposit dorso- on several occasions in M. sangui- ventrally flattened chorin-covered nipes. In both Canada and the larvae in the intersegmental body United States M. packardii and folds of the host. AF and HAB : M. sanguinipes are generally the Unknown. 42 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Mating may occur anytime were unable to show any feeding after emergence, followed by a 10- on grasshopper eggs, even when to 14-day gestation period. During larvae were placed in close associ- this time the adult flies can be ation with freshly deposited egg found among the grasshopers, al- pods. though they are indifferent to Description.—Eggs are small, their activities. Each female pro- 0.7 mm. long by 0.17 mm. wide, duces on an average 34 eggs per translucent white, and tapering litter with a maximum of 88. toward both ends from the middle These develop at varying rates (389). Larvae (first instar) are 1 into larvae while still in the egg mm. long by 0.15 mm. wide, trans- sac. lucent white, and dorsoventrally The female fly attacks its vic- flattened. Spines are very small tim on the ground, generally in a and inconspicuous on the body but stalking manner. When within more conspicuous on the floor of striking range, the fly brings the the posterior cavity where there tip of the abdomen forward either are two rows of nine or more between its legs or along one of its spines. A few coarse spines also sides and places a larva on the surround the anal opening. side of the grasshopper, usually Heavily pigmented platelike spots within one of the body folds. It are located on the mid-dorsal sur- quickly enters the host's body face on pleural areas of segments through the softer parts of the 9 and 10, often with a black basal integument and remains there for ring. Segments 6 to 9 are devoid about 8 to 9 days while passing of spines on the pleural regions. through the usual three larval Oral hooks are scalpel-like in pro- stages. When mature the larva file and the infrahypostomal emerges from the host and bur- bridge articulates with the hypos- rows into the soil where it either tomal sclerites. Within the pos- pupates several days later or en- terior cavity is a pair of single- ters diapause. Those that pupate lobed, sharp spiracles. There is no become adults in about 15 days, clawlike process. (309, pp. 86, and those in diapause require a 96-97) temperature of 8° C. for 50 days Pupae are 5.25 mm. long by 2.0 before proceeding with develop- mm. wide and dull yellowish red. ment. (183, p. 9; 307, pp. 229- Tubercles are not visible on the 231 \ 323, p. 118)389) edge of the posterior cavity or Treherne and Buckell {336, pp. around the anal opening. Anterior 31, 33; 389) reported the only spiracles have nine small yellow case of larvae of B. hunteri being lobes on a deep-reddish base. reared from grasshopper egg Within the posterior cavity is a pods. In tests at the Bozeman pair of dark-red spiracular plates. laboratory, however, workers Each plate consists of three dis- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 43 tinct lobes and in each lobe is a males. The frontal stripe is yellow yellow spiracular slit somewhat or brassy and the antennae are broader at the base than the almost yellow. Stripes on the pointed apex. The anal opening is light-gray thorax are hardly dis- small and depressed. (124^, pp. U, tinguishable. The abdomen is also 10) light gray with a median indistinct Adult males are 6 to 7 mm. stripe. The genital segment is re- long. Antennae are reddish brown. tracted and not visible, but the The third joints are slender and orifice is closed by a distinct twice as long as the second. larvipositor, which is spatulate or Aristae are long and plumose for duckbilled at the apex. There are slightly more than half their also two bristles on the middle length. Palpi are yellow and the tibia, one smaller than the other. proboscis is brown. On the back (Í5^, pp. 207-210) of the head are three rows of black Discussion.—Although B. hurt- hairs. Outer vertical bristles are teri is apparently of little impor- absent. The thorax is gray polli- tance in Canada, it is one of the nose and has the usual three to most common parasites in Mon- five black stripes. There are two tana. Since only one record exists, pairs of anterior acrostichals but its role as an egg predator is only three postsutural dorsocen- questionable but not impossible. trals. A distinct median black line Prematurely emerged larvae may and a changeable lateral line on use eggs to supply enough nutri- each side run along the dorsal tion to complete development, part of the whitish-gray abdomen. although I doubt whether mature The genitalia are minute and larvae seek additional food. black. Forceps are slender, yel- low, delicate, and squarish at the Blaesoxipha impar (Aldrich) tip. Possibly one of the most (Sarcophaga) 1916: 220-222 characteristic features of this Synonymy.—Sarcophaga im- species is the tuft of hair that par Aldrich (320, p. 9^6). emerges from the forceps and that Recorded distribution.—South- can be seen even when the geni- ern States from New Mexico east talia are in repose. to include Florida and north to Legs are black and the hind- include Nebraska and Iowa (5, p. tibiae lack villosity. The middle 222; 320, p. H6). (Map 14.) tibia has only one bristle on the Host species.—"Lubber grass- outer front side, whereas a few hoppers" (5, p. 222). stout spines are located on the Incidence of parasitism.—None middle femur. Wings are subhy- located. aline and lack costal spines. Fe- Life history and habits.—SO: males measure about 5 mm. and Eggs, larvae, and pupae unknown. are more ashy pollinose than the Adults not given. LOD : Unknown. 44 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE each side. {167, pp. 438-^39) Pupae unknown. _;, Adults measure 8 to 10 mm. long. Antennae are brownish black. The third joints are twice - as large as the second. Aristae are plumose for slightly more than half their length. Palpi and pro- * boscis are ordinary and black. Three rows of black hairs stand along the back of the head as do -s the outer vertical bristles, which MAP 14.—Recorded distribution of are not well developed. The tho- Blaesoxipha impar (Aldrich). rax is gray pollinose, slightly i yellowish tinged, and on which are the usual three to five black OWS, G/Y, SA, N/H, and MOA: stripes. Although there are four Not given. AF: Aldrich (5) postsutural dorsocentrals, the an- reared flies on "food, beer, and terior two are small. Anterior meat." HAB : Unknown. (5, pp. acrostichals are distinct. The ab- 220-222.) domen is gray pollinose and tes- , Description.—Eggs unknown. sellated. It has three distinct, Larvae (mature) are slightly changeable stripes. The fourth ' more than 3 mm. long and are segment is reddish and gives rise heavily armed similar to Sarco- to a marginal row of about a phaga sarracenioides Aldrich in dozen bristles along the hind edge. form. The anterior borders of First segment of the male hypo- segments 2 to 12 are completely pygium is large, broad, and encircled with spines, which be- pollinose, with a yellow to dark- come more widely spaced and brown ground and with a row of - reduced in size on segments 11 about eight stout bristles. The and 12. The posterior borders of second segment is red, globose, segments 5 to 12 have complete and slightly pollinose. It has bands of spines, although reduced erect black hairs. Probably the on segment 6 and especially 5. most characteristic structure of The posterior cavity is small and this species is the forceps. Front surrounded by two pairs of small and back edges are parallel in tubercles on the upper border and profile, whereas near the tip they two pairs (one weakly defined) suddenly bend at almost right on the lower. Posterior spiracles angles. Legs are black. On the lack a peritreme. Anal area is outer front side of the middle small and flanked with two lat- tibia are two bristles, and on the erally directed tubercles, one on hindtibia are a few erect hairs ARTHROPOD AND NEMATODE PARASITES IN AMERICA 45

on the inner and outer side. There (Scudder), M. packardii Scudder, is also a comb on the middle M. sanguinipes (Fabricius), and femur. Wings are hyaline. (5, M. spretus (Walsh). In addition to VP. 220-222) other orthopteran families, hosts Discussion.—Because of its include species of Coleóptera and limited host range and because the Lepidoptera. (5, pp. 263-265; host is of little economic impor- 37, pp. 115-116; 38, p. 6; 77, p. tance, this parasite has little value 12; 78, p. 98; 123, pp. ^6^-485; as a natural control agent. 132, p. 519; 136, pp. 116-117; 16i, pp. Í35-U6; 168, pp. U7-1A8; Blaesoxipha kellyi (Aldrich) 183, p. 9; 297, p. 22U; 299, p. 28; (Sarcophaga) 1914: 443 307, pp. 225-228; 318, pp. 30, 32, Synonymy.—Sarcophaga kellyi 3Í-35; 325, pp. 5, 7-9, li; 326, Aldrich (in Kelly (16^)), Kelly- p. 26; 380, pp. 291-306; 389) myia kellyi (Aldrich), S. veri- Incidence of parasitism.—B. cauda Coquillett (320, p. 9A6). kellyi is able to build up rapidly Recorded distribution.—South- and overtake host populations ern Canada west of Quebec and near the end of the season. This the United States west of the species is therefore credited with Atlantic Coast States {320, p. the elimination of many grass- 9i6). (Map 15.) hopper populations of economic Host species.—Camnula pellu- concern. Recorded percentages, cida (Scudder), vi- however, were not located. (307, ridifasciata (DeGeer), pp. 225-228) Carolina (Linnaeus), D. longipen- Life history and habits.—SO: nis (Thomas), Melanoplus bivit- Eggs, larvae, and pupae late tatus (Say), M. differentialis spring to fall. Adults may be (Thomas), M. marginatus found in the field in Canada any- time after June and several weeks earlier in more southern areas. The first killing frost terminates their presence. LOD: Gestation generally requires 13 to 19 days although a range of 8 to 60 days has been recorded. Eggs mature and hatch in the uterus. Larvae mature in 4 to 5 days. Pupae 11 days. Adults: Smith (307) re- ported an adult female 90 days old that produced viable larvae. Under controlled laboratory con- MAP 15.—Recorded distribution of ditions adults have lived as long Blaesoxipha kellyi (Aldrich). as 150 days. OWS : Larvae in the 46 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE soil. G/Y: From two to four de- has paired oral hooks and an artic- pending apparently on location, ulated infrahypostomal bridge. climate, and availability of hosts. Upper and lower borders of the In addition, as many as five litters posterior cavity each contain two are produced per female. SA: pairs of small tubercles ; the inner Nearly all. N/H: Generally only pair on the lower border is small one. MOA: B. kellyi will attack and inconspicuous. Within the almost anything moving in the air cavity is a pair of large spiracles. from a host to a thrown dirt clod. The upper end of the spiracular Also it sometimes deposits larvae slits end in a spinelike point, on stationary or dead hosts. AF: which is lighter in color and Unknown. HAB : Grasslands near difficult to see. Clawlike processes grasshopper outbreaks. and spines on the floor are absent. Mating may occur anjrtime (167, pp. U9, JÍ25, U2\ 309, pp. after the fly becomes an adult. 86, 98) Smith (307) found that in the lab- Pupae are 8 mm. long by 2.5 oratory a female may produce up mm. wide and dull dark red. to five litters with 7-day intervals Tubercles on the outer side edge between litters when food was in are indistinct and anal tubercles good supply. When the host is in are absent. Anterior spiracles fiight, the larva is generally near the cephalad end on the placed on the underside of its puparium each contain six small wings, often near the base. The yellow lobes on a reddish-brown parasite quickly penetrates the base. Within the large nearly integument and completes its round posterior cavity is a pair larval development in about 3 of deep reddish-black spiracular days within the host's hemocoel. plates, each one containing three (307, pp. 225-228; 389) narrow yellow slits. The middle Description.—Eggs unknown. slits are nearly parallel with each Larvae (first instar) are 2.5 to other. Around each plate is an 2.7 mm. long by 0.7 mm. wide, extension that nearly encircles it translucent white, and tapering and a wrinkled area on the inner toward the anterior end. Larvae lower edge. The anal opening is are not heavily armed. The an- small, depressed, and surrounded terior borders of segments 2 to 7 on each side by a depressed area. have a complete band of spines (167, pp. U19, k25, U2) as do the posterior borders of seg- Adult length ranges from 6.5 ments 9 to 11. There is a broad to 10 mm. Antennae are black. ventral band on segments 4 to The third joints are twice as long 11 with a transverse spineless as the second. Aristae are plumose area in the center. On segment for half their lenert:h. Both palpi 2 the spines are mostly furcate. and proboscis are black. There are The buccopharyngeal armature three distinct rows of black hairs ARTHROPOD AND NEMATODE PARASITES IN AMERICA 47 on the back of the head. The ing other Orthoptera and species grayish pollinose thorax has the of Coleóptera and Lepidoptera. usual three to five black stripes. Third, it is able to survive in a Anterior acrostichals are well de- dead host. Aldrich (5) reported veloped, and there are four large it from dead grasshoppers and postsutural dorsocentrals. On the Smith (307) reared it on dead tessellated whitish-gray pollinose grasshoppers. In 1924 C. C. Wil- abdomen are three distinct longi- son after a poisoning experiment tudinal black stripes. The fourth collected many dead grasshoppers, abdominal segment contains a which yielded larvae that were marginal row of about 10 lateral reared to maturity for identifica- bristles. The fifth sternite is red- tion. B, kellyi is itself not without dish brown and deeply incised in its own group of parasites and the middle. The first segment of predators. Recorded enemies are the hypopygium is black, polli- the hyperparasites Aphaereta sar- nose, and tessellated. It has a row cophagae Gahan (Braconidae) of stout bristles on the posterior and Eupteromalus sarcopfiaga margin. The second segment is Gahan (Pteromalidae) and the yellow, globose, and not bristly. predator Cyrptogogon wilMstonia Forceps are long, slender, and Curran ( Asilidae). (307 ; 323, p. black with a brown base. The 117; 325, p. 26; 389) outer side of the forceps has nu- merous short, stubby, erect hook- Blaesoxipha opifera CCoquillett) lets except for the basal third. (Sarcophaga) 1892: 22-24 Legs are black and hindtibiae Synonymy.—Sarcophaga opi- lack villosity. On the middle fera Coquillett, Opsophyto opifera femur is a short comb and on the (Coquillett) (Í, p. 513; 320, p. middle tibia are two bristles on H5). the outer front side. Wings are Recorded distribution.—Brit- hyaline and lack costal spines. ish Columbia, Alberta, Saskatche- The genital segment of the female wan, Manitoba, and most of the is red and the orifice is transverse. United States west of the Missis- There are two distinct concavities sippi River (307, pp. 228-229; side by side on the red genital 320,p.9í5). (Map 16.) tergite. The middle femur of the Host species.—Amphitornus female lacks the comb. (5, pp, coloradus (Thomas), Camnula 263-265; 16JÍ, pp, Í35-U6) pellucida (Scudder), Melanoplvs Discussion.—B. kellyi is un- hivittatus (Say), M. confiisus usual in several ways. First, it has Scudder, M. dawsoni (Scudder), two to four generations per year M. devastator Scudder, M. differ- with up to five litters per female. entialis Thomas, M. femurrubrum Second, it attacks many hosts (DeGeer), M. marginatus other than the Acrididae, includ- (Scudder), M. plumbeus (Dodge), 48 TECHNICAL BULLETIN NO. 14<50, U. S. DEPT. OF AGRICULTURE stages. N/H: Two to three can live successfully within one host at the same time. MOA : Attacks stationary hosts. AF and HAB: Unknown. Mating occurs anytime after eclosión. Each female produces as many as 57 larvae. Attack on the host is accomplished in several ways, each method requiring that the host is quiet during the at- tack. If the ñy is facing toward MAP 16.—Recorded distribution of the host, it flips its ovipositor Blaesoxipha opifera (Coquillett). forward and under or along either side of the thorax and thus hurls and M. sanguinipes (Fabricius) the larva at the grasshopper (5, pp. 119-121; 6i, pp. 22-2Í; where it sticks upon contact. If 123, p. Â85; 306, pp. 28-33; 307, facing away, the fly brings its pp. 228-229; 325, pp. 7, H; 326, abdomen between its legs and p. 21 ; 336, p. 31 ; 380, p. 306 ; 389). flips it back, also hurling the larva In addition, Treherne and Buck- at the grasshopper. A third ell (336, p. 31 ) reported that the method entails the fly pouncing larvae of B. opifera attack the on the back of the grasshopper, eggs of C. pellucida. dropping the larva, and quickly Incidence of parasitism.—Ca- jumping away. While the grass- nadian references indicate 40 hopper attempts to rub the area percent of M. sanguinipes and 5 touched by the fly, it neglects percent of C. pellucida are para- that area where the larva is bur- sitized in British Columbia. In rowing. Montana up to 27 percent of M. B. opifera completes the usual sanguinipes has been recorded. three larval stages within the (307, pp. 228-229; 389) host, emerges, and burrows into Life history and habits.—SO: the ground where it pupates Eggs, larvae, and pupae late within several days or enters dia- spring to fall. Adults can be found pause. Smith (307) discovered in the field in Canada and Mon- that a temperature of 8° C. is tana from about mid-June to mid- needed for 77 days in order to September. LOD: Gestation 8 to break diapause. (307, pp. 228- 9 days. Eggs are hatched in the 229; 389) uterus. Larvae 7 to 9 days. Pupae Description.—Eggs (nonferti- 12 to 13 days. Adults unknown. lized) are 0.45 mm. long by 0.16 OWS : As larvae in the soil. G/Y : mm. wide, translucent white, and Two to three in Canada. SA : All tapering toward both ends from ARTHROPOD AND NEMATODE PARASITES IN AMERICA 49 the middle. The chorion often ap- joint thickened on its basal two- pears sculptured. (389) Larvae fifths. Proboscis is short and (first instar) are about 1.3 mm. brown to blackish and the palpi long by 0.5 mm. wide and trans- are yellow to yellowish brown. lucent to ivory white. The body is Thorax is light gray and has five without spines except for the to seven black stripes. There are anterior margin of the first tho- three postsutural dorsocentrals, racic segment. There are two or and the anterior acrostichals are three apical spines on each of the strong. The abdomen is densely two clawlike processes in the gray pollinose with changeable posterior cavity and several reflections and three rather con- around the anal opening. The stant blackish stripes. The fourth infrahypostomal bridge articu- segment contains about a dozen lates with the hypostomal sclerite. hairs in a row. Hypopygium is The two fine oral hooks are small small, red, and retracted. The first and sicklelike and curve gently segment is pollinose and the sec- downward. The posterior cavity ond two are red and support is shallow and contains a pair of coarse hairs. Forceps are red ex- two-lobed, sharp-tipped spiracles, cept on the extreme tips. The the outer lobes of which are about distal part is bare, curved for- twice as long as the inner. (309, ward, and ends in a small point. pp. 86, 96) Accessory plate tapers to a Pupae are 5 to 6.5 mm. long slender tip and is yellow. Basal by 1.5 to 2 mm. wide and dull segment of the penis is yellow and yellow to red. The pupal case is the distal blackish. not wrinkled, and the anal tuber- Legs are brown to black and cles are absent. Each anterior without villosity. Coquillett (6JÍ) spiracle contains six well-sepa- stated that all femora and tibiae rated yellow lobes on a dull dark- are bristly. There is a comb near red base. Within the small ellipti- the tip on the hind side of the mid- cal posterior cavity are two dle femora and two bristles on the deep-reddish spiracular plates on outer front side of the middle which are three short and nearly tibia. Wings are hyaline and lack straight spiracular slits. (12Jí, costal spines. pp. A, 9-^12) Females diflfer from males in Adult females are 5i/^ to 61^ having the thorax and the abdo- mm. long. Antennae are red to men more densely pollinose, the dark brown. The second joints are stripes more faint on the thorax nearly as long as the third. and nearly absent on the abdo- Aristae are black, plumose for men, and two to four orbitals. two-thirds their length, and two- The fourth abdominal segment is jointed, with the basal joint as red at the tip and the genital broad as it is long and the second segment is all red. They lack a 50 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE larvipositor, which is found in liotti (Thomas), Camnula pellu- many of this group. The last cida (Scudder), Chorthippus Ion- - sternite is oval and hairy. (6i, gicornus (Latreille), Melanopltis , pp. 22-2A) angustipennis (Dodge), M. bivit- Discussion.—This species can tatus (Say), M. confusiis Scudder, ~ often be found with other Blaesox- M. dawsoni (Scudder), M. differ- ipha species and several tachinids entialis (Thomas), M. femurru- attacking grasshoppers. Its peak brum (DeGeer). M. foedus * population periods are often offset Scudder, M. infantilis Scudder, - by those of other species so that M. packardii Scudder, M. san- the host species feel a steady guinipes (Fabricius), M. seudderi ^ pressure of parasitism rather (Uhler), Orchelimum sp. (family than a fluctuating one. This is Tettigoniidae), and Phoetaliotes another effective controlling nebrascensis (Thomas) (5, pp. * agent. 135-136; 38, p. 6; 123, p. Í85; ■- 132, p. 521; 306, pp. 28-33; 307, Blaesoxipha reversa (Aldrich) pp. 233-235; 318, pp. 31-32, 3A; ' (Sarcophaga) 1916: 135-136 389). Synonymy.—Sarcophaga re- Incidence of parasitism.—The versa Aldrich (320, p. 9i5). highest parasitism recorded, ac- ' Recorded distribution.—South- cording to Smith (307), was 47.3 ern Canada and northern United percent of M. bivittatus in Can- States above and including Cali- ada. He also declared this species fornia, Colorado, Missouri, and to be one of the most important Massachusetts (320, p. 9-Í5). of the sarcophagid parasites of (Map 17.) grasshoppers in western Canada Host species.—Aeropedellus (307, pp. 23Jf-235). clavatus (Thomas), Ageneotettix Life history and habits.—SO: deorum (Scudder), Aulocara eU Eggs, larvae, and pupae late spring to fall. Adults appear in ' the field in Canada early in June and remain until September. LOD : Gestation requires about 9 * days. Eggs hatch in the uterus. , Larvae mature in 14 to 15 days. Pupae mature in 11 days. Adults live up to 60 days in the labora- tory. OWS : As larvae in the soil. G/Y: Two to three per year in Canada and Montana. SA: AU stages. N/H : Usually one to two, MAP 17.—Recorded distribution of sometimes three. MOA: The fe- Blaesoxipha reversa (Aldrich). male either alights on or runs ARTHROPOD AND NEMATODE PARASITES IN AMERICA 51 over a stationary grasshopper, segments 3 to 10. Several short depositing one or two larvae, or spines surround the anal opening. attacks flying grasshoppers and Located on the ventral surface of deposits a larva on the dorsal part the posterior segment are two of the abdomen, usually near the pairs of protuberances, the upper tympanum. AF: This species pair of which is longer than the feeds on the honeydew from more rounded lower pair. Oral aphids, which infest Artemisia hooks are sickle shaped and the cona Pursh in the field, and sugar infrahypostomal bridge appears cubes in the laboratory. HAB: to be fused with the hypostomal Not located. In Montana it can sclerite. The strong pair of oral be found on rangeland. hooks curves downward and out- Mating generally occurs soon ward, terminating in sharp tips. after eclosión. The average num- A pair of sharp-tipped, two-lobed ber of eggs per female fly is about spiracles is within the posterior 44, although 80 were found in one cavity, the outermost lobes of specimen. After deposition larvae which are the longest. (306, pp. enter the hemocoel through the 85-87) Pupae unknown. integument and pass through the Adult flies are 7 to 9 mm. long. usual three instars. Upon emer- Antennae are dark brown or gence from the host, they burrow blackish. The third joints are into the soil for several days and nearly double the length of the either pupate or enter diapause. second. Aristae are plumose for Smith (307) found that an expo- slightly more than half their sure of 8° C. for 67 days would length. Proboscis is fleshy and the break diapause and that exposure palpi are black. The thorax is gray to this temperature for 200 days pollinose and displays the usual produced no apparent harmful three to five stripes. Anterior effects. (307, pp. 233-^239; 389) acrostichals are well developed Description.—Eggs are translu- and there are only three distinct cent white and taper toward both postsutural dorsocentrals. The ends (389). Larvae (first instar) abdomen is gray pollinose and tes- are 2.0 mm. long by 0.5 mm. wide sellate and has a rather distinct and translucent white ; they have median blackish line. The fourth the normal tapering shape. Only segment is red at the apex and the first five segments bear spines contains a row of more than a on the dorsal half. The ventral dozen bristles. Hypopygium is parts of all segments, except the entirely yellow and shiny, the first second, have dark spines. The segment medium sized, and the band of spines on the underside second globose. Forceps are red of segments 5 to 11 are divided to the apex and contain fine hair transversely and the bristles are around the base. scattered on the pleural areas of Legs are black and the hind- 52 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE tibiae lack villosity. There is a short comb below the posterior edge of the middle femur and there are three bristles on the outer front side of the middle tibia. Females are much grayer than the males, and their abdomens are strongly tessellated. The oviposi- tor is red, elongate, straight, and trough shaped. (5, pp. 135-136) Discussion.—Although M. bi- vittatus is one of the preferred MAP 18.—Recorded distribution of hosts, B. reversa suffers a high Blaesoxipha taediosa (Aldrich). degree of mortality within it, sometimes reaching 60 percent (307, p. 235). Stage of develop- Eggs, larvae, and pupae unknown. ment of the host seems to aifect Adults not given. LOD : Unknown. survival. Larvae entering third-, OWS, G/Y, SA, N/H, and MOA: fourth-, and fifth-instar nymphs Not given. AF : Unknown. HAB : and very young adults generally Not given. survive and eventually kill their Description.—Eggs, larvae, and hosts. Those entering adult hosts pupae unknown. Adult flies are die, whereas the host lives. B. 6 to 9 mm. long. Antennae are reversa is one of the more efficient dark brown to blackish. The third grasshopper parasites. joints are nearly double the length of the second. Aristae are plumose Blaesoxipha taediosa (Aldrich) for slightly more than half their (Sarcophaga) 1916: 145 length. Palpi are long and the Synonymy.—Sarcophaga acu- proboscis is short and fleshy. leata var. taediosa Aldrich (320, Outer vertical bristles are absent. p. 9U). The usual three to five stripes are Recorded distribution.—Brit- located on the gray poUinose tho- ish Columbia, Wyoming, South rax. Anterior acrostichals are well Dakota, Kansas, and New Mexico developed and there are only three (320, p. 9U). (Map 18.) postsutural dorsocentrals. The Host species.—Camnula pellu- abdomen is gray pollinose and cida (Scudder) and Melanoplus tessellated; it has a distinct me- sanguinipes (Fabricius) (38, p. 6; dian blackish line. The fourth 123,p..í8i;318,p. 37). segment is entirely black and has Incidence of parasitism.—None a row or more than 12 bristles. located. Hypopygium of the male is en- Life history and habits.—SO: tirely yellow. The first segment ARTHROPOD AND NEMATODE PARASITES IN AMERICA 53 supports a row of bristles across the hind margin and the second segment bears numerous erect, coarse black hairs. Forceps are thin. Legs are black and the hind- tibiae lack villosity. The middle femur supports a short comb on the posterior edge, and the middle tibia has two bristles on the outer front side. Wings are hyaline and have small costal spines. (5, p. U5)

Discussion.—B. taediosa very MAP 19.—Recorded distribution of closely resembles B. aculeata and Blaesoxipha uncata (Wulp). was described by Aldrich (5) as a variety of B. aculeata. This similarity probably accounts for N/H, MOA, AF, and HAB: Not the scattered collection records given. (389). and lack of additional informa- Description.—Eggs and larvae tion. Its effectiveness as a grass- unknown. Pupae are medium hopper controlling agent is un- sized, 7 to 7.5 mm. long by 3.0 known. mm. wide, yellowish red, and slightly arched along the dorsum. Blaesoxipha uncata (Wulp) There are no outstanding tuber- (Sarcophaga) 1895: 268-269 cles on the edge of the posterior Synonymy.—Sarcophaga un- cavity or around the anal opening. cata Wulp; S. marginata Aldrich There is, however, a depression U,p. 5H; 320, p. 9i7). just above the posterior cavity. Recorded distribution.— Greene (124) did not observe any United States (320, p. 9^7). anterior spiracles. The posterior (Map 19.) cavity is medium sized, elliptical, Host species.—Dissosteira Car- and centrally located on the hori- olina (Linnaeus), Melanoplus bru- zontal axis. Within are a pair of neri (Scudder), shiny red spiracular plates, each haldemanii (Scudder), and Xan- with three yellow slits, the middle thippus corallipes (Haldeman) and inner ones of which are (318,p.31;389). slightly convex to each other and Incidence of parasitism.—None of about equal length. The outer located. slit is straight and about two- Life history and habits.—SO: thirds as long as the one in the Eggs, larvae, and pupae unknown. middle. The anal opening is small Adults first of June in Montana. and indistinct. (124, pp. 5, 18- LOD : Unknown. OWS, G/Y, SA, 19) 54 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Adult flies are 6 to 8 mm. long. and contain small costal spines. Antennae are dark brown to (5, pp. 136-137; 381, pp. 265- " black. The third joints are twice 272) as long as the second. Aristae are Discussion.—Although widely plumose for a little over half their dispersed, B. uncata has been length. Palpi are black, long, studied very little, probably be- cylindrical, and with short hairs. cause of its generally low popula- Proboscis is black with brown tion in most areas. Its effective- terminal tip, short, and fleshy. ness as a grasshopper controlling 1 Three rows of black hairs line the agent is unknown. back of the head, although the outer vertical bristles are absent. Boettcheria cimbicis (Townsend) The thorax is gray pollinose and (Sarcophaga) 1892: 126 has the usual three to five stripes. Synonymy.—Boettcheria fer- " Anterior acrostichals are well naldi Parker, Sarcophaga cim- developed, and there are three bicis Townsend U, V- 511 ; 320, postsutural dorsocentrals. The p. 948). ■* abdomen is gray pollinose and Recorded distribution.—Wash- tessellated. It has a rather dis- ington, Idaho, California, and ., tinct median black line. The north-central and eastern United fourth abdominal segment is en- States (320, p. 948). (Map 20.) * tirely black. Hypopygium of the Host species.—Reared from the , male is entirely yellow. The first orthopteran host Chortophaga segment is medium sized and has viridifasciata DeGeer and many a row of bristles across the hind lepidopteran hosts (5, pp. 79-81 ; margin. The second segment is 164, p. 441 ; 318, p. 30 ; 325, p. 8 ; globose and has numerous erect, 326, p. 24). coarse black hairs. Forceps are entirely yellow or with only the extreme apex being slightly dark and are surrounded by black hair at the base. Beyond the middle on each side is a white fringe or ruffle, and on the back is a deep cut made by a transverse notch in which the color is whitish. Legs are black. There is a short comb below on the posterior edge of the middle femur, two bristles on the outer front side of the middle tibia, and the hindtibiae MAP 20.—Recorded distribution of lack villosity. Wings are hyaline Boettcheria cimbicis (Townsend). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 55

Incidence of parasitism.—None basal half. Palpi are stout and located. blackish ; proboscis is brown. The Life history and habits.—SO: thorax is silvery white with three Eggs, larvae^ and pupae unknown. heavy black stripes. There are Adults occur from May to June in only three postsutural dorsocen- South Dakota. LOD: Unknown. trals and the anterior acrostichals OWS : Not given. G/Y : Unknown. are well developed. The abdomen SA, N/H, MOA, and AF: Not is silvery to gray marble, about given. HAB : Unknown. (155, p. the same width as the thorax, and 33; 160, p. 252; 33i, pp. 126-127) has a fairly distinct median dark Description.—Eggs unknown. line. Hypopygium is red with the Larvae : B, cimbicis so closely re- first segment often dark at the sembles J?. latisterna Parker in base. The second segment is red the larval stage that Knipling subshining, globose, and without (167) only indicated the differ- bristles. Forceps are short, wide, ences between the two in describ- and support a tuft of very striking ing this species. First-instar long black hair at the basal half. larvae are about 3 mm. long and Fifth sternite contains two pad- taper slightly in both directions like structures that face forward from the middle. They are well at the base of the "V." armed with bands of spines that Legs are black and the femora cover the anterior borders of seg- silvery white. There are two en- ments 9 to 12. The spines are tire rows of bristles on the outer distinctly divided transversely on side of the hindfemora above the the ventral surface of segments middle part. Wings are grayish 5 to 7. Spines on segments 3 and hyaline and longer than the abdo- 4 are long and slender as they are men. Haltères are a tawny yellow. on the lateral and dorsal regions Females are 6 to 10 mm. long. of the other segments. The There is a changeable reñection cephalopharyngeal mechanism is spot along the base of the an- similar to that of B, latisterna but tennae. Acrostichals are well de- smaller, whereas the spiracular veloped and there are three slits of the posterior spiracles are postsutural dorsocentrals. Abdo- larger. (167, pp. U25, U2S, UUU, men is black except the fourth UU7-U5U) Pupae, unknown. segment, which is sometimes Adult males are 7 to 12 mm. slightly red as is the genital long. Antennae are blackish with segment. The genital opening is the third joints not quite as long nearly circular and fringed with as the second. Aristae are of uni- hairs, whereas the red sternite form color, three segmented, ex- below is distinctly ñat. (5, pp. tremely long, thickened on the 78-81 ; 167, pp. Í25, U28, UÍ, U7- basal third, and plumose on the U5U; 335, pp. 126-127) 56 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Discussion.—Only one ortho- Life history and habits.—SO: pteran host has been reported and Eggs, larvae, and pupae unknown. this was probably incidental. B. Adult flies are active in the ñeld cimbicis is apparently a parasite early in June in Michigan and primarily of Lepidoptera. remain through September, and early in July in California and Helicobia rapax (Walker) Washington. LOD: Eggs and (Sarcophaga) 1849: 818 larvae unknown. Pupae 6 to 12 Synonymy.—Sarcophaga rapax days. Adults unknown. OWS, Walker, S. helicis Townsend (4, G/Y, SA, N/H, and MOA: Not p. 512; 320, p. 9A9). given. AF and HAB : Unknown. Recorded distribution.—South- After leaving their host, larvae ern Canada and the United States burrow into the soil and pupate (320, p. 949). (Map 21.) within 4 to 9 days. (5, pp. 158- Host species.—Camnula pellu- 161; 167, pp. 419, 427, 442-443, cida (Scudder), Melanoplus bi- 447-454) vittatus (Say), M. differentialis Description.—Eggs unknown. (Thomas), M. femurrubrum (De- Larvae (ñrst instar) are 2.5 mm. Geer), M. marginattus (Scud- long, narrow, and tapering. They der), M. sanguinipes (Fabricius), are not heavily armed but are the mantid Stagomantis Carolina covered by many spines. Spines Johan, and many species of Cole- completely encircle segments 2 to óptera (5, pp. 158-161; 38, p. 6; 9 on their anterior borders, al- 123, p. A68; 152, pp. 37-38; 168, though reduced on segment 9, and pp. 147-148; 224, p. 25; 318, pp. ring segments 9 to 11 on the 30, 32, 35; 32.':, p, 28; 325, p. 26). posterior borders. The cephalo- Incidence of parasitism.—None pharyngeal mechanism is small. located. There are two pairs of small pa- pillae on each of the upper and lower borders of the small pos- terior cavity. Within the cavity are two posterior spiracles with short broad slits. The anal area is also small. {167, pp. 419, 427, 442-443, 447-454) Pupae un- known. Adult length varies from 3 to 8 mm. Antennae are black. The third joints are nearly twice as long as the second. Aristae are plumose, and both the palpi and

MAP 21.—Recorded distribution of proboscis are black. The thorax is Helicobia rapax (Walker). gray pollinose with the usual ARTHROPOD AND NEMATODE PARASITES IN AMERICA 57

three to five stripes. Anterior dontis (Townsend) U, p. SU; acrostichals are lacking and there 320, p. 955). are only three postsutural dorso- Recorded distribution.—From centrals. The abdomen is black, southeast New Mexico east covered with silver, and strongly- through Florida. Also reported tessellated. Along the dorsal part from the West Indies and South are three black stripes, and the America. {320, p. 955) (Map fourth segment is red. Hypoygium 22.) of the male is small and the first Host species.—Schistocerca segment is blackish with a few americana (Drury), S. cancelata strong hairs. The second segment (Serv.), Ramalea microptera is red with scattered stout hairs. (Beauv.), and a wide variety of Forceps are yellow at the base and Hemiptera, Lepidoptera, and Co- curved forward, tapering rapidly leóptera (-í, pp. 265-267; 6, pp. to a simple point. (5, pp. 158- 590-592; 325, p. 18; 326, pp. 26- 161 ; 3^7, p. 818) 27; 33A, pp. 105-106). Discussion.—H. rapax is at- Incidence of parasitism.—None tacked by the hymenopterous located. hyperparasite Aphaereta sarco- Life history and habits.—SO: phaga Gahan in the Sacramento Eggs, larvae, and pupae unknown. area of California. Both the fly Adults not given. LOD : Unknown. and its parasite are apparently OWS, G/Y, SA, N/H, MOA, AF, very hardy, since both emerged and HAB: Not given. (171, p. from poisoned grasshoppers and 31) survived. (389) Description.—Eggs and larvae This species is very common unknown. Pupae are 7 mm. in throughout Canada and the length by 2.5 mm. in diameter and United States and generally can subshining dark red. Each an- be collected in fair numbers. It attacks at least three orders of insects, including a half-dozen species of short-horned Orthop- tera. However, its effectiveness as a grasshopper parasite is un- known.

Sarcodexia sternodontis Townsend 1892: 106 Synonymy.—Sarcodesia lam- bens Wiedemann, Sarcophaga MAP 22.—Recorded distribution of lambens (Wiedemann), S. sterno- Sarcodexia sternodontis Townsend. 58 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE terior spiracle is about as high as turned forward), and ending in broad and dark dull red; each a broadly truncate form. The contains 14 small yellow lobes. genital segment of the female is The large, round, deep posterior golden pollinose and apparently cavity contains two deep-red red in ground color with a spiracular plates, each with three rounded orifice. reddish-yellow slits. The middle Legs of both sexes are black. slit of each plate is nearly parallel The middle femur lacks a comb, with each of the others. There is whereas the middle tibia contains a broad extension on the upper two stout bristles on the front outside edge of each plate, and side. The hindtibiae are not vil- there is a short ridge on the lower lous. Wings are hyaline and lack edge of the posterior cavity. The costal spines. (5, pp. 265-267 \ anal opening is depressed and 33^, pp. 105-106) surrounded by two tubercles. Discussion.—This is a southern {12JÍ, pp. i, 11-12) species of limited distribution in Adults: Antennae are black. the United States. It attacks a The third joints are twice as long wide variety of insect species as the second. Aristae are plumose other than grasshoppers and ap- for nearly two-thirds of their parently shows no preference for length. Both the palpi and pro- the latter. Therefore I believe this boscis are black and the bucea is species to be of only minor im- covered with pale hairs. There are portance as a grasshopper para- three rows of black hair above site. the back of the head that narrow to one opposite the lower part of Sarcophaga haemorrhoidalis the eye. The thorax has the usual (Fallen) three to five black stripes and is a (Musca) 1817: 237 grayish-white poUinose. There are Synonymy.—Sarcophaga geor- three pairs of anterior acrosti- gina Wiedemann, S. aegra Walker chals and four postsutural dorso- (320, p. 959). centrals. There is a median black Recorded distribution.—C o s - shiny stripe on the abdomen and mopolitan (320, p. 958). (Map two lateral shifting ones. The 23.) fourth abdominal segment has a Host species.—Melanoplus dif- marginal row of about a dozen ferentialis Thomas and Melano- bristles. The first segment of the plus spp. in the United States; hypopygium is rather small, sub- Dociostaurus maroccanus Thnb., shiny, black, and without bristles, Locusta migratoria L., Schisto- whereas the second segment is cerca cancellata Serv., and S. globose, shining red, and without gregaria Forsk. elsewhere. It also bristles also. Forceps are yellow, attacks a lepidopteran host in ad- very wide (with the apical part dition to dead meat, decaying ARTHROPOD AND NEMATODE PARASITES IN AMERICA 59 spiracles lack spiracular plates and the anal area has two pairs of tubercles, the larger directed laterally. {167, pp. A20-A21, U25-U26, lt36, U7-]t5k) Pupae are large, 9 to 10 mm. -.V j • ^1 . ^!"? .. -.^y long by 2.5 to 4 mm. wide, and a

■ '\) '•■.=■■ • ;•■ dull brownish red. The pupal case is slightly wrinkled. Two pairs of rounded tubercles are located above the center and one pair be- low. On the deep-red base of each anterior spiracle are 14 small MAP 23.—Recorded distribution of yellow lobes. The deep, nearly Sarcophaga haemorrhoidalis (Fallen). rounded posterior cavity is lo- cated centrally on the horizontal matter, and human excrement. axis and is widest on a horizontal (5, pp. 189-193; 168, pp. 1U7-1U8; plane. Within the cavity are two 325, p.lU; 326, p. 25) dark-red spiracular plates, each Incidence of parasitism.—None with three narrow yellow slits. located. The inner slit is bent slightly just Life history and habits.—SO: above the middle, and on the in- Eggs unknown. Larvae are pres- ner edge of each plate near the ent from early to mid-July in the base is a small ridge. {12U, pp. United States. Pupae unknown. 5,1U) Adults appear in the field around Adults are 10 to 14 mm. long. the last part of June. LOD : From There are four postsutural dorso- deposition to adult is about 16 centrals, the posterior pair well days. Eggs, larvae, pupae, and developed and the anterior pair adults not located. OWS, G/Y, SA, indistinct and hardly differenti- N/H, and MOA: Not given. AF ated from the surrounding hairs. and HAB: Unknown. (93, p. The hypopygium of the male is 586; 9A, p. 808; 167, pp. mO-A21, usually black on the apical half !i.25-U26, U36, U7-U5U; 277, p. and red on the basal half. The 228; 318, p. 30) black section is pollinose with a Description.—Eggs not given. variable row of bristles on the Larvae (first instar) are about hind margin. Numerous erect 2.9 mm. long and are heavily hairs cover the red globose sec- armed with small spines. The an- ond segment. Forceps are black terior border of segments 2 to 10 and nearly straight in profile and the posterior borders of seg- with a notch behind near the base. ments 6 to 12 are encircled by The genital segment of the fe- bands of these spines. Posterior male is deep red with a rounded 60 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE opening, which has a notch above. pp. 318-320; 167, pp. A20-Í21, Outer verticals and orbitals are 425-426, 436, 447-454) indistinct. Sarcophaga harpax Pandelle Legs are black; the hindtibiae have long villosity that spreads 1898: 189 both ways. Only one or two Synonymy.—Sarcophaga tu- , bristles rise from the front side berosa var. harpax Pandelle (320, of the middle tibiae, whereas short p. 958). spines are near the tip of the Recorded distribution.—South- i middle femora. Wings are hyaline ern Canada, northern United with the first vein bare. (5, p-p. States, and south through Ari- 189-193; 108, p. 237) zona, Kansas, and New York {320, Discussion.—This fly has also p. 958). (Map 24.) been reported as breeding on hu- Host species.—Camnula pellu- " man excrement and being passed cida (Scudder) and Melanoplus -^ with stools of man. With ref- sanguinipes (Fabricius) (38, p. 6; erence to this, Aldrich (5) re- 123, p. 485; 152, pp. 37-38; 168, ported, "These cases raise the pp. 147-148 ; 316, p. 500 ; 318, pp. suspicion that intestinal myiasis 30-32). ,. in man may be often or generally Incidence of parasitism.—None caused by this insect. The habit located. involves not only the deposit of Life history and habits.—SO: , larvae on human food, which Eggs, larvae, and pupae unknown. probably occurs at times in Adults occur as early as May in several scavenger species of Sar- Utah. LOD: Eggs, larvae, and cophaga, but also the ability of the pupae unknown. Adults not given. larva to develop almost or quite OWS, G/Y, SA, N/H, andMOA: ' to full size within the alimentary Not given. AF and HAB: Un- . canal of man, which has not as yet been proved in any other species." Keilin (162) proposed that S. haemorrhoidalis accidentally in- gested with food is not a parasite of man but is able to survive within his digestive tract and to feed on the partly digested and decaying food therein. The only damage caused, he claims, is the scratching of the gut wall by the oral hooks. It appears to be un- important as a parasite or grass- MAP 24.—Recorded distribution of hopper. (5, pp. 189-193; 162, Sarcophaga harpax Pandelle. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 61

known. (163, pp. SiS-SU; 168, therefore, of the American species pp.lW-li8;300,p. 60) is Sarcophaga harpax according Description.—Eggs, larvae, and to Stone et al. (5, pp. 171-173; pupae unknown. 168, pp. H7-1A8; 318, pp. 30, 32; Adult males are 8 mm. long. An- 320, p. 958) tennae are brownish black. The Sarcophaga sarraceniae Riley third joints are twice the size of the second, reaching four-fifths 1874: 238-240 the way to the vibrissae. Aristae Synonymy.—Sarcophaga car- are plumose for slightly more than naria var. sarraceniae Riley, S. half their length. Palpi and pobos- carnaria L. (^, p. 513; 320, p. cis are black and ordinary. The 959). thorax is whitish pollinose and Recorded distribution.—New has the usual three to five black York, North Carolina, South Car- stripes. There are four post- olina, Georgia, and Alabama (320, sutural dorsocentrals, but the an- p. 959). (Map 25.) terior acrostichals are not differ- Host species.—Melanoplus san- entiated. Abdomen is also silvery guinipes (Fabricius), Chorto- pollinose and has fine hairs but phaga viridifasciata (DeGeer), no bristles. The second segment several other species of Orthop- is globose and shining with abun- tera, several species of Lepidop- dant erect hairs. Forceps are tera, and one of Coleóptera (325, black with dense black hair. p. 8; 326, p. 28; 389). Legs are black. There are two Incidence of parasitism.—None series of stout spines below on the located. middle femur, and on the middle Life history and habits.—SO: tibia is a single bristle on the outer Eggs, larvae, and pupae unknown. front side with sometimes an Adults not given. LOD : Unknown. added small one. There is a very dense villosity which spreads both ways on the hindtibia. Wings are subhyaline and lack costal veins. (5, pp. 171-173; 2U3, p. 199) Discussion.—Aldrich (5), Spencer and Buckell (315), and others termed this species Sarco- phaga tuberosa var. harpax Pan- delle, and Knowlton and Janes (166) referred to it as Sarco- phaga tuberosa. However, S. tuberosa does not occur in the MAP 25.—Recorded distribution of United States. The present status. Sarcophaga sarraceniae Riley. 62 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

OWS, G/Y, SA, N/H, MOA: Not three well-developed rows of given. AF and HAB: Unknown. bristles. Pulvilli are long and (5^ V. US\ 16U, VV' UUl-UJi'2\ 22U\ brown. Wings are subhyaline and 26i,p.32A\265,p.262) lack subcostal spines. (261, pp. Description.—Eggs, larvae, and 23 8-2 W) pupae unknown. Discussion.—There is consider- Adults are 8 to 12 mm. long. able doubt that this species is an Antennae are reddish brown. The insect parasite. Riley (264') gave third segments are twice as large ^^veral grasshopper species as as the second and reach three- hosts that were later reported in quarters the distance to the vib- Thompson (325). However, Aid- rissae. Aristae are plumose. Palpi rich (5) stated that Riley mis- and proboscis are black. There are identified his material and that three rows of black hairs behind the true S. sarraceniae has never the eye on the back of the head. been reared from anything but the The thorax has five to seven black cups of the plant Sarracenia. stripes with intermediate poUi- Nothing was located later than nose stripes divided by black when Aldrich's reference to clarify this viewed from behind. There are position. (5, pp. 86-89; 26i, p. four postsutural dorsocentrals al- 32Jf\ 325, p. 8; 326, p. 28) though the second from the suture is sometimes reduced, giving the Sarcophaga sarracenioides appearance of only three. The ab- Aldrich domen is entirely gray and tessel- 1916: 227 lated with the tessellation more Synonymy.—Sarcophaga tu- concentrated posteriorly. Hypopy- berosa var. sarracenioides Aid- gium of the male is red and rather rich, S. misera var. sarracenioides large. The middle and posterior A\áv\(í]\(320,p.959). regions of the first segment are Recorded distribution.—South- pollinose and lack bristles. The ern Canada and the United States, second segment is shining, glo- (320, p. 959). (Map 26.) bose, hairy, and also without Host species.—Camnula pellu^ bristles. Forceps are stout and cida (Scudder), Chortophaga vir- taper on the outer side with the idifasciatus DeGeer, Melanoplus tip prolonged in an oblique tooth bivittatus rSay), M. differentialis inwardly. On the anterior lateral (Thomas), M. femurrubrum (De- edge are several teeth. Geer), M. sanguinipes (Fabri- Legs are black and the hind- cius), and species from the orders tibia has long dense villosity that Coleóptera, Lepidoptera, and spreads in both directions. On the other Orthoptera (5, pp. 227-232', middle femur are rows of bristles 38, p. 6; 93, p. 586; 123, p. 485; below which form combs on the 12Jf, pp. U, 12; 325, p. 8; 326, p. apical half. On the hindfemur are 28; 336, p. 31; 389) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 63 Incidence of parasitism.—None encircled with spines, which are located. short, broad, and similar in form Life history and habits.—SO: and color on all segments. The Eggs, larvae, and pupae unknown. border of the posterior cavity is Adults appear early to mid-June sparsely armed with the usual in Montana, North Dakota, and long hairlike spines, and the South Dakota, early May in upper border has two pairs of Texas, and April in British Co- tubercles. Within the cavity the lumbia. LOD: Eggs unknown. posterior spiracular slits are not Larva to adult requires 20 days. surrounded by peritreme. (167, OWS, G/Y, SA, N/H, and MOA: pp. Jf25-A26, ASS, U7-U5U; 277, Not given. AF: Larvae can be p. 228) reared on decaying meat. HAB: Pupae measure 7 to 10 mm. long Unknown. Each female produces by 3.5 mm. wide and are dull to on an average 38 larvae. (5, pp. dark red. Numerous distinct 227-282; 167, pp. U25-U26, A38, wrinkles are noticeable on the kU7-U5U) edge of the posterior cavity, and Description.—Eggs unknown. there is a rugose conical tubercle Larvae (first instar) are about on each side of the anal opening. 3.3 mm. long, taper slightly pos- Each anterior spiracle has 18 yel- teriorly, and are heavily armed low lobes surrounding the reddish with spines. The anterior borders rugose center. Most of the pos- of segments 2 to 10 are completely terior cavity is below the hori- encircled with spines, whereas on zontal axis. Within are two the ventral part of segment 11 reddish-black shining spiracular the spines extend well up laterally plates that contain three yellow and stop. The posterior borders slits each; the middle slits con- of segments 6 to 11 are completely verge slightly toward the apex. The anal opening is located below the posterior cavity. {12A, pp. U, 12) Adults are 8 to 13 mm. long. Antennae are black with the third joints 1.5 times as long as the second. Aristae are plumose for three-fifths of their length. Both the palpi and proboscis are ordi- nary and black. There are three rows of black hairs on the back of the head. Outer verticals are absent. The thorax is gray polli- MAP 26.—Recorded distribution of nose and has the usual three to Sarcophaga sarracenioides Aldrich. five black stripes. Postsutural 64 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE dorsocentrals number four, but longs to S. sarracenioides. This the anterior acrostichals are lack- parasite attacks many hosts in ing. The abdomen is gray and several orders, but its importance tessellated, with the fourth seg- as a parasite of grasshoppers is ment largely red and bearing a unknown. (5, pp. 227-232) marginal row of about 14 bristles. There is long dense hair emanat- Sarcophaga shermani Parker ing from the ends of the ventral 1923: 124 parts of the tergite. The hypopy- Synonymy.—Sarcophaga ex- gium of the male is red. The first überaus Pandelle, S. tuberosa var. segment is pollinose and lacks exuberans Pandelle (320, p. 959). bristles on the hind edge. The Recorded distribution.—South- second segment is shiny red and ern Canada and the United States globose without bristles. Forceps south to the southernmost region are black and curve forward. (320, p. 959). (Map 27.) Front and back edges of the for- Host species.—Melanoplus san- ceps are nearly parallel, and there guinipes (Fabricius) and possibly is an apical tooth formed by the other grasshoppers (38, p. 6; 168, excision of the front edge. At the p. 1Í8; 318, pp. 30, 32). point of attachment of the forceps Incidence of parasitism.—None is a covering of dense black hair. located. The genital segment of the fe- Life history and habits.—SO: male is reddish yellow, with al- Eggs, larvae, and pupae unknown. most a triangular orifice, which is Adults not given. LOD : Unknown. fringed with a few small bristles OWS, G/Y, SA, N/H, and MOA: on the upper part. Not given. AF and HAB: Un- Legs of both sexes are black known. (316, p. 500) and the hindtibiae contain a dense Description.—Eggs, larvae, and villosity. A rather loose comb of pupae unknown. spines is located on the lower hind Adult males are similar to S. side of the femur near the tip, sarracenioides and are separated and a well-developed lower row of from the latter by differences in bristles is noticeable on the outer genitalia, hypopygium, and for- side of the hindf emora. Wings are ceps. S. shermani males are 8 to hyaline and lack costal spines. 13 mm. long and are generally (5, pp. 227-232) gray pollinose. Antenna are black. Discussion.—S, sarracenioides The third joints are 1.5 times was listed as a variety of tuberosa longer than the second. Aristae by Aldrich (5) and later given as are plumose for three-fifths their a variety of misera by Knipling length. Palpi and proboscis are (167), Aldrich also believed that hairy and ordinary. There are the Riley mistakenly attributed infor- usual three rows of black hairs mation to S, sarraceniae that be- along the back of the head, 4r ARTHROPOD AND NEMATODE PARASITES IN AMERICA 65

middle tibia and a well-developed lower row of bristles is on the outer side of the hindf emur. Hind- tibiae have a dense villosity. Wings are hyaline and lack the costal spines. (5, pp. 227-229; 237, p. 124.) Discussion.—Knowlton and Janes (168) reported that S. tuberosa is parasitic on grass- hoppers in Utah. Since S. tuberosa does not occur in Canada or the United States, but two species ^ MAP 27.—Recorded distribution of Sarcophaga shermani Parker. that were originally described as varieties do, I believe their ref- erence to mean either this species, whereas the outer verticals are S. shermani, which is the same as absent. Thorax is a gray pollinose S. exuberans, or S. sarracenioides. and displays the usual three to This species is believed to be of five black stripes. There are four minor importance as a grass- postsutural dorsocentrals and the hopper parasite. (168, p. US) anterior acrostichals are either small or absent. The abdomen is Sarcophaga sinuata Meigen gray and tessellated. Its fourth 1826: 22 segment is largely red, usually on the posterior half, and has a mar- Synonymy.—S arc ota chin ella ginal rovir of approximately 14 intermedia Townsend, S. sinunta bristles. A mat of dense, long hair (Meigen) (318, p. 959). - is on the underside of the tergite. Recorded distribution.— The first segment of the hypopy- Alaska, most of Canada, and the gium is of median size, shining United States except the south- black, and without bristles along ernmost region. Also reported the hind edge, although it does from Germany. (320, p. 959) occasionally contain some small (Map 28.) " hairs. The second segment is red, Host species.—Camnula pellu- . globose, and also without bristles. cida (Scudder), Melanoplus bivit- Forceps are yellow and very tatus (Say), M. differentialis slender. (Thomas), and M. sanguinipes Legs are black and the middle (Fabricius) (38, p. 6; 123, p. 485; femur has a rather loose comb of 137, p. 521 ; 164, p. 441 ; 168, pp. spines on the lower back side near 147-148; 300, p. 60; 306, pp. 28- the tip. Two bristles are located 33; 307, p. 241; 318, pp. 30, 32; on the outer front side of the 325, p. 14; 326, p. 29). 66 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE 1 dorsocentrals and the anterior acrostichals are generally absent. ^ The abdomen is untapering, black, and tessellated. Both segments of -- the hypopygium are shiny black and have erect hairs, which be- come almost bristlelike on the hind margin of the first. Palpi are slightly clávate in the female, the , abdomen is convex and broad, and ^ the genital segment is black with • a vertical slit guarded by a row of MAP 28.—Recorded distribution of bristles. Sarcophaga sinuata Meigen. Legs of both sexes are black. • There is no comb of spines on the ^ Incidence of parasitism.—^Un- middle femora although a few < known. long slender hairs are found on ' Life history and habits.—SO: the inner front side of the hind- Eggs, larvae, and pupae unknown. tibia. Wings are nearly hyaline, Adults appear in the field from and the costal spine is distinct. ■ early June to late August in Can- (5, pp. 67-69; 217, p. 22) -i ada. LOD : Unknown. OWS, G/Y, Discussion.—This species is a SA, and N/H, Not given. MOA: parasite of the grasshopper Doci- ', Attacks hosts in flight. AF and ostaurus maroccanus Thnd. in HAB: Unknown. Smith (307, p. Spain and is also reported from 2U) believed this species to be Germany. similar in life history and habits to Blaesoxipha kellyi. Tachinidae Description.—Eggs, larvae, and The Tachinidae is a large eco- "; pupae unknown. nomically important, cosmopoli- Adult flies are 5 to 9 mm. long. tan family, which apparently The head has a pale-yellow sheen contains more species than any extending smoothly down the other family of muscoid flies. parafacial. Antennae are black. Adults are usually medium to The third segment is less than large, robust, and very good fliers. double the length of the second. Many are similar in appearance * Aristae are long and plumose for to the house and flesh flies, but about half their length. Palpi and they are distinctive in having both proboscis are black. The thorax the hypopleural and pteropleural is gray and displays the usual three blackish stripes, which are bristles developed and the post- generally visible only from the scutellum prominent. (Fig. 9.) back. There are three postsutural The Tachinidae is the most ef- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 67

FIGURE 9.—Adult tachinid fly. fective family of the Díptera in their hosts. Larvae emerging pre- biologically controlling other in- maturely generally die, indicating sects. They are common parasites no feeding occurs in the soil. of caterpillars and of adult and Mating generally occurs shortly young Coleóptera and Hemiptera. after emergence, often within Only a very few species attack minutes, and the male may mate grasshoppers, and of these only with several females. Gestation about three species are considered varies from 2 days to 4 weeks, important. with the young laid either as un- Adult food generally consists of incubated eggs, incubated eggs, or plant and insect secretions, liquid living first-instar larvae. Eggs are organic material, and sometimes the blood of the host. Larvae of commonly deposited on vegetation, the grasshopper parasites feed but most of those attacking grass- exclusively on the hemolymph, the hoppers deposit their eggs di- reproductive organs and their rectly on the host. These hatch products, and the fat body of almost immediately and the larvae 68 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE quickly burrow into the hemocoel Winter is usually passed as pupae where they develop through at in the soil, although Ceracia den- least three instars. Most emerge tata (Coquillett) is reported to in late summer or during the fall, over-winter as a larva within its killing the host, and enter the soil host. When diapause occurs, it is where pupation eventually occurs, usually in the pupal stage.

KEY TO ADULT TACHINIDS PARASITIC ON GRASSHOPPERS la. Abdomen with macrochaetae ^ lb. Abdomen without macrochaetae or long bristles; sides of face bare; apical cell closed; hindtibiae with row of erect bristles Trichopoda plumipes (Fabricius) 2a. Facial ridges bristled on lower half or less 3 2b. Facial ridges bristled on at least lower two-thirds; apical cell ends before apex of wing; bend of fourth vein with distinct appendage or fold; first vein bare; eyes hairy Euphorocera claripennis (Macquart) 3a. Apical crossvein present (fig. 10) 4 3b. Apical crossvein absent; posterior crossvein present; antennae do not reach below lower one-third of face Hemithrixion oestriforme B. & B. 4a. Apical cell without petiole 5 4b. Apical cell with petiole about one-third length of apical crossvein; vibrissae above oral margin Leucostoma simplex (Fallen) 5a. Posterior pair of presutural acrostichals located slightly behind posterior presutural dorsocentrals ; two pairs of presutural dorsocentrals 6 5b. Posterior pair of presutural acrostichals located much behind posterior pair of presutural dorsocentrals and very close to suture ; always three pairs of presutural dorsocentrals 7 6a. Prosternum with hair Ceracia dentata (Coquillett) 6b. Prosternum bare Acemya tibialis CoquiUett 7a. Second and sometimes third abdominal segments display some rufous or red at sides Lespesia anonyma (Riley) 7b. Not as above; 6 to 7 mm. in length; displays grayish pollinose stripe on shiny black mesonotum Lespesia frenchii (Williston)

Acemya tibialis Coquillett Host species.—Ageneotettix de- 1897- 116 orum (Scudder), Aeropedellus ^., . ,. clavatus (Thomas), Aulocara Synonymy-Acemym UUahs ^^^ ^^ ^^^^^^ ^^^^^.

(Coquillett) (^, V. 267; 320, p. ;,^^J,,^^, (LatrelUe), Melano- ^'^Recorded distribution.-Sonth- V^""' angustipennis (Dodge), M. ern Canada and the United States bivittatus (Say), M. boreahs south to the southeastern region (Fieber), M. bruñen (Scudder), (320, p. 1036). (Map 29.) M. confmus Scudder, M. dawsoni ARTHROPOD AND NEMATODE PARASITES IN AMERICA 69

costal vein. apical crossvein.

FIGURE 10.—Tachinid wing showing apical crossvein.

(Scudder), M. femurrubrum pupae. G/Y: Three to four in (DeGeer), M. foedus Scudder, M. Canada. SA: All stages. N/H, infantilis Scudder, M. keeleri MOA, AF, and HAB: Unknown. (Dodge), M. packardii (Scudder), Each female produces up to 150 M. sanguinipes (Fabricius), and eggs per litter. Although the host Phoetaliotes nebrascensis may survive the attack, its re- (Thomas) (123, pp. 469, 486 ; 183, productive capacity is destroyed. p. 9; 227, pp. 935-936; 306, pp. (183, p. 9; 300, pp. 60-61; 307, 28-33 ; 307, pp. 246-248 ; 308, pp. pp. 246-248; 316, p. 501) 187-199; 389). Description.—Eggs unknown. Incidence of parasitism.—In Larvae (second instar) are 1.8 Canada the highest rates reported mm. long by 0.5 mm. wide and were 53.6 percent of M. packardii, 55 percent of M. femurrubrum, 65 percent of M. bivattatus, 32 percent of M. sanguinipes, and 16 percent of C. pellucida (307, p. 247). Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults in Canada are in the field early in June until late September. LOD : Eggs and larvae unknown. Pupae 9 to 17 days. Diapause lasts 135 days regardless of tempera- MAP 29.—Recorded distribution of ture. Adults unknown. OWS: As Acemya tibialis Coquillett. 70 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE ivory white. They are cylindrical hosts, whereas M. femurrubrum with the ends rounded. The dorsal and M. packardii are also heavily ^ part supports a scattering of very parasitized. (307, pp. 2A6-2Í8) fine minute dots, and there is a Ceracia dentata (Coquillett) band of spines on the anterior- ventral margin of segments 2 to (Acemyia) 1895: 311 10, although these are inconspic- Synonymy.—Acemyia dentata uous on segments 3 and 4. On seg- Coquillett (^ p. 267; 320, p. , ments 2 to 10 are two to three 1037). rows of large scalelike spines Recorded distribution.—South- ^ with a single sharp tip at the ern Canada and the United States , apex of each. Posterior spiracles (320, p. 1037). (Map 30.) are small and are projected on a Host species.—Ageneotettix de- tip of a feltlike chamber at the orum (Scudder), Aulocara elliotti -^ posterior surface of the last ab- Thomas, Camnula pellucida dominal segment. The bucco- (Scudder), Campylacantha oli- i pharyngeal armature bears a vacea (Scudder), Chortophaga , single median tooth, with the an- viridifasciata (DeGeer), C. spp., terior margins finely serrated Corthippus spp., neg- (309, pp. 99-100) Pupae un- lectus (Thomas), Dichromorpka ■; known. viridis (Scudder), Melanoplus _ Adult females are 4 mm. long angustipennis (Dodge), M. bivit- and blackish. Palpi, apex of the tatus (Say), M. confusus ^ proboscis, and the first two joints (Scudder), M. dawsoni of the antennae, as well as the (Scudder), M. differentialis trochanters, apices of the femora, (Thomas), M. femurrubrum and the entire tibiae are yellow. (DeGeer), M. gracilis (Bruner), The third joints of the antennae M. infantilis Scudder, M. pack- are 1.5 times as long as the second. ardii Scudder, M. sanguinipes -' The aristae are thickened at the basal fourth. The thorax is gray pollinose and marked with four black stripes. The abdomen is also gray pollinose. Hairs are de- pressed and at the base of each is a black dot. Haltères are whitish, and wings are hyaline. (70, p. 116) Discussion.—A. tibialis is con- sidered in both Canada and the United States as a good natural control agent. M. bivattatus and MAP. 30—Recorded distribution of M. sanguinipes are preferred Ceracia dentata (Coquillett). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 71

(Fabricius), Phoetaliotes nebras- tached by one end to the posterior censis (Thomas), Psoloessa deli- end of the larva and by the op- catula ( Scudder), Spharagemon posite end to a large trachéal spp., and Syrbula admirabilis trunk in the host's thorax. (280 ; (Uhler) (70, pp. 9, 117; 128, pp. 281, pp. 83-87; 307, pp. 2UU-2U6; JÍ69, US7\ 132, p. 519; 152, pp. 3i- 308, pp. 182-199; 323, p. 122) 37; 22JÍ, p. 25; 227, pp. 935--936; Description.—Eggs are 0.4 mm. 281, pp. 83-87; 306, pp. 28-33; long by 0.2 mm. wide, flattened, 307, pp. 2U-2JÍ6; 325, pp. 8, 9; and elongate-oval. The chorion is 326, p. JÍ9; 389). thick dorsally and thin ventrally Incidence of parasitism.—Re- on the side of attachment to the corded incidences are very low, host. (307, pp. 22h'-2U6) Lar- ranging from 1.7 percent in Can- vae (first instar) have a single ada to 3 percent in Montana and median oral hook dorsad of a well- Missouri (152, pp. 3^-37; 227, developed buccopharyngeal plate. pp. 935-936). Second-instar larvae are 2.5 mm. Life history and habits.—SO: long by 0.7 mm. wide, ivory white, Eggs, larvae, and pupae unknown. and cylindrical with the ends Adults first appear in June in rounded. Spines are minute. The Canada and remain until the first single median tooth is finely ser- killing frost. LOD: Eggs and rated on the anterior margin. larvae unknown. Pupae require Posterior spiracles are bilobed about 2 weeks at room tempera- and appear at the tips of elon- ture. Adults unknown. OWS: As gated felt chambers, which are larva in the host. G/Y: Three to present on the posterior surface four in Canada. SA : Older nymphs of the last abdominal segment. and occasionally young adults. Third and fourth instars are N/H: Unknown. MOA: An incu- larger but similar to the second. bated egg is deposited directly on (281, pp. 83-87; 307, pp. 100- the grasshopper where it remains 102) fastened by a gelatinous secretion Pupae: The puparium varies until hatching. AF: Plant secre- greatly in size and is dull reddish tions and honeydew. HAB: Un- brown. The horizontal fracture known. line denotes the location where Up to 200 eggs are produced by the puparium may split at the each female and are incubated in end of pupation. Two heavily the uterus. They are deposited on sclerotized spiracular projections the host where they adhere until are located ventrally to the base the larva has hatched and gained of the anal scar. (281, pp. 83- entrance into the hemocoel where 87) Adults are 6 to 7 mm. long. it becomes free living. Later lar- Tibiae and antennae, except the val instars receive outside air by apex of the second joint, are black. means of a cuplike funnel at- (69, p. 311 ; 70, pp. 9, 116) 72 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Discussion.—I am not certain why this species has not been re- ported as a more effective grass- hopper parasite. Possibly higher rates of parasitism are not re- corded because of misidentifica- tion or possibly it is not important where it has been collected. The placement of the eggs directly on the host, the production of nearly 200 eggs per female, three to four generations per year, the wide VYT.^-^ grasshopper host range, and its MAP 31.—Recorded distribution of extensive geographical distribu- Euphorocera claripennis (Macquart). tion all indicate its great poten- tial. Possibly such hyperparasites species of Lepidoptera, several of as Perilampus hyalinus Say, Hymenoptera, and one species of '■ Brachymeria tegularis (Cresson), Coleóptera (132, p. 519; 151, p. and Brachymeria coloradensis U3; 326, pp. 97-98). (Cresson) are effective in keeping Incidence of parasitism.—None the fly population low. A lesser located. possibility is high mortality rates Life history and habits.—SO: ^ within the hosts, although this Eggs, larvae, and pupae unknown. seems unlikely. Adults unknown (may over- winter). LOD: Unknown. OWS: Euphorocera claripennis As pupae or adults. G/Y, SA, (Macquart) N/H, and MOA: Not given. AF (Phorocera) 1848: 209 and HAB: Unknown. (93, p. ^ 582; 225, p. 25) Synonymy.—Phorocera clari- Description.—Eggs, larvae, '• pennis Macquart, P. edwardsii and pupae unknown. Adults are (Williston)??, Euphorocera vi- 5 to 14 mm. long, palpi are yellow, brissata (Brauer and Bergen- and the head at the vibrissae is stamm) ?, Podotachina vibrissata noticeably shorter than at the (Brauer and Bergenstamm), base of the antennae. Facial • Eurygaster septemtrionalis ridges are bare of bristles on the « Walker (4-, PP. 259-260; 320, p. outer side. There are four post- lOSA). sutural dorsocentrals on the tho- Recorded distribution.—South- rax, but the diskal bristles are ern Canada and the United States absent from the second and third i320,p. 105Jf). (Map 31.) abdominal segments. (70, pp. Host species.—Melanoplus dif- 11-12, 102; 213, p. 209) ferentialis (Thomas) plus many Discussion.—Some references ARTHROPOD AND NEMATODE PARASITES IN AMERICA 73 of E. claripennis are believed to lum (Say), Camnula pellucida be confused with those of E. (Scudder), Chorthippus longi- edwardsii (Williston), a very cornis (Latreille), Melanoplus closely related species. E. clari- bivittatus (Say), M. bruneri pennis is a prominent species and Scudder, M. dawsoni (Scudder), has a very wide host range. It is M. femurrubrum (DeGeer), M. noted as a parasite of the tus- foedus Scudder, M. infantilis sock moths (Hemerocampa leucos- Scudder, M. packardii Scudder, tigma A. & S., Stilpnotia salicis L., and M. sanguinipes (Fabricius) Porthetria dispar L., and Nygmia (123, pp. A67, 469, Í76, 487; 152, phaeorrhoea Don.), but appar- pp. 36-37 ; 227, pp. 935-936 ; 239, ently it shows no preference for p. 492; 306, pp. 28-33; 307, pp. any grasshopper species; there- 248-249). fore it is probably of little value Incidence of parasitism.—In as a grasshopper parasite. Canada the parasitism rarely ex- ceeds 5 percent with a maximum Hemithrixion oestriforme Brauer and Bergenstamm of 18.2 percent, but this species is considered very important in 1891: 357 certain areas. In Montana the Synonymy.—None (^, p. i.26', highest rate recorded was 1 per- 320, p. 1037). cent. (227, pp. 935-936 ; 307, pp. Recorded distribution.—From 248-249; 389) British Columbia through Mani- Life history and habits.—SO: toba and south through Texas Eggs, larvae, and pupae unknown. (320, p. 1037). (Map 32.) Adults in Canada are present in Host species.—Ageneotettix de- the field from mid-June to late orum (Scudder), Boopedon nubi- September. LOD : Eggs unknown. Larvae require 19 to 21 days. Pupae 9 to 20 days. Adults un- known. OWS: As pupae in the soil. G/Y: Two to three in Can- ada. SA: All stages. N/H: Un- known. MOA : Not given. AF and HAB : Unknown. (307, pp. 248- 249 ; 323, p. 122) Description.—Eggs unknown. Larvae (second instar) are about 1 mm. long, cylindrical with rounded ends, and ivory white. On the anterior ventral margin of

MAP 32.—Recorded distribution of segments 2 to 10 is a band of Hemithrixion oestriforme Brauer and spines, which on segments 3 and Bergenstamm. 4 consists of one to two rows of 74 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE small inconspicuous spines. Dor- souri (259, pp. 129-131; 26^, p. . sal surface of segments is covered 323; 320, p. 1101). (Map 33.) with very minute black dots. The Host species.—"Locusts." Also buccopharyngeal armature con- species of Lepidoptera. (122, p. sists of a single median tooth with A87; 26h, p. 323, 326, pp. 32-33) a finely serrated anterior margin. Incidence of parasitism.—None A dorsal accessory tooth is appar- located. ent but small. On the posterior Life history and habits.—SO: surface of the last abdominal Eggs, larvae, and pupae unknown, j segment are located small two- Adults not given. LOD: OWS: ' lobed spiracles. Antennae are well Not given. G/Y, SA, N/H, MOA, pigmented with the diameter AF, and HAB : Unknown. twice that of its length. {309, p. Description.—Eggs, larvae, 99) Pupae unknown. Adult flies and pupae unknown. Adults: Al- are approximately 6 mm. long. though Riley (259) in 1872 stated Antennae are short. The third that he would not describe this joint beneath is narrow and species at that time, his short pointed. Forehead of the female notations in that publication were is narrow. Tarsi are elongated. the only descriptions located. He Vibrissae contain numerous stated, "It belongs, apparently, to bristles, which are broad, sep- Macquart's genus Masicera. I arated, and flat. (26, p. 357) have bred precisely the same spe- . Discussion.—Although little is cies from Citheronia regalis, and known of the life history and others that differ only in being habits of this species, Smith smaller, from Prodenia autumn- (307) indicated that the type and alis, an undescribed noctuan, Cyn- number of larval stages in the thia cardui, Heliothis armígera, host are similar to those of Cer- Datana ministra, and Danais < acia dentata and that the habits and development are probably similar for both C. dentata and Acemya tibialis. In Canada this species is considered very impor- tant in certain areas as a grass- hopper control agent. (307, pp. 2^8-249)

Lespesîa anonyma (Riley) (Tachina) 1872: 129 Synonymy.—Tachina anonyma Riley (Â, p. ^69; 320, p. 1101). MAP 33.—Recorded distribution of Recorded distribution.—Mis- Lespesia anonyma (Riley). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 75

archippus; while I have others ^ from Sphinx Carolina which differ only in having the apical joint of abdomen rufous. The face appears quite golden on the top and the second and sometimes the third abdominal joints incline to rufous at sides." (259, pp. 129-181) Discussion.—There is some ' doubt whether this is a valid spe- cies. The poor description and damaged type specimen make establishment of this name un- MAP 34.—Recorded distribution of certain. The consensus of some is Lespesia frenchii (Williston). that this is a synonym of Lespesia -- frenchii (Williston). pupae unknown. Adults are 6 to 7 mm. long. The ground color is Î Lespesia frenchii (Williston) scarcely visible although the sides of the front are a distinct yellow. (Masicera) 1889: 1923 Antennae are broad and com- Synonymy.—Masicera frenchii pletely black. Aristae are thick- Williston, fachina longicornis ened for at least half their length. Wiedemann, Achaetoneura Palpi are yellow. There is a hespersus Brauer & Bergen- grayish poUinose stripe on the stamm, M. sphingivora Townsend, shiny black mesonotum, and the Frontina malacosomae Curran, F. black abdomen is covered with sórdida Curran (4, p. 4631 ; 320, gray. Legs are black. The wings p. 1102). are gray hyaline and the haltères Recorded distribution.—Most are white. {70, pp. 15-17, 107; of Canada and the United States 376, p. 1923) {320, p. 1102). (Map 34.) Discussion.—The synonymy of Host species.—Dissosteira Car- this species may also include olina (Linnaeus) {123, pp. 467, Lespesia anonyma (Riley). 487; 260, p. 139). Incidence of parasitism.—None Leucostoma simplex (Fallen) £> located. {Ocyptera) 1820: 8 Life history and habits.—SO: Synonymy.—Ocyptera simplex Eggs, larvae, and pupae unknown. Fallen, Leucostoma atra Town- Adults not given. LOD : Unknown. send {4, p. 438; 320, p. 976). OWS, G/Y, SA, and N/H: Not Recorded distribution.—South- given. MOA, AF, and HAB : Un- ern Canada and the United States known. south to the southeastern region Description.—Eggs, larvae, and {320, p. 976). (Map 35.) 76 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE and three postsutural dorsocen- trals. The abdomen is also desti- ^ tute of pollen but supports a pair of marginal bristles on each side > of the first two abdominal seg- « ments. (109, p. 8) Discussion.—There is not enough available information to ' evaluate this species. . Trichopoda plumipes ^ (Fabricius)

MAP 35.—Recorded distribution of (Threva) 1805: 220 Leucostoma simplex (Fallen). Synonymy.—Threva plumipes ^ Fabricius, Trichopoda histrio Host species.—Melanoplus san- Walker, T. trifasciata Loew (i, ^■ gunipes (Fabricius) (123, p. -i-S?; pp. 425-A26; 320,p. 966). 306, pp. 28-33). Recorded distribution.—From Incidence of parasitism.—None Kansas east through Connecticut, located. south through Florida, and west - Life history and habits.—SO: through Texas (320, p. 966). , Eggs, larvae, and pupae unknown. (Map 36.) Adults not given. LOD : Unknown. Host species.—Dissosteira ve- OWS, G/Y, SA, N/H, and MOA: nusta (Stäl) (70, pp. 21, 23, Jf8; Not given. AF and HAB: Un- 123, p. Í87). known. Incidence of parasitism.—None Description.—Eggs unknown. located. Larvae (second instar) are 2.0 Life history and habits.—SO: mm. long by 0.65 mm. wide, ivory white, and cylindrical with rounded ends. There are minute spines on the ventral surface of segments 3 to 10, which are in one or two rows and dotlike. No spines are on segment 2. The infrabuccal plate has the lateral tips of the marginal arc directed toward the base of the stem. (306, p. 100) Pupae unknown. Adults are 4 to 5 mm. long and black in basic color. Palpi are yellow. The thorax is shining and almost without MAP 36.—Recorded distribution of pollen but has three sternopleural Trichopoda plumipes (Fabricius). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 77

Eggs, larvae, and pupae unknown. cies reported here attack only the Adults not given. LOD: Eggs, eggs of the Acrididae. larvae, and pupae unknown. The scelionids practice the Adults not given. OWS, G/Y, SA, habit known as phoresy. As the and N/H : Not given. MOA, AF, host female becomes gravid and and HAB : Unknown. the eggs approach maturity, the Description.—Eggs, larvae, and female parasite attaches itself to pupae unknown. Adults are 7 to the host female and adheres 11 mm. long and generally yellow thereon until oviposition, when it to black. There is an opaque in- departs to attack freshly laid terrupted line of yellowish pollen eggs. Scelio species will attack the on the second, third, and fourth eggs from the time of deposition abdominal segments. The thorax until 24 hours later. They may or is shining black and the scutellum may not bite into the eggs of the and femur are yellow. That part host before depositing their own in front of the transverse suture eggs and usually parasitize all the is yellowish pollinose and marked eggs of a pod. with three to four black stripes. The parasite larva feeds on the The lower calypters are yellowish. yolk of the egg although the de- The wings are brown with a yel- veloping host embryo is not dis- low costal margin and a hyaline turbed until the parasite is in the hind margin. (70, pp, 21, 23, US ; second larval instar. During pu- 105, p, 220) pation the head of the parasite is Discussion.—Of minor impor- generally pointed toward the tance as a grasshopper parasite. anterior part of the egg. Under ideal conditions the duration of a life cycle ranges in most species Hymenoptera from 8 to 15 days; one-half to two-thirds of the time is passed in Scelîonîdae the pupal stage. It has not been Some of the smallest insects determined at what stage of known belong to the family development the parasite over- Scelionidae. Most or all of them winters. are parasitic on the eggs of in- The eggs are all stalked, with sects belonging to the orders the main body ovate to spindle Lepidoptera, Hemiptera, Orthop- shaped and tapering. Eggs are tera, Diptera, Coleóptera, and slender with no distinct line of Neuroptera. Both exotic and en- demarcation between the stalk demic species have been used and the main body. They increase successfully for biological control. the size during incubation as the Each genera is usually restricted result of the stalk being absorbed. to the hosts of one or two orders, First-instar larvae of Scelio are genera, or species. The Scelio spe- teleaf orm, completely lacking vis- 78 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE ible segmentation, but with the the anterior margin of the abdo- body divided by a sharp constric- men. It varies considerably in tion into two sections. Mandibles number and distribution among of all species are external, exceed- species. (Fig. 11.) ingly large, widely spaced, curved, The few descriptions of second- and sharply pointed. Antennal and third-instar larvae indicate processes are usually large and they are similar to the first in- conical, widely spaced, and aris- star. In several species only two ing immediately from the base of larval instars have been noted, the mandibles. In addition, there possibly because the second and is a large fleshy lobe or process third instars are similar. on the medial ventral line of the Adult wasps are characterized céphalothorax below the mandi- by having 11- to 12-segmented bles that is considered the labrum. antennae, or if the antennae are The abdomen is rather globular seven- to eight-segmented, the and terminates in a small horn or antennal club is unsegmented. tail. There is a partial or complete Front wings usually have mar- transversal ring of long hairs on ginal and stigmal veins. The

FIGURE 11.—Teleaform Scelio larva. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 79 abdomen is rather sharply keeled cell is present on the wings. Base laterally, and hindwings are nar- of the abdomen is constricted. The row without the anal lobe seg- abdomen is rather petiolate. mented by a deep incision. An- Only six Scelio species para- tennae are inserted below the sitic on grasshopper eggs are mouth close to the margin of the mentioned in the literature. C. F. clypeus, whereas the mandibles W. Muesebeck (personal corre- are normal, the tips meeting when spondence) noted a seventh closed. The pronotum is laterally species, Scelio fuscipennis Ash- extended to or nearly to the te- mead, which attacks the grass- gula, its posterior angles usually hopper Schistocerca americana are not lobed, or, if so, the tegula Drury in Mexico. These two in- is absent. The ovipositor of the fe- sects occur in the United States, male usually issues from the tip but records of attack here were of the abdomen but lacks a pair not located. It is highly probable, of external sheaths. Trochanters however, that attacks do occur are one-segmented and a costal here.

KEY TO SCELIO ADULTS la. Species lacking parapsidal furrows 2 lb. Species with parapsidal furrows 3 2a. Stigmal veins and radius lacking S, hyalinipennis Ashmead 2b. Stigma with stigmal vein S. ovivorus (Riley) Coxae dark brown or black; rest of the legs brownish yellow to reddish yellow 4 3b. Coxae and legs pale brownish yellow 5 4a. Wings hyaline, stigma reaching to costa ; antennae brown in male S. oedipodae Ashmead 4b. Wings subfuscous, an oblique stigma at end of marginal vein; antennae yellowish to yellowish red except the brown club S. bisulcus (Ashmead) 5a. Stigma lacking stigmal vein S. calopteni Riley 5b. Stigma with short stigmal vein S. luggeri Riley

Scelio bisulcus (Ashmead) Carolina, and Florida {225, p. (Sceliomorpha) 1893: 240 702). (Map 37.) Synonymy.—Sceliomorpha bi- Host species.—Dichromorpha sulca Ashmead, Scelio hisulcatus viridis (Scudder), Melanoplus Kieffer, S. monticola Brues {225, confusus Scudder, M. femurru- p. 702), brum (DeGeer), and M. keeleri Recorded distribution.—Ari- luridus (Dodge) {123, p. Ul ; 176, zona, South Dakota, Kansas, pp. 213-21Í; 177, pp. 8Ji-87\ 359, Texas, Illinois, Michigan, South p. 308). 80 TECHNICAL BULLETIN NO. 14^0, U. S. DEPT. OF AGRICULTURE

Wasps of this species become firmly attached to the grasshopper * with their legs and mandibles at an area where the femur attaches ^ to the body. Lanham and Evans (176, 177) believed that the stim- ulus for attachment is the angle formed by the femur with the ■* thorax, since dislodged wasps would always return to the same area. Lanham and Evans noted that female wasps attached them- MAP 37.—Recorded distribution of selves only to female grass- Scelio bisulcus (Ashmead). hoppers. (36, pp. 136-HO; 176, ' pp. 213-214; 177, pp. 84-87) Incidence of parasitism.—Lan- Description.—Eggs, larvae, and ham and Evans (176, pp. 213- pupae unknown. Adult females 21 Jf) indicated that about 60 are 8.4 mm. long and brownish percent of the eggs of M. confusus to black. The integument is finely were parasitized early in the year wrinkled and sparsely pubescent and about 10 percent each of two on the scutellum. The head is ' later species, M. femurrubrum wrinkled and the face subconvex. - and M. keeleri. However, the total Cheeks are short and margined populations of the last two were and the eyes ovate. Although the considerably higher. antennae are short, they are 12- Life history and habits.—SO: jointed. They are yellow except Eggs, larvae, and pupae unknown. the club, which is brown. The ' Adults appear from July to Sep- pronotum is formed into a collar » tember in Michigan. LOD and anteriorly, and there are two dis- OWS: Unknown. G/Y: At least tinct furrows in the mesonotum. one, possibly two. If only one, The metathorax is closely punc- then there are two peaks or tate and has a median furrow. The strong periods of emergence. SA : venter of the fusiform and de- - Eggs. N/H: One per egg. MOA: pressed abdomen is shining and Adult female wasps attach them- faintly lineated. Legs are brown- ' selves to the abdomen of gravid ish yellow to reddish yellow except ' adult grasshoppers and await the black coxae. Wings are sub- their oviposition. They then enter fuscous and the marginal vein the pods and insert an egg into ends in an oblique stigma. (11, - each grasshopper egg. AF : pp. 240-241) Among other things, the contents Discussion.—Lanham and of some of the host's eggs. HAB : Evans (176, pp. 213-214; 177, pp. Unknown. 84-87) showed that under proper ARTHROPOD AND NEMATODE PARASITES IN AMERICA 81

conditions parasitism can be ex- in each grasshopper egg. AF : May tremely effective. ingest some of the grasshopper egg contents. HAB: Unknown. Scelio calopteni Riley {77, p. 12; 299, p. 28). 1893: 242, 246 Description.—Eggs, larvae, and Synonymy.—None (in Ash- pupae unknown. Adult females mead 1893) (11,pp.2U2, 2U6; 225, are 3 to 3.4 mm. long. The head, - p. 702). thorax, and abdomen are black Recorded distribution.—From and wrinkled. There are faint Idaho east through New Hamp- but distinct furrows on the mes- shire, south through Louisiana onotum. On the face above the and eastern Texas, and north into antennae is a smooth, shiny Montana {225, p. 702). (Map space with fine lines on each side ^ 38.) converging toward the mouth. ^ Host species.—Camnula pellu- Mandibles are pale and reddish. "^ cida (Scudder), Melanoplus bivit- Antennae are 12-jointed and '^ tatus (Say), and M. sanguinipes brownish black. Scape and pedicel (Fabricius) {11, pp. 2U2, 2U6; are yellow. The club of the an- 118, p. 21 ; 123, p. Jt88 ; 177, p. 86 ; tennae is slightly compressed » 300, p. 58; 325, pp. 7, H; 329, p. from above and the joints are ^ 630; 336, p. 32). closely conjoined. The thorax and Incidence of parasitism.—Not tegulae are light red. The legs in- ''■ given. cluding the coxae are pale brown- Life history and habits.—SO ish yellow. Abdomen is fusiform Eggs, larvae, and pupae unknown and longer than the head and ^ Adults not given. LOD and OWS thorax combined. Wings are hya- ^ Unknown. G/Y; Not given. SA line, with the submarginal vein Eggs. N/H: Only one. MOA ending in a round stigma but lack- ^ Practice phoresy and deposit eggs ing the stigmal vein. {11, pp. 2Í2, 2^6) Discussion.—Practices phoresy on three species of grasshoppers of economic importance. The ef- fect of control, however, is un- known.

Scelio hyalinipennis Ashmead 1887: 119 Synonymy.—None (225, p. 702). Recorded distribution.— MAP 38.—Recorded distribution of Florida {10, p. 119; 225, p. 702). Scelio calopteni Riley. (Map 39.) 82 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Host species.—Chortophaga vi- low the eyes. Mandibles, scape, ridifasciata (DeGeer), Melano- and pedicel beneath in the female *" plus dijferentialis (Thomas), and are brownish yellow as are the Schistocerca obscura (Fabricius) legs and coxae in the male. An- (123, p. A88; 22U, pp. 23-2U). tennae are 12-jointed, brownish Incidence of parasitism.—None black, and with the pedicel about located. one-third longer than the first Life history and habits.—SO: joint. Posterior angles of the ■"' Eggs, larvae and pupae unknown. metathorax are prominent and : Adults occur from May 1 to the subacute. Wings and veins are . last part of June. LOD and OWS : hyaline and the stigma is slightly Unknown. G/Y: Not given. SA: tinged with brown. The abdomen Eggs. N/H: One. MOA: Insert is scaly and wrinkled and the eggs in hosts' eggs. AF: May in- venter of the female is also often gest the contents of the host's wrinkled. (10, p. 119; 11, p. egg. HAB : Unknown. Larvae are 243) generally located in the eggs at Discussion.—Although S. hy- the lower end of the egg pod and alinipennis utilizes four hosts, it emergence is delayed until the appears to be restricted to Flor- , young grasshoppers are partially ida, possibly by environmental grown. (160, p. 270) factors. Description.—Eggs, larvae, and pupae unknown. Adults are 4 to Scelio luggeri Riley 4.5 mm. long, black, coarsely 1893: 242-246 wrinkled, and punctate. The mes- onotum lacks furrows, whereas Synonymy.—None (in As\i- the face is striate above the in- mead 1893) (11, pp. 2^2, 246-,225, ; sertion of the antennae and be- p. 702). , Recorded distribution.—Min- * nesota (225, p. 702). (Map 40.) Host species.—Melanoplus san- guinipes (Fabricius), M. spretus (Riley), and Melanoplus STp-p. (11, - p. 246; 123, p. 488; 264; 325, p. 14; 329, p. 630). ' Incidence of parasitism.—Not " given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. ' Adults not given. LOD and OWS : Unknown. G/Y: Not given. SA:

MAP 39.—Recorded distribution of Eggs. N/H: One. MOA: Insert Scelio hyalinipennis Ashmead. eggs in the hosts' eggs. AF : May ARTHROPOD AND NEMATODE PARASITES IN AMERICA 83 Scelio oedipodae Ashmead ^S^/^*^ 1893: 242, 246 Synonymy.—None {225, p. 703). Recorded distribution.—Vir- ginia {11, pp. 2U2, 2A6; 225, p. 707). (Map 41.) Host species.—Chortophaga vi- ridifasciata (DeGeer), Oedipodi- nae spp., Melanoplus differentialis (Thomas), and Schistocerca ob- scura (Fabricius) {11, pp. 24-2, MAP 40.—Recorded distribution of Scelio luggeri Riley. 2Jf6; 123, p. i88; 22Jf, pp. 23-2A). Incidence of parasitism.—None ingest fluid from host's egg. HAB : located. Unknown. {336, p. 31 ) Life history and habits.—SO : Description.—Eggs, larvae, and Eggs, larvae, and pupae unknown. pupae unknown. Adult females Adults first appear from May 1 are 3 to 3.2 mm. long, black ex- to late June. LOD and OWS : Un- cept the scape and legs, which are known. G/Y: Not given. SA: brownish yellow, and the head and Eggs. N/H: One. MOA: Insert antennae, which are brown to eggs in hosts' eggs. AF: May in- black. The body surface is wrin- gest fluid of grasshopper egg. kled and there are two furrows on HAB : Unknown. Usually found in the mesonotum. The pedicel of the those eggs of the host lowest in antenna is distinctly longer than the egg pod. Emergence is much the first flagellum joint; joints later than hatching of host's eggs. 2 to 4 are very short. There are {22^, pp. 23-2A) four complete longitudinal me- dian ridges on the metanotum. The abdomen is fusiform and about one-third longer than the head and thorax combined. Wings are not quite spindle shaped. They are pale at the base and contain a stigma with a short stigmal vein. {11, p. 2A6) Discussion.—Since its host spe- cies are of economic importance and are and were widespread, I believe its distribution to be con- siderably larger than in Minne- MAP 41.—Recorded distribution of sota. Scelio oedipodae Ashmead. 84 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Description.—Eggs, larvae, and pupae unknown. Adult wasps are 3.5 to 4.0 mm. long and black. The integument is wrinkled. The pedi- cel and legs are brownish yellow. In the male the antennae are brown and the face is vertically striated. Femora are brownish and the tibiae and tarsi are yel- lowish. There are longitudinal grooves on each side of the meso- scutellum, especially noticeable in the male but wanting anteriorly MAP 42.—Recorded distribution of in the female. Metathorax is trun- Scelio ovivorus (Riley). cate behind. The abdomen is longer than the head and thorax combined and is striated. The Life history and habits.—SO: first segment is well separated Eggs, larvae, and pupae unknown. from the second by a strong con- Adults not given. LOD and OWS : striction and is twice as wide as Unknown. G/Y: Not given. SA: > it is long. Wings are hyaline. (11, Eggs. N/H: One. MOA: Insert . pp. 242, 2i6) eggs in hosts' eggs. AF : May in- Discussion.—Why it has not gest contents of grasshopper eggs. been reported from a larger geo- HAB: Unknown. (2W, p. 306; graphical region is puzzling, es- 265, p. 270) pecially when its host range in- Description.—Eggs, larvae, and cludes at least four species from pupae unknown. Adults are 3.6 . a wide geographical area. to 4.2 mm. long. Females are black, very coarsely wrinkled, and Scelio ovivorus (Riley) punctured. The mesonotum lacks furrows and there is a large pol- (Caloptenobia) 1878: 307 ished space on the face above the Synonymy.—Caloptenobia ovi- antennae with coarse lines con- ^ vorus Riley (225, p. 703). verging toward the mandibles on Recorded distribution.—South- either side of the face. Antennae , ern Canada and Massachusetts are 12-jointed, brownish black; (225, p. 703). (Map 42.) the scape and pedicel are pale Host species.—Dissosteira car- rufous or brownish yellow. The otina (Linnaeus) and Melanoplu^ thorax is very coarsely rugose, ; spretus (Walsh) (123, p. A88; except for fine sculpturing on the 26A, p. 306; 265, p. 270). metathorax. Tegulae are yellowish Incidence of parasitism.—None as are the legs. The abdomen is given. fusiform and the dorsum and ARTHROPOD AND NEMATODE PARASITES IN AMERICA 85 venter are coarsely striated longi- brownish yellow. Wings are hya- tudinally. The first segment is line. The submarginal vein termi- short, transverse, and slightly nates in a rounded stigma with a longer than half the length of the short stigmal vein. The abdomen second. Wings are subf uscous and is distinctly striated and the api- the stigma of the submarginal cal edges of the segments are vein is large. The head and thorax smooth. (10, p. 119; 26^, p. 306) of the male are coarsely wrinkled, Discussion.—The gap between although the thorax lacks fur- eastern Canada and Massachu- rows. Antennae are pale brown setts may indicate incomplete dis- and 10-jointed. The legs are tributional, records here.

INTERNAL PARASITES Nematoda Mermithidae Identification of nematodes is extremely difficult, primarily be- At least three species of mer- cause of the limited taxonomic mithids (Nematoda) are para- characters that are available and sitic in grasshopper nymphs and reliable and because most of the adults. Several other species are keys refer to adult male speci- recorded in the literature from mens. The description of A. mel- Canada and the United States, anopli, on the other hand, is for but, according to Poinar (38^), the immature stage and thus adds these are synonyms of the three to the difíiculty. Even when ma- valid species named here. Mermis terial is in excellent condition, the nigrescens Du jar din does not in- experienced taxonomist has diffi- habit North America and there- culty in making positive identifica- fore the references attributed to tion. Therefore descriptions are it probably belong to M. subni- not included and the reader is ad- grescens Cobb, a very closely re- vised to utilize life-history and lated species. Mermis acuminata habit information when attempt- is a synonym of Agamermis de- ing to identify these species. caudata Cobb, Steiner, and The distribution of these nem- Christie. The remaining species, atodes is limited to the Midwest, including A. pachycethala S., Hexamermis spp., and M. albicans Northeast, and spotted western are thought to belong to A. de- locations. Adequate moisture is caudata, Agamospirura melanopli apparently necessary for success- Christie, and M. subnigrescens, ful development. Where present, the three nematodes associated nematodes can be very effective with grasshoppers. controlling agents, infesting more 86 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE than 60 percent of preferred Host species.—Aeropedellvs grasshopper species. However, clavatus (Thomas), Camnula pel- most grasshoppers of economic lucida (Scudder), Chorthippus importance are found in crop and longicornis (Latreille), Melano- rangelands that become extremely plus hivittatus (Say), M. daw- dry at various times because of soni (Scudder), M. femurrubrum low rainfall or low relative hu- (DeGeer), M. gladstoni Scudder, midity, a situation unacceptable M. infantilis Scudder, M. pack- to nematode development. There- ardii Scudder, M. sanguinipes fore the effectiveness of nema- (Fabricius), the long-horn grass- todes as grasshopper controlling hopper Conocephalus saltans agents is greatly restricted to (Scudder), and the cricket Acheta very localized areas by strict mois- assimilis (Fabricius) (27, pp. 73- ture limitations. (22, p. 268) 76; 137, pp. 521-522; 307, pp. 253-256). Agamermis decaudata Cobb, Incidence of parasitism.—None Stiener, and Christie located. 1923: 924-926 Life history and habits.—SO: Synonymy.—Mermis acumi- Eggs are deposited in the spring. nata (^38^). Larvae are in the field in May and Recorded distribution.—South- June. Adults occur from fall to eastern Canada and northeastern spring. LOD: Eggs generally re- United States, south through Vir- quire 50 to 60 days or more until ginia, west through Nebraska, warm weather. Larvae need about and north through North Dakota 80 days within the host, followed (27, pp. 73-76; 51, pp. 161-198). by 9 to 11 months in the soil. (Map 43.) Adults live for 2 or possibly 3 years. OWS : As larvae and adults. G/Y: One generation lasts 2 to 4 years. SA: Generally very young nymphs. N/H : One to six or more. MOA: Penetrate the integument of the host. AF: None. HAB: After leaving the host, the nema- tode migrates 3 to 20 cm. deep into the soil where it remains un- til its death years later. Both the eggs of the host and the parasite hatch about the same time so that the nematode nymphs are able to attack young grass- MAP 43.—Recorded distribution of Ago- mermis decaudata Cobb, Steiner, an-l hopper nymphs. Penetration Christie. through the integument requires ARTHROPOD AND NEMATODE PARASITES IN AMERICA 87

2 to 10 minutes, during which Life history and habits.—SO: time a part of the nematode tail Eggs and larvae unknown. Adults is broken off and left outside. not given. LOD, OWS, G/Y, SA, Upon exit from the host, the para- N/H, MOA, and AF: Unknown. site enters the soil and the host HAB: Pastures. {50, pp. 127- dies. The larva locates a spot, con- 130; 177, p. 179) structs a cell, and remains there Descriptions.—The reader is until the following summer when directed to the work of Christie it molts to an adult. Eggs are de- for the nymphal description {50, posited on the walls of the cell pp. 127-130). until the advent of cold tempera- Discussion.—A. melanopli has tures. Egg laying also commences only been described in the larval early the following spring and stage and only one report was lo- continues through the summer. cated pertaining to research on Very few live to the following any phase of this species. Only a summer. {27, pp. 73-76; 51, pp, few specimens were mentioned in 161-198; 52, pp. 35S-S5Jf; 53, p. it. Therefore it is concluded that k6; 58, pp. 921-926; 152, p. U; A. melanopli is insignificant as a 300, p. 60; 306, pp. 28-33; 307, parasite of grasshoppers. pp. 255-256; 308, pp. 187-199; 323, pp. 90, 93, 98, Jf6Jf) Mermis subnigrescens Cobb Description.—For a complete 1926: 67-69 description, the reader is directed Synonymy.—Mermis nigres- to the work of Christie, who gives cens Dujardin {387). development by stages {51, pp. Recorded distribution.— 161-198). Throughout northeastern United Discussion.—This parasite is States from the Atlantic coast effective in controlling grasshop- west through Minnesota, Iowa, pers in localized areas. Its dis- and Missouri, north into Quebec tribution is limited by the timing and Manitoba, and south as far and amount of yearly moisture. as Washington, D.C. {52, pp. 353- 365; 53, pp. Jf3-Jf8). (Map 44.) Agamospirura melanopli Host species.—Arphia sul- Christie phur ea (Fabricius), Camnulapel- 1928: 127-130 lucida (Scudder), Chorthippus Synonymy.—None {387). longicornis (Latreille), Chorto- Recorded distribution.—Not phaga viridifasciata (DeGeer), given in the literature. (Bur- Host species.—Melanoplus fe- meister), Melanoplus bivittatus murruhrum (DeGeer) {50, pp. (Say), M. femurrubrum (De- 127-130). Geer), M. luridus (Dodge), M. Incidence of parasitism.—None sanguinipes (Fabricius), Orphu- located. lella pelidna (Burmeister), 0. 88 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE posited on the vegetation and in- gested when the host consumes it as food. Eggs hatch within the gut and the larvae migrate into the hemocoel through the gut wall. AF : Unknown. HAB : Moist pas- ture-grass type complex. Oviposition occurs in the spring. Rain and continued mois- ture are the stimuli that cause the gravid females to climb vegeta- tion where the eggs are laid. MAP 44.—Recorded distribution of Christie (52) reported that one Mermis subnigrescens Cobb. gravid female 85 mm. in length contained approximately 1,400 speciosa (Scudder), Romalea mi- eggs. The females continue laying croptera (Beauvais), Spharage- eggs until they complete oviposi- mon collare (Scudder), and two tion or the weather becomes dry, species of Tettigoniidae (27, pp. when they fall to the ground and 73-76; 52, pp. 353-365; 53, pp. migrate into the soil. The arrival A3-U). of moist conditions again stimu- Incidence of parasitism.—Up lates climbing and oviposition. to 10 percent (52, pp. 353-365). The length of life of the adult fe- Life history and habits.—SO: male that completes oviposition Eggs can be found on vegetation was not determined by Christie, in early spring to early fall. Lar- but those that did not oviposit re- vae (young) are within the host mained alive for 2 years. beginning about early June and Eggs remain viable on the fo- remain until fall. Adults enter the liage throughout the summer. soil from middle to late fall and When the eggs are ingested, the remain until they die several digestive juices of the gut dis- years later, generally during the solve parts of the nematode's egg- winter. LOD : Eggs are in the soil shell, allowing the escape of the over winter. Larvae need several worm. Within a short time, it months within the host, then they migrates through the gut wall molt several months after leaving into the hemocoel where it is free the host and enter the soil. Adults living. Since the eggs are viable Unknown. OWS: As larvae and throughout the summer, hosts adults. G/Y: One generation ex- may be infected many times and tends from 2 to 4 years. SA : All. nematodes of various stages of N/H: Generally only one or two, development may exist within a although 40 or more have been single host. Males usually remain recorded. MOA: Eggs are de- within the host from 4 to 6 weeks, ARTHROPOD AND NEMATODE PARASITES IN AMERICA 89 whereas females require 8 to 10. Adults usually are found 6 to 8 As the mature larvae complete inches below the soil surface, al- the feeding stage, they leave the though some have been seen as host and migrate into the soil deep as 24 inches. They are not (fig. 12). Emergence from the gregarious and are usually found host results in its death. Many of singly except during copulation. the immature nematodes that are As the eggs mature within, the forced to emerge prematurely female becomes brownish and by eventually die in the soil, possibly September appears black ex-ipt because of starvation. for a small area at the extremities The nematode overwinters as a of the body. mature larva and the final molt As the female crawls and en- occurs the following spring. twines around grass stems, it

■4 ^*:w^

PN-2730 FIGURE 12.—Nematode emerging from host. 90 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE leaves behind a trail of eggs. ened areas formed by the attach- These eggs contain an area of at- ment of an entangled appendage tachment, called a byssus, which called a byssus, the size and shape aids in fastening them to the veg- of which varies. Color of the outer etation. All types and sizes of veg- shell is lighter at the poles than etation are utilized. (27, pp. elsewhere. This area becomes 73-76 \ 52, pp. 353-365; 53, pp. transparent as digestion occurs ^3-í8 ; 56, pp. 66-72 ; 57, pp. JÍ79- within the grasshopper gut. {56, ^80;125, pp. 12-15; 132, pp. 521- pp. 67-69) 522; 152, p. U; 179, p. 179; 307, Discussion.—The effectiveness pp. 253-256; 308, pp. 187-199; of this parasite on grasshoppers 323, pp. 89, 9i, 98, i6Jf) is dependent primarily on climatic Description.—Because of the conditions. During dry weather difficulty of , the reader very few eggs are deposited and is directed to the works of Cobb very few hosts are parasitized. {56). One of the outstanding However, when the spring is characteristics of the nematode is moist, allowing more oviposition, its egg. The egg measures ap- this parasite exerts a very definite proximately 50 to 56 ¡JL and is effect on reducing the grasshop- composed of an inner shell cover- per population in the areas where ing that is brown. The outer shell it is found. is divided into two cuplike parts The life history is very similar by a groove around the circum- to that of Agamermis decaudata ference. At the poles are thick- but differs in the oviposition site.

Gordiacea The systematic position of the been little work done with those horsehair worms, hair snake- associated directly with grass- worms, or Gordian worms (from hoppers other than notes of occur- the Gordian knot of ancient Greek rence. C. V. Riley {262) reported mythology) is somewhat confused Gordius seta and G. robustus and has been for many years. The within the Rocky Mountain lo- most popular present-day con- cust {Melanoplus spretus cepts put them in either the class (Walsh) ) and several others, such Gordiacea or Nematomorpha and as G. villoti Rosa and Gordius separate them from the Nema- spp., have been reported as at- toda. There are 80 known species, tacking Orthoptera {321). G. one of which is marine in nature. robustus has also been reported Horsehair worms attack many as attacking Anabru^ simplex hosts, but they are mostly noted Haldeman {329). However, since from crustaceans, , crick- specific identification of these ets, and grasshoppers. There has worms is extremely diflScult, many ARTHROPOD AND NEMATODE PARASITES IN AMERICA 91 incidences probably go unre- how the worms enter the host. ported. One possibility is that the worm Gordiacea occurs from the hot encapsulates itself, is ingested tropical areas to the cold temper- with the food of the host, emerges ate climates, including some within the alimentary tract, and mountain areas above timberline. burrows its way into the hemo- Full-grown worms range from 6 coel. It is more probable that the to 48 inches in length but are less worm penetrates directly into the than 1 mm. in diameter. Worms hemocoel of the host through the are dioecious and the digestive integument by means of a pro- tract is vestigial. Males are trusible proboscis, which is armed smaller than the females and are with spines. Once in the host the more rounded at the posterior worm develops through an un- end. Adults can be found in known number of molts over sev- ponds, brooks, or other quiet wa- eral months. Food is absorbed di- ter and even in damp soil from rectly and not in the gut, and October to May. Males are gen- tissues of the host are dissolved erally active, swimming about in in the vicinity of the worm. The whiplike fashion, whereas the fe- worm leaves in the fall and ap- males simply writhe. Copulation parently only when the host is occurs in the spring, about May, near water. This emergence gen- and the eggs are deposited in the erally kills the host, although water in gelatinous strings. The some hosts apparently drown. eggs hatch over many months. Gordiacea is common only in The young larvae swim about in areas near water. Therefore its the water for a time and then effect on grasshopper populations apparently make their way to is extremely limited. More study vegetation near the water's edge. is needed to determine its dis- There is some question as to tribution and abundance.

EXTERNAL PARASITES

Hymenoptera Sphecidae ilies. Hosts include spiders, lo- custs, grasshoppers, cockroaches, This is a large family of wasps, most of which either make their and lepidopterous larvae. Twenty- nest in the soil or build mud cells nine species of the genera Solie- on foliage or other ridged frame- relia, Sphex (fig. 13), Tachysphex, work. Many are host specific in and Tachytes are recorded attack- their attacks, but some utilize ing grasshoppers in Canada and members of several related fam- the United States. The adults 92 TECHNICAL BULLETIN NO. 1460, U. S. DEFT. OF AGRICULTURE

^ ^'^B,"*

PN-2731 FIGURE 13.—Adult of Tachysphex sp.

themselves are not parasitic but establish its location. Once near take the prey to be utilized by the the entrance of the burrow, the , young larvae, which develop as wasp completes the design of the external parasites. The adults cell to fit the size and shape of the ' construct a burrow or nest within host. It then drags the host inside, , the soil. The angle, size, and shape deposits an egg usually near the - are characteristic for the species. attachment of the femur to the Either before or immediately body, fills in the entrance with , after the constructing of the bur- soil, and camouflages it. row, the female wasp seeks a prey Upon hatching, the larva be- - of a particular size. Some species gins to consume the host, which - utilize several small prey, whereas is generally paralyzed but still ' others require only a large one. alive. Sometimes the host is killed If the burrow is constructed first, purposely by severing the head , the wasp apparently gains its ini- from the body prior to departure tial bearings by flying around the of the female wasp. Upon comple- ■ open nest prior to searching. tion of development, the larva After stinging and paralyzing the leaves the dead remains of the prey, it drags it back to the bur- host and pupates within the cell. row, often taking to the air to re- Little is known about most spe- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 93 cies. As a group, however, they Since larval descriptions are mea- may exert considerable effect on ger or nonexistent, a key is given reducing grasshopper populations, only for adults.

KEY TO ADULT SPHECIDS la. Abdomen with long petiole ; middle tibiae with two apical spurs 32 lb. Abdomen petiolate, but somewhat sessile; middle tibiae with only one spur 2 2a. Hindocelli obsolete or indicated by scarlike structure ; mandibles usually emarginated on underside 3 2b. Hindocelli normal, distinct; second cubital cell petiolate or triangular; marginal cell truncate with appendage; hindtibiae smooth, not serrated 4 3a. Body stout, no notch at base of anterior femora of male; tarsi of female lacking long flexible spurs; pygidium hairy 6 3b. Anterior femora of males notched near base beneath ; female tarsi have long, flexible spines; hypopygium is naked 18 4a. Abdomen of both sexes entirely black Solierella plenoculiodes similis (Bridwell) 4b. Abdomen red 5 5a. All tibiae of females with white stripe exteriorly Solierella striatipes (Ashmead) 5b. All tibiae black; third and fourth antennal joints equal Solierella fossor (Rohwer) 6a. Appearance beelike; thorax of female densely pubescent; forecoxae of male simple 7 6b. Appearance not beelike; thorax of female not densely pubescent; fore- coxae of male with elongated process 12 7a. Female 8 7b. Male 9 8a. Abdomen red in part; head and thorax clothed with dense, silvery pubescence Tachytes fulviventris fulviventris Cresson (in part) 8b. Abdomen black with four silvery bands dorsally; front silvery; legs black; tibiae and tarsi with silvery pile; apical part of tarsi ferrugineous Tachytes pepticus (Say) (in part) 9a. Basal joints of flagellum rounded out beneath; apical joints normal; abdominal segments with transverse reddish band Tachytes exornatus (Fox) 9b. Basal joints of the flagellum not rounded out (except for Tachytes pepticus) or scarcely so beneath; aplical joints abnormal 10 10a. Two apical joints of antennae narrow and elongate, much narrower than joints 9 to 11, which are noticeably broader Tachytes fulviventris fulviventris Cresson (in part) 10b. Apical joints of antennae normal 11 11a. Base of mandibles covered with white pubescence; face with silvery hair; 10th, 11th, and 12th joints of antennae normal Tachytes sayi Banks lib. Base, of mandibles lacking white pubescence; face with little or no silvery hair; antennal joints 10 and 11 widened, 12 narrow Tachytes pepticus (Say) (in part) 94 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

12a. Female 13 12b. Male 17 13a. Greater part of femora reddish 14 13b. Greater part of femora or legs entirely black 15 14a. Apex of first three dorsal segments only with silvery pile Tachytes distinctus (Smith) (in part) 14b. Apex of first four dorsal segments only with silvery pile Tachytes elongatus elongatus Cresson 15a. Space between eyes at top about equal to length of antennal joints 2 and 3 combined; abdomen reddish or red and black Tachytes abdominalis (Say) 15b. Space between eyes greater than antennal joints 2 and 3 combined 16 16a. Metanotum furrowed Tachytes obductus Fox 16b. Metanotum not furrowed Tachytes rufofasciatus Cresson (in part) 17a. Wings yellow; apical areas of wings dark; apex of femora, tibiae, and tarsi yellow ferrugineous Tachytes distinctus Smith (in part) 17b. Abdomen and legs black; body not densely silvery sericeous; wings not yellow; metanotum slightly furrowed Tachytes rufofasciatus Cresson (in part) 18a. Female 1Ö 18b. Male 28 19a. Color is dark ferrugineous except head, which is black; entire body is covered with silvery pubescence, which nearly obscures ground color; heart-shaped impressed space caudad of posterior ocelli is shiny and possesses medial longitudinal depression ._Tac%sp/iea; propinquus Viereck 19b. Specimen not as above 20 20a. Space between eyes at top about equal to antennal segments 2 and 3 combined 21 20b. Space between eyes at top greater than antennal segments 2 and 3 com- bined 26 21a. Abdomen entirely red; metathorax strongly reticulated above Tachysphex texanus (Cresson) (in part) 21b. Abdomen not entirely red 22 22a. Metathorax coarsely wrinkled above, abdomen about equally red and black Tachysphex quebecensis (Provancher) (in part) 22b. Metathorax not coarsely or strongly reticulated or wrinkled above, at most finely striated or granulated 23 23a. Greater part of abdomen reddish 24 23b. Abdomen black or equally black and red 25 24a. Metathorax above finely granulated; front rather coarsely granulated; dorsulum shining and with separate punctures Tachysphex semirufus (Cresson) 24b. Dorsulum very closely punctured; front finely granulated; metathorax above finely leathery Tachysphex tarsatus (Say) (in part) 25a. Apical abdominal segment red or black, silvery pubescence as usual Tachysphex similis Rohwer 25b. Thorax and abdomen nude of silvery pubescence, first three abdominal segments or first 2% segments bright red Tachysphex tenuipunctus Fox 26a. Anterior margin of clypeus produced medially into short lobe 27 26b. Anterior margin of clypeus not as above ; usually last two segments red Tachysphex terminatus (Smith) (in part) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 95

27a. Last abdominal segment red Tachysphex fusus Fox (in part) 27b. Abdomen all red Tachysphex plenoculiformis Williams 28a. Abdomen red or black and red 29 28b. Abdomen entirely black, except for maybe posterior apex 30 29a. Space between eyes at top distinctly greater than length of antennal joints 2 to 4 combined; anterior margin of clypeus slightly rounded out, not produced medially Tachysphex terminatus (Smith) (in part) 29b, Space between eyes at top not greater than length of antennal joints 2 to 4 combined; clypeus produced medially Tachysphex fusus Fox (in part) 30a. Metapleurae coarsely striated Tachysphex texanus (Cresson) (in part) 30b. Metapleurae not coarsely striated 31 31a. Space between eyes at top about equal to joints 3 and 4 combined; abdomen entirely red Tachysphex tarsatus (Say) (in part) 31b. Space between eyes at top less than combined length of antennal joints 3 and 4; metathorax coarsely wrinkled above Tachysphex quebecensis (Provancher) (in part) 32a. Abdomen red or yellowish red 33 32b. Abdomen black, purple, or blue 3g 33a. Pile silvery to gray 34 33b. Pile not silvery or grayish 35 34a. Wings hyaline Sphex'thomaeFahricius 34b. Wings black with violet iridescence Sphex striatus (Smith) 35a. Pile yellow Sphex pubidorsus (Costa) 35b. Pile black Sphex rufiventris rufiventris Cresson 36a. Spiracles of first abdominal segment set on tubercles; color bronzed o/it. ^ P^rpl® Sphex aerarius Patton dob. Color black; spiracles not on tubercles 37 37a. Part of thorax shining gold color Sphex ichneumoneus ichneumoneus (Linnaeus) 37b. Thorax entirely black 33 Females of medium size, about 17 mm. long Sphex atratus LePeletier 38b. Females large, 25 to 28 mm. long Sphex laviventris Cresson

Solierella fossor (Rohwer) Life history and habits.—SO: (Niteliopsis) 1909: 116 Eggs, larvae, and pupae unknown. Synonymy.—Niteliopsis fossor ^^^^^^ not given. LOD : unknown. Rohwer, N. fosser Rohwer, N. ^^S and G/Y: Not given. SA: foxii Viereck {225, p. 9^3). Nymphs. N/H: One. MOA: Cap- Recorded distribution.-^Co\0' t^^es, stings, and paralyzes its rado. New Mexico, and Kansas P^^^- ^^ ^^^ HAB: Not given. {225, p. HS). (Map 45.) (^^^) Host species.—Species of Oedi- Description.—Eggs, larvae, and VoaindLQ {128, p. JÍ89\ 257, p. 185 \ vnpae unknown. Adult females 868, pp. 209-211 ). are 5 mm. long and basically black Incidence of parasitism.—None except the abdomen, which is red. located. Tarsi are reddish and very 96 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE gon and California {225, p. 9^3). (Map 46.) Host species.—Grasshoppers {123, p. ^89; 366). Incidence of parasitism.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS, G/Y, SA, N/H, MOA, AF, and HAB : Not given. Description.—Eggs, larvae, and MAP 45.—Recorded distribution of pupae unknown. Adults are 6.5 Solierella fossor (Rohwer). mm. long, basically black, and covered with an appressed silvery pubescence, which is very con- sparsely covered with a silvery- spicuous on the clypeus and face. pile. Anterior margin of the cly- The tegulae, tarsi, and margins peus has an acute "V"-shaped of the abdominal segments are tooth. Head is granular with a brownish with some silvery pubes- front groove. Antennae are slen- cence on the base and sides of the der; the third joint is slightly tergites and the sides of the pro- longer than the fourth. Prothorax podeum. The clypeus is strongly is rounded above and slightly carínate and the eyes are strongly notched. Scutellum is slightly im- convergent above. The front is pressed with a few striae at the strongly and closely punctured, base of the metanotum. Abdomen becoming striated in the depres- is shining and finely punctured. Pygidial area is indistinct. Wings are hyaline with the margin some- what dusky and the veins brown. Males are 4.5 mm. long and simi- lar to the females except for a spine beneath the third and fourth antennal joints and a white stripe on the external area of the pos- terior tibia. (27h VV- 109-116) Solierella plenoculiodes similis (Bridwell) (Silaon) 1920: 402-403

Synonymy.—Silaon similus MAP 46.—Recorded distribution of Sol- Bridwell {225, p. 9A3). ierella plenoculoides similis (Brid- Recorded distribution.—Ore- well) . ARTHROPOD AND NEMATODE PARASITES IN AMERICA 97 sion preceding the front ocellus. The head behind the ocelli and the occiput are strongly striate-punc- tate. The pronotum is short and notched in the middle and the mesanotum is closely punctured. The abdomen is shiny; the first tergite is strongly and finely punc- tured and the remainder to a less extent. Wings are subhyaline and the veins yellowish. The stigma and costa are black. (28, pp. k02-k03) MAP 47.—Recorded distribution of Discussion.—The economic im- Solierella striatipes (Ashmead). portance of the hosts is unknown. have a silvery pubescence. Legs Solierella striatipes (Áshmead) are black and the tarsi somewhat (Niteliopsis) 1899: 9 reddish with a brownish base. All tibiae contain a white stripe on Synonymy.—Niteliopsis stri- the outer side. Wings are hyaline. atipes Ashmead (225, p. 94-2). (13, p. 9; 274, p. 108) Recorded distribution.—Cali- Discussion.—M. ligneolus is of fornia (225, p. 9Jt2). (Map 47.) minor importance. Host species.—Melanoplus ligneolus Scudder (123, p. A89; Sphex aerarius Patton 229, p. 958). 1879: 133-134 Incidence of parasitism.—Not Synonymy.—Sphex coerulea given. Drury, Chlorion cyaneum Dahl- Life history and habits.—SO: bom, C. aerarium Patton, S. Eggs, larvae, and pupae unknown. (Chlorion) nearcticus Kohl, S. Adults not given. LOD : Unknown. (Chloriort.) occultus Kohl (225, OWS, G/Y, SA, and N/H: Not V.974). given. MOA: Adults capture, Recorded distribution.—United sting, and paralyze their prey. AF States (225, p. 974). (Map 48.) and HAB: Not given. (371, pp. Host species.—Melanoplus 355-417) spretus (Walsh), crickets, and Description.—Eggs, larvae, and spiders (257, p. 44; 264, pp. 318- pupae unknown. Adults are 6Va 319; 330, p. 177; 331, p. 160). mm. long, the head and thorax are Incidence of parasitism.—None black, and the abdomen is red. located. Both the head and thorax are Life history and habits.—SO: finely and closely punctured. Face, £70^0^8, larvae, and pupae unknown. temples, pronotum, and pleura Adults not given. LOD : Unknown. 98 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE brown as is the clypeolabral su- ture. Sides of the maxillae and " the base of the labium are blotched with brown. There are scattered punctures on the clypeus and a few on the lower area of the head. <: Both mandibles and maxillae with the palpi are stout. The surface of the mouth parts in general is cov- ' ered with scattered punctures, ' some of which have strong setae. {98, pp. 2JÍ2-2U). Pupae un- - MAP 48.—Recorded distribution of known. ^ Sphex aerarius Patton. Adult wasps are 25 mm. long and bronzed purple. Antennae, OWS and G/Y: Not given. SA: mandibles, tegulae, and feet are ^ Young grasshoppers. N/H: Not black and the postscutellum is given. MOA : Stings prey paralyz- blue. Head and thorax are smooth - ing it, buries it in cells in the soil, with scattered and shallow pune- ^ and deposits a small egg at the tures. The metathorax is trans- juncture of the hindfemur and the versely striate throughout. Spir- body. It then departs and seals the acles of the first segment of the i entrance. AF: Apparently likes abdomen are situated on tuber- willow sap. HAB: Not given. cles. The clypeus terminates in a (153, pp. 2h6-2k7; 2U2, pp. 85- row of five teeth ; the lateral teeth ; 93; 2Jf3, pp. 256-261) are more remote from the inter- Description.—Eggs unknown. mediate teeth than are the inter- Larvae in diapause are 25 mm. mediate teeth from the median , long and cylindrical, tapering to- tooth. {75, p. 211; 238, pp. 133- ' ward both ends, more gradually 13^) so anteriorly. The integument is Discussion.—Although this . covered with minute spinules species will attack grasshoppers, and the dorsum is transversely it appears to prefer crickets and < wrinkled. Thoracic spiracles are spiders. smaller than the abdominal spir- Sphex atratus LePeletier acles. Head is blotched with brown above and below. The an- 1845: 355 tennal orbitals in the depression Synonymy.—Chlorion atratum areas in the center of the front (LePeletier), Priononyx brun- ' and on the clypeus and the pari- ñipes Cresson, Sphex atrata Le- • etal bands are strongly pigmented. Peletier, S. labrosa Harris {225, The median apical part of the p. 973). labrum and mandibles are a dark Recorded distribution.—Most ARTHROPOD AND NEMATODE PARASITES IN AMERICA 99 of Canada and the United States 308-310 ; 330, p. 179 ; 331, p. 189 ; including Alaska. Also in Mexico. 369, pp. 227-230). {97, pp. 178-181; 225, p. 973; Incidence of parasitism.—None 306, pp. 28-33; 369, pp. 227-230). located. (Map 49.) Life history and habits.—SO: Host species.—Ageneotettix de- Eggs, larvae, and pupae unknown. orum deorum (Scudder), Arphia Adults are in the field from June corinata Scudder, A. xanthoptera to September. LOD : Eggs require Burmeister, Aulocara elliotti 3 to 5 days. Larvae and pupae un- Thomas, Dissosteira Carolina (Lin- known. Total development per naeus), D. longipennis (Thomas), summer generation is less than 1 Melanophis arizonae Scudder, M. month. Adults, unknown. OWS: angustipennis Dodge, M. bispino- Not given. G/Y: Several. SA: sus Scudder, M. bivattatm (Say), Adults and nymphs. N/H: One. M. devastator Scudder, M. differ- MOA: As the female searches entialis (Thomas), M. femoratics for a victim, it may fly over a (DeGeer), M. femurrubrum (De- small area or rest for time to time Geer), M. foedus Scudder, M. lak- on vegetation until one is located. inus (Scudder), M. sanguinipes It apparently attacks only moving (Fabricius), M. spretus (Walsh), hosts. When the prey is captured, Mermeria neomexicana (Scud- it is held within the wasp's legs der), Pardalophora phoenicoptera and mandibles while it is stung Burmeister, and several times. The grasshopper citrina Scudder (97, pp. 178-181 ; becomes paralyzed and is stung a 101, pp. 137-14-2; 123, pp. U7U, few times more before being re- U89 ; 2Jtl ; 252, p. 23 ; 257, pp. 24, leased. The sting may occur any 62, 159-175; 264, p. 318; 303, pp. place on the body from the ven- 8-11; 306, pp. 28-33; 322, pp. tral side of the abdomen to the dorsal neck membrane. The wasp may then feed on some of the exuded fluids from the puncture wounds and may cut off the jump- ing hindlegs with its mandibles. After a short period of rest and preening, the wasp seeks a suit- able spot to construct a burrow. AF and HAB : Adults frequent the plants of Asclepias, Aster mul- tiflorus, Melilotus, Petalostemon, blue veronica, buckbrush, iron- weed, and white snakeroot to feed MAP 49.—Recorded distribution of on the nectar of the flowers. These Sphex atratus LePeletier. same plants often serve as resting 100 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE places during the night and the ging is repeated several times un- heat of the day, as perches from til the burrow is complete and the which to search for prey, and, as grasshopper is directly in front. noted by Rau (257), locations for The wasp backs down the tun- courtship. nel, pulling the victim in after her. Courtship is apparently quite The cephalad end of a small egg is simple, consisting mainly of the cemented next to the hindcoxae of female waiting on a plant such as the grasshopper. In a manner as Melilotus and attracting males rapid as the excavation, the wasp within a 15-foot radius. Copula- scoops the earth chips from its tion is apparently of short dura- deposit pile and carries them into tion. the tunnel where they are de- S. atratus may sample several posited and pounded into place areas before it locates a suitable with the head and clypeus. It then spot for its burrow, generally chips pieces from the surrounding within a 5-foot radius of the vic- soil, places them on top, and taps tim. By using its mandibles, it them into position by standing on chips away chunks of earth. The its head and acting like a pile size and shape are characteristic driver. A completed burrow is of the species. These are then car- then difficult to locate unless, as ried out of the tunnel between the reported by Rau and Rau (257), forelegs and clypeus and deposited cinders of a foreign color are ac- in a small pile a short distance cidentally deposited over the tun- from the entrance. Smaller pieces nel entrance. and dust are kicked out by the Upon hatching, the larva be- hindlegs. When the burrow is only gins to feed through the mem- partly complete, the wasp rests brane above the hindcoxae with briefly, takes its bearings on the the body of the wasp at first re- surroundings, and seeks its vic- maining external and only the tim. It often may have difficulty head within the grasshopper's in relocating the paralyzed grass- body cavity. After a 4-day larval hopper, flying beyond it and back period, the wasp is ready to spin several times. It then straddles its cocoon. the prey. Facing in the same di- There is some disagreement as rection and grasping the antennae to the sequence of constructing in its mandibles and the head or and obtaining the host. Rau neck in its forelegs, it begins to (252), Williams (369), and Evans drag the grasshopper toward the (97) noted that the wasp first burrow. seeks its victim and then begins The wasp often leaves its vic- construction on the burrow, tim, returns to its burrow, and whereas Peckham (2J^2) and continues with the construction. Strandtmann (322) cited several This pattern of digging and drag- instances when the grasshopper ARTHROPOD AND NEMATODE PARASITES IN AMERICA 101 was parasitized after construction unicatus (Say) locates nests of S. had begun. Smith (303) reported atratus that have been closed, re- that in nearly, every instance the excavates them, enters, deposits grasshopper that is placed in the an egg of its own, and destroys nest is stung before the excava- the egg of S. atratus by crushing tion is undertaken, unless the it between its mandibles. For wasp is not satisfied and stings some reason it does not close the several prey during construction. burrow completely and often In this situation the last victim is leaves the work half completed. used to stock the nest and all (12, pp. 56-66; 97, pp. 178-181; others are discarded. Once stung, 101, pp. 137-H2; 238; 252, p. 23; the grasshopper seldom if ever 257, pp. 2k, 62, 159-175; 26Jf, p. will survive for more than 2 or 3 318; 300, pp. 8-11 ; 322, pp. 308- days. 310; 369, pp. 227-230) The burrow entrance is 6.5 to Description.—Eggs are slender, 15 cm. long by 1 cm. wide and ex- whitish, and strongly curved. tends down to a horizontal cell 4 They are 6 mm. long. Larvae to 9 cm. under the soil surface. (young) are 4 mm. long, increas- Each burrow differs in the length ing to 23 mm. just prior to pupa- of the tunnel and the angle of in- tion. Anterior section of the body cline, which Evans (96) indicated is slender and the pleural lobes as being individual and ecological are weakly developed. Minute variations. Cells are 2.5 to 5 cm. spines arise from platelike bases long and 1.2 to 1.8 cm. wide; the on the integument. Head is 1.15 size apparently is dependent on mm. wide and 1.25 mm. high, the size of the prey. The cell is light straw colored, and without large enough for a small air sup- blotches or other pigmented areas. ply but not large enough to allow The parietal bands and margin of the grasshopper to turn around. the labrum are scarcely pig- S, atratus is not without its mented, whereas the mandibles own parasites and predators. and pleurostomal and hypostomal Adams (2) reported Metopia leu- thickenings are brown. Each an- cocphala Rossi ( Sarcophagi dae) tennal orbit has three small sen- hovering about the nests prior to sory cones. There are a few small their completion. Williams (369) scattered punctures on both the reported a small fly that lurked front and clypeus. The labrum is about the construction of a S. about three times as wide as high atratus burrow. When the wasp and emarginated. Punctures are entered, the fly quickly deposited more numerous on the anterior larvae on the grasshopper, which margin than the posterior. There was to act as food for the S. atra- are four sensory pores on each tus larva. side of the epipharynx located The hjonenopteran Stizoides well back of the margin. The re- 102 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE mainder of the epipharynx is cov- ered with spines. Spinules on the side are directed forward, whereas those in the median area are directed posteriorly. On the apex of the mandibles are three large teeth and a smaller fourth at the base of these. (97, pp. 178-181 ; 101, pp. 137-H.2). Pu- pae unknown. Adult females are 17 mm. long and generally black. Head, thorax, and abdomen are all covered with MAP 50.—Recorded distribution of black hair. The metathorax is con- Sphex ichneumoneus icheumoneus cave and the abdomen is glossy. (Linnaeus). Feet and legs are black and con- tain hair and spines of the same Host species.—"Orthoptera," color. Hair around the anus is Melanoplus spretus (Walsh), and also black. Wings are black with many species of crickets (116; a violet reflection and scales are 26i; 267, pp. 37A-385). a ferrous black. {62; 6A; 75, p. Incidence of parasitism.—Not 213;202, p. 355) given. Discussion.—On occasion this Life history and habits.—SO: species may limit the grasshopper Eggs, larvae, and pupae not given. populations in a given area. It is Adults early summer. LOD: Ges- not host speciñc but seems to con- tation requires 2 to 3 days. Eggs fine its activities to grasshopper unknown. Larvae 6 to 9 days. hosts. Its own population appears Pupae and adults not given. OWS : to be restricted by certain re- Not given. G/Y: One. SA: Adults. quirements of soil type and con- N/H: Not given. MOA: Stings dition. prey to provision nest. AF and HAB : Not given. Sphex ichneumoneus Females begin digging their ichneumoneus (Linnaeus) nests within 3 days after mating. The nests are 5 to 12 inches deep (Apis) 1758: 578 and contain one to three compart- Synonymy.—Apis ichneumon- ments. When the nests are com- eus Linnaeus, Vespa jamaicensis pleted, or nearly so, the wasp Drury, ? Sphex ornata LePeletier seeks a prey, paralyzes it, and (225, p. 972). grasping the body with her fore- Recorded distribution.—South- legs and the antenna in her man- ern Canada and the United States dibles she flies or drags the prey (225, p. 972). (Map 50.) toward the nest. She drags the ARTHROPOD AND NEMATODE PARASITES IN AMERICA 103

victim until it is nearly touching clypeus. Mandibles are stout and the opening, releases it, and en- support four strong teeth near the ters the burrow, apparently for a apices. {101, pp. 139-140) Pu- final inspection. When satisfied, pae not given. Adults' rostum is the wasp drags the prey inside hard and straight. The tongue where she lays a single egg on the sheath is straight and the tip ventral side of the prey between forked. Antennae are black. Tho- the forecoxae and the midcoxae. rax is black with part a shining One or more prey may be stored golden color. Abdomen is black in each compartment of the nest, and petiolate. (116, pp. 1043- but only one egg will be found 1062) there. The wasp then leaves and Discussion.—Preference by far the hole is sealed with dirt from appears to be for crickets. the surface. ^ The egg hatches in 2 to 3 days Sphex laeviventris Cresson and the young larvae feed on the 1865: 463-464 chorion during the first day. Fully Synonymy.—Palmodes laevi- developed larvae usually remain ventris (Cresson), Sphex morio in the nest until the following Kohl, Epyris laeviventris (Cres- summer. (1, pp. 255-258; 110, son), Ischius laeviventris (Cres- pp. Jt58-Jt60; 116, pp. 1043-1062; son) (225, p. 974). 2U, pp. 33-41 ; 252; 264; 267, pp. Recorded distribution.—^West- 374-385; 323, p. 149) ern United States {225, p. 974). Description.—Eggs not given. (Map 51.) Larvae (mature) are 33 mm. long, Host species.—Stipator spp., fusiform, and pale yellowish probably stevensonii {225, p. 974 ; white. Spiracles are light brown 323, pp. 196-472; 369, p. 227). with a dark rim. The body lacks evident setae, but the integument appears granular. Thorax is strongly tapered toward the small head capsule. There is a rounded pleural lobe on each side of the abdominal segments. Thoracic spiracles are slightly smaller than abdominal spiracles. The head is straw colored, with the top and sides of the cranium a light blotchy brown. Mandibles are dark brown except the tips, which are nearly black. Palpi are also brown- ish. Epistomal suture is weak and MAP 51.—Recorded distribution of there are many punctures on the Sphex laeviventris Cresson. 104 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Incidence of parasitism.—None Sphex pubidorsus (Costa) located. (Enodia) 1862: 69 Life history and habits.—SO: Synonymy.—Enodia pubidor- . Eggs, larvae, and pupae unknown. sum Costa, Priononyx bifoveolata Adults not given. LOD : Unknown. Taschenberg, P. canadensis Pro- OWS, G/Y, SA, and N/H: Not vancher, Sphex excisus Kohl, ■- given. MOA: Adults attack, cap- Chlorion (Priononyx) bifoveola- ture, sting, and paralyze, then tum Fernald, C. pubidorsum Fer- bury and oviposit on host. AF and nald {225, p. 973). HAB : Not given. {178, pp. 7AS- Recorded distribution.—United ^ ÍS 3) States {225, p. 973). (Map 52.) Description.—Eggs, larvae, and Host species.—Melanoplus fern- - pupae unknown. Adults are large, urrubrum (DeGeer), M. scudderi robust, and completely shining Uhler, Trimerotropis citrina black. Females are 25 to 28 mm. Scudder, and Sciretetica marmo- > long, males approximately 21 raiaPicta (Scudder) {97, pp. 183- ^ mm. Abdomen is smooth and pol- 18Í ; 123, p. A89 ; 252, p. 23). ished. Females are sparsely cov- Incidence of parasitism.—None - ered with a black pubescence. The located. ; clypeus is indented on each side Life history and habits.—SO: anteriorly with a central truncate Eggs, larvae, and pupae unknown. ^ process. Mandibles are large and Adults not given. LOD : Unknown, i deeply grooved. Antennae are OWS and G/Y: Not given. SA: black and sericeous. Thorax above Adults and nymphs. N/H: One. is smooth and shining, and the MOA: Prey is usually collected , pleura are finely sculptured. Te- before the burrow is constructed. gulae are shining black. Abdomen is large, ovate, convex, and very robust with a smooth and polished surface. Petiole is short and ro- bust. Wings vary from subhyaline to fuliginous and often are tinged with a yellow fuscous. Veins are black, legs are black, very robust, shining, and with tibiae sparsely and tarsi densely spinose. Male resembles the female but is much smaller, the face more narrow, and it is covered with a fine sil- very cinereous pile. {7A, PP- MAP 52.—Recorded distribution of A63-A6A) Sphex pubidorsus (Costa). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 105

The wasp captures a grasshopper, though most of the host species stings it, and often stores it in a are seldom abundant and of little clump of grass. It then constructs economic importance. a burrow in the nearby sand by Sphex rufiventris ruñventris digging and shoveling out the Cresson grains, similar to the manner of the other species of this genus. It 1872: 211 then returns to its prey, straddles Synonymy.—Palmodes rufiven- it, and grasping the antennae in tris (Cresson) (225, p. 97i). its mandibles drags it back to the Recorded distribution.—South- burrow. It then enters and pulls western Canada and western the grasshopper inside, deposits United States (225, p. 97^). an egg at the base of the hindleg, (Map 53.) departs, fills in the burrow, and Host species.—Stipator, prob- covers the entrance with sticks ably stevensonii (369, p. 227). and other debris. Grasshoppers Incidence of parasitism.—Not paralyzed but left generally die located. after about 5 days. AF : Not given. Life history and habits.—SO: HAB: Roadways, near rivers, Eggs, larvae, and pupae unknown. sandy areas, and grassy areas. Adults are found in August. LOD : Attacked by the hyperparasite Unknown. OWS, G/Y, SA, N/H, Senotainia rubiventris (Ma- and MOA: Not given. AF: Seen quart) . Í97,pp.l83-18i;252,p. on sumac flowers. HAB: Not 23; 316, p. 500) given. (75, p. 211) Description.—Eggs, larvae, and Description.—Eggs, larvae, and pupae unknown. Adults are about pupae unknown. Adult females 15 mm. long, black, and covered are 23 mm. long and a deep vel- with a golden pile. Face, clypeus, pronotum margins above, spots under the wings, all are gray sil- ver. The abdomen is dark red rust. The first segment of the petiole has a black oblong spot on the top. The second segment has a large black oval spot. Wings are a gray hyaline with the veins black pitch. The tegulae are black, and the tibia is a tawny fomentóse behind. (71, p. 69) Discussion.—It is widespread MAP 53.—Recorded distribution of and has a good host range, al- Sphex rufiventris rufiventris Cresson. 106 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE vety black. The head, scutellum, (Cyrtacanthacridinae). (97, p. and abdomen are shiny ; the meso- 18i) thorax and scutellum are closely Incidence of parasitism.—None covered with minute punctures. located. Petiole is long, and the abdomen Life history and habits.—SO: oblong, oval, and entirely yellow Eggs, larvae, and pupae unknown. ferrugineous. Legs are black with Adults not given. LOD : Unknown. the tibiae brown within. Wings OWS, G/Y, and SA: Not given. are fuscous. (75, p. 211) N/H : One. MOA : S. striatus para- Discussion.—Only one host rec- lyzes the prey before constructing ord was noted. the burrow which takes approxi- mately an hour. Although the Sphex striatus (Smith) grasshopper, a Xylius species, is (Priononyx) 1856: 266 considerably larger than the wasp, Synonymy.—Priononyx stri- the latter holds the base of the atus Smith, Sphex (Priononyx) grasshopper's antennae in its laerma Cameron (225, p. 973). mandibles and the body with its Recorded distribution.—Ne- front claws while walking on its vada, Arizona, New Mexico, and middle legs and hindlegs and vi- Texas. Also south to Argentina. brating its wings. The wasp en- (225, p. 973). (Map 54.) ters first, pulling its prey inside Host species.—None reported by its antennae. An egg is de- in the United States. However, in posited on the grasshopper at the Mexico it attacks Xylius spp., base of the hindcoxae, after probably centralis Rehn (Oedi- which the wasp exits and closes podinae), and in Argentina Schis- the entrance. AF : Flowers of the tocerca cancellata (Serville) black mangrove (Avicennia nitida Jacq. ) in Texas. HAB : Not given. (97, p. 18i) Description.—Eggs, larvae, and pupae unknown. Adult females are 21 to 23 mm. long and are basically black. The face is cov- ered with a silver pile and there is a small notch in the middle of the anterior margin of the clypeus. Mesothorax and metathorax are finely strigose and are covered thinly with short cinereous pubes- cence. Abdomen is ferrugineous and the petiole black. Tibiae and MAP 54.—Recorded distribution of tarsi are strongly spinose ; the an- Sphex striatus (Smith). terior tarsi contain long cilia on ARTHROPOD AND NEMATODE PARASITES IN AMERICA 107 the outside. Wings are black with a violet iridescence; the apical margins are lighter. The male dif- fers from the female only in be- ing smaller. {302, p. 256) Discussion.—It appears to pre- fer the desert and requires warmer, drier areas than are found in most of Canada and the United States.

Sphex thomae Fabricius 1775: 346 MAP 55.—Recorded distribution of Synonymy.—Priononyx thomae Sphex thomae Fabricius. (Fabricius), Pepsis crusts Fabri- cius (225, p. 973). S. thomae may attempt to dig Recorded distribution.—United in several locations before the de- States west of the Rocky Moun- sired site is found, usually in an tains and southeastern United area where sand or cinders are States {225, p. 973). (Map 55.) present. Chunks of the soil and Host species.—Amphitornus cinders are carried with the species, (Bur- mouth parts as the wasp backs meister), Aulocara species, Dis- out of its burrow with its load sosteira Carolina (Linnaeus), En- and drops it near the orifice of coptolophus subgracilis texensis the tunnel. This continues until a Bruner, Orphulella pelidna pe- small pile of debris is accumulated lidna (Burmeister), Paraidemona near the entrance, at which time species, and Trimerotropis vicu- the wasp rapidly kicks and lata (Rau) (97, pp. 181-183; 129; brushes the material to a neat 257, pp. 62, 175-193; 369, p. 277). little pile some distance away. Incidence of parasitism.—None After the tunnel has been con- located. structed downward to a distance Life history and habits.—SO: of about its own body length, the Not given. LOD: Eggs require 2 wasp begins constructing the cell to 3 days. Larvae develop in sev- in a horizontal direction. Each eral days. Pupal stage lasts for 1 tunnel leads to only one cell, which month. Adults not given. OWS: lies 1 to 3 inches below the soil Not given. G/Y : Possibly several. surface. Upon completion of the SA: Not given. N/H: One. MOA: construction, the wasp departs Stings and paralyzes prey for and attempts to relocate the host, food for its young. AF : Not given. a feat that is sometimes accom- HAB: Chambers constructed in plished with difficulty. soil of sand, cinders, or both. When found, the wasp mounts 108 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE the grasshopper so that it is fac- Petiole is elongated and the ma- ing in the same direction as its culae are large and black. Wings prey, and grasping the antennae are hyaline. (75, p. 213; 102, p. with its mandibles attempts to 3A6) drag it toward the burrow with Discussion.—The split distri- the aid of its wings. The grass- bution may indicate incomplete hopper is drawn into the burrow collection records but more likely and placed with the head toward a retreat of the species to favor- the far wall, after which a small able areas. egg is deposited near the hind- coxae. The wasp then departs and Tachysphex fusus Fox fills the burrow entrance with 1893: 519 sand and cinders but does not at- Synonymy.—None (225, p. tempt to pack the material down 951). with the head as does S. atratus. Recorded distribution.—United For the next 2 or 3 days the egg States except Alaska and Florida. increases in size and changes Also found in Hawaii. (225, p. from brown to green. The larva 951) (Map 56.) consumes the food supply in sev- Host species.—Oxya chinensis eral days and forms a light brown (in Hawaii) and Melanoplus spp. pupa. (97, pp. 181-183; 130; in the United States and Hawaii 257, pp. 92, 175-193; 369, p. 227) (123, p. 4-90 ; 225, p. 951 ; 368, pp. Description.—Eggs unknown. 201, 211; 370, p. 17). Larvae (full grown) are approxi- Incidence of parasitism.—None mately 18 mm. long and appear located. similar to S. atratus larvae in Life history and habits.—SO; shape. The apical margin of the Eggs, larvae, and pupae unknown. labrum has several circular pores Adults not given. LOD : Unknown. on each side of the median line, whereas the epipharyngeal sen- sory area consists of about seven pores on each side. Mandibles, maxillae, and labrum are very similar to those of S. atratus as are the spinulae. Antennal orbits are slightly eccentric. {101, pp. 137, 1A2-H3) Pupae unknown. Adults of S. thomae appear simi- lar to S. atratus adults although slightly smaller. Length is about 50 mm. The abdomen is red, and the head and thorax are black MAP 56.—-Recorded distribution of and covered with a silvery pile. Tachysphex fusus Fox. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 109

OWS, G/Y, SA, and N/H: Not be very finely punctured ven- given. MOA: Females provision trally. In color and appearance the their burrovi^s v^ith grasshopper male is very similar to the female. hosts and can sometimes be found {115, pp. 519-520; 273, pp. 222- vs^aiting in ambush in ant lion pits. 223) AF: Not given. HAB: Sandy Discussion.—Although widely areas vs^ith sparse vegetation. located, the host range appears {228, p. 615; 257, pp. 1^9-150) limited. Description.—Eggs, larvae, and Tachysphex plenoculiformis pupae unknovs^n. Adult females Williams are approximately 8 mm. long and are characterized by having the 1913: 167-168 last abdominal segments bright Synonymy.—None (225, p. red. Clypeus is rounded anteriorly 952). with the apical margin slightly Recorded distribution.—West- uneven. Front mesopleura, pectus, ern Kansas {225, p. 952 ; 368, pp. and the vertex are strongly punc- 167, 201-202, 211). (Map 57.) tured. Dorsulum, scutellum, and Host species.—Tryxalid locusts postscutellum are shining and {368, pp. 167, 201-202, 211). have separated punctures. Both Incidence of parasitism.—Not the mandibles and scape beneath given. However, will use as many the apex are reddish. Front, face, as six immature grasshoppers per and clypeus are densely covered burrow. with a silvery pubescence, the Life history and habits.—SO: thorax is sparsely covered and the Eggs, larvae, and pupae unknown. abdomen has a rather dense silver Adults not given. LOD : Unknown. pile. The last two abdominal seg- OWS and G/Y: Not given. SA: ments are bright red, and the Nymphs. N/H : Not given. MOA : pygidium of the male is sparsely punctured. Legs are well spined and tarsi are reddish. Wings are subhyaline and iridescent with the veins black. Males are 7 to 8 mm. long. Front and vertex are coarse and contain confluent punc- tures. Antennae are rather long; the flagella are slightly thickened medially with the first joints about one-third shorter than the second. Upper surface of the metathorax is transversely rugose granulate and the sides are rather MAP 57.—Recorded distribution of strongly striated. Abdomen may Tachysphex plenoculiformis Williams. lio TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Stocks burrow with immature lo- Incidence of parasitism.—None custs. AF and HAB: Not given. located. (228, pp. 615-619; 368, pp. 167, Life history and habits.—SO : 201-202, 211) Eggs, larvae, and pupae unknown. Description.—Eggs, larvae, and Adults not given. LOD : Unknown. pupae unknown. Adult females OWS and G/Y: Not given. SA: are black, slender, and 7 mm. in Mature and immature grasshop- length. The frons has shallow pers. N/H, MOA, and AF: Not punctures as does the polished given. HAB : Desert. (368, pp. vertex. The ocellar depression is 202-203, 211) deep. Antennae are covered with Description.—Eggs, larvae, and a silvery pile. The fourth anten- pupae unknown. nal joint is longer than the third, Adult females are 10 mm. long which is about twice as long as and are noted for their silvery the second. The scutellum is punc- ornamentation and color. Clypeus tured and the sides are strongly is shining with both large and striate. Abdomen is red with the small punctures on the surface, pygidial area well defined and which are separated by irregular covered with punctures. There is spaces. Small, confluent, imper- a sparse white pile covering the fect punctures on the front give body. Legs are slender. Venation the integument a rugose appear- of the wings is weak, and the mar- ance. The first joint of the an- ginal cell is not obliquely trun- tenna is a little more than two- cate. (368, pp. 167-168) thirds the length of the second. The posterior ocellus is a heart- Tachysphex propinquus Viereck shaped impressed space, which is shining and has a medially longi- 1904: 87-88 tudinal depression. Sculpturing of Synonymy.—None (225, p. 952). Recorded distribution.—West- ern United States east to central Montana, central Kansas, and central Texas (225, p. 952). (Map 58.) Host species.—Ageneotettix spp., Cordillacris spp., Drotmena spp., Mesotobretma spp., and Opeia spp. It is also reported prey- ing on mature A. deorum (Scud- der), C. crenulata (Bruner), and

Trachyrhachys kiowa (Thomas). MAP 58.—Recorded distribution of (368, pp. 202-208, 211) Tachysphex propinquus Viereck. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 111 the thorax is similar to that of the frons. The scutellum is convex and slightly depressed medially, whereas the postscutellum is dull and the metathorax finely and uniformly leathery. Abdomen is dullish, long, and very finely sculptured. The pygidial area is shining, slightly convex, and with a few punctures on the sides. Color is a dark ferrugineous with the head black except the clypeus. A silvery pubescence covers the MAP 59.—Recorded distribution of whole insect and almost obscures Tachysphex quebecerisis (Provancher). the reddish and black ground col- ors. Host species.— (368, Legs are black and of the same p. 211). color as the abdomen. Pulvilli and Incidence of parasitism.—Not the large spur of the four poste- given. rior tibiae are black; the anterior Life history and habits.—SO: tarsi contain approximately 15 Eggs, larvae, and pupae unknown. bristles to the comb. The middle Adults not given. LOD: Eggs, tarsi support eight spines or more larvae, and pupae unknown. on the first joint between the base Adults not given. OWS and G/Y: and the apex, whereas the poste- Not given. SA: Immature grass- rior tarsi contain only one or two hoppers. N/H: Unknown. MOA: spines on the first joint between Not given. AF : Not located. HAB : the base and the apex. Wings are Not given. (368, p. 211 ) hyaline and the veins dark brown Description.—Eggs, larvae, and to h\ack.(343, pp. 87-88) pupae unknown. Discussion.—It has a good host Adult females are 10 to 11 mm. range. long and are generally black. Mandibles in the middle, the un- Tachysphex guebecensis der part of the scape, and the (Provancher) apical area of the tarsi are red- dish. There is a silvery pile on the (Larra) 1882: 150 abdomen; the first two segments Synonymy.—Larra quebecensis are red. The front part of the head Provancher (225, p. 952). and the thorax have a silvery Recorded distribution.—Que- pubescence. First joint of the an- bec, Ontario, Maine, and Mas- tennal flagellum is slightly shorter sachusetts (225, p. 952). (Map than the second joint. The dor- 59.) sulum is finely and closely pune- 112 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE tured and the scutellum is dis- tinctly impressed. The above area of the metathorax is coarsely wrinkled and the metapleurae are striated. The pygidial area is strongly punctured and is nearly 214 times longer than it is broad at the base. Wings are dark sub- hyaline and the nervures are blackish. Males are approximately 9 mm. long and are colored similarly to the females but with a more pro- MAP 60.—Recorded distribution of fuse pubescence on the head and Tachysphex seviirufus (Cresson). > thorax. The antennae are rather p long and slender; the first joint Life history and habits.—SO: of the flagellum is about one-third Eggs, larvae, and pupae unknown. ' shorter than that of the second. Adults not given. LOD : Unknown. í The metathorax is more strongly OWS, G/Y, SA, N/H, MOA, and ^ sculptured than that of the fe- AF: Not given. HAB: Mountain male, but the striations of the shrubbery. (228, p. 615; 368, p. • metapleurae are not so strong. 211) {115, pp. 527-528; 251, p. 150) Description.—Eggs, larvae, and Discussion.—It may be re- pupae unknown. Adult females stricted in both geographical and are 7 to 9 mm. long. This species . host ranges. is probably most closely related to Tachysphex tarsatus (Say) and • Tachysphex semirufus (Cresson) can easily be distinguished from ^ (Larrada) 1865: 464 it by size, punctures, and sculp- Synonymy.—Larrada semirufa turing on the metathorax. Ante- Cresson, Tachysphex punctulatus rior margin of the clypeus is , Smith, T. puncticeps Smith, T. slightly rounded outward and giffardi Rohwer, L. semirufa Pat- armed on each extreme side by ton (225, p. 952). two distinct teeth. On the vertex Recorded distribution.—West- are fine punctures. The first joint ■ ern United States (225, p. 952). of the flagellum is scarcely one- (Map 60.) fourth shorter than the second. Host species.—Melanoplus Sides of the metathorax are spretus (Walsh) and katydids striated and granulate. Abdomen (228, p. 615; 262, pp. 317-318; is impunctate above ; the last ven- , 368, p. 211). tral segment is strongly punc- Incidence of parasitism.—None tured. Pygidial area is well de- located. fined and sparsely punctured. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 113

General color is black except the turns to the burrow with the prey, scape, beneath at the apex, the often flying, opens the burrow, middle of the mandibles, and part and enters. She emerges head of the tarsi, which are an obscure first, pulls the grasshopper inside, reddish. The abdomen is bright deposits an egg, and departs, seal- red, whereas the face, front, cly- ing the entrance again. Each cell peus, and thorax are sparsely cov- may contain a dozen grasshoppers ered with a silvery pubescence, and there may be four cells to a which is absent on the abdomen. nest. The grasshopper nearest the Legs are strongly spined. The entrance receives the egg near the wings are slightly dark brown and coxa of the jumping leg. iridescent. Veins are black. {7A, AF: Males eat honey, nectar, p. A6i) and pollen. HAB: Males prefer Discussion.—Only one grass- open sandy areas in which they hopper host was located. dig short burrows for resting dur- ing the night and hot, inclement Tachysphex similis Rohwer weather. These burrows are at 1910: 51 an angle of about 30° to 40°, about Synonymy.—Tachysphex simi- 2.3 cm. long, and 4 mm. in diam- lans Rohwer, Solierella similans eter. When closed inside the male (Rohwer) (225, p. 952). faces the entrance. Later he Recorded distribution.—From leaves the burrow and may not re- Utah and Massachusetts south turn, but he may construct a new through Texas and Florida (225, burrow when needed. (172; 173; p. 952). (Map 61.) 371) Host species.—Acridid grass- Description.—Eggs, larvae, and hoppers (123, p. U89; 371). pupae unknown. Adults are 5 to Incidence of parasitism.—None 6 mm. long and are generally located. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults are in the field from April to May in Texas and to August in New Jersey. LOD: Unknown. OWS and G/Y: Not given. SA: Nymphs. N/H: One per host or up to one per 12 hosts. MOA: The female excavates a slanting burrow in the sand that terminates in one to four cells. She closes the entrance, departs, and locates her prey, which she MAP 61.—Recorded distribution of paralyzes with a sting. She re- Tachysphex similis Rohwer. 114 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE black. The occiput of the female is shining and surrounded with punctures. The third antennal joint is approximately one-third shorter than that of the fourth. The dorsulum also contains close, distinct punctures as does the scutellum, which is also im- ^ pressed. Metanotum is closely and .r:^' \ . Í ; , » I coarsely granular and separated ■ from the transverse striated pos- V.I «i terior face by a furrow that is broken in the middle. Legs are MAP 62.—Recorded distribution of Tachysphex tarsatus (Say). covered with weak spines. The apical abdominal segment is red or black and covered with a sil- Metator nevadensis (Bruner), very pubescence. Wings are Camnula pellucida (Scudder), faintly dusky hyaline and irides- Oedaleonotus enigma (Scudder), cent with dark-brown venation. Dissosteira spurcata (Saussure), The male is very similar to the Melanoplus devastator Scudder, female except the venation of the and many other species of the wings is pale brown and the wings slant-face, , and spur- are clear. (28, pp. Í02-Í03; 275, throated grasshoppers. Several pp. 51-52) species of Mantidae are also in- Discussion.—The works of cluded. (123, p. ^90; 220, p. 10; Kurczewski (172, 173) cover its 225, p. 952; 265, p. 270; 368, pp. life history and habits as well as 203-20Jt, 211) possible hyperparasites. Incidence of parasitism.—None located. Tachysphex tarsatus (Say) Life history and habits.—SO: (Larra) 1823: 78 Eggs, larvae, and pupae unknown. Synonymy.—Larra tarsata Adults not given. LOD : Unknown. Say, Tachysphex hitteri Rohwer, OWS and G/Y: Not given. SA: T. zimmeri Michel (225, p. 952). Prefers third instars. N/H: Not Recorded distribution.—South- given. ern Canada and the United States MOA : The life history and hab- (225, p. 952). (Map 62.) its are similar to those of the Host species.—Xanthippus neg- other Tachysphex species that are lectus (Thomas), Trimerotropis parasitic on grasshoppers. The montícola (?), Platuibothrus burrow is constructed, the female brunnea (?) (these last two spe- wasp leaves in search of prospec- cies of questionable identifica- tive victims, one is located, a tion), (Say), struggle ensues during which the ARTHROPOD AND NEMATODE PARASITES IN AMERICA 115 grasshopper is stung one to sev- striated and the dorsal part also eral times, and the wasp returns contains very close punctures. to the burrow with the prey. It The metathorax above and on the is placed inside the nest and on it sides is finely coriaceous. Legs are is placed an egg. The nest is feebly spineous. eventually closed and the female The second segment of the fe- leaves to repeat the cycle. male antennae is about three T. tarsatus would probably be times as long as the first. The a much more effective parasite if dorsulum is finely punctured and it were not for two metopiid flies, depressed at midpart anteriorly. Sphenometopa (Ettaraba) ter- Scutellum is not impressed. Dis- gata Coquülett and Taxigramma tinct striations can be seen on the heteroneura Meigen. When fe- metapleura and on the face. Ab- males of either species observe T, domen is also impunctured and tarsatus dragging its prey, they the ventral areas of the last two lay in wait until the victim has segments have sparse punctures. been placed in the burrow and the Midpart of the mandibles and the wasp has departed for a few sec- apical joints of the tarsi are ob- onds of rest. These flies then dart scurely reddish. The abdomen into the burrow and deposit living varies from a bright red to nearly maggots on the grasshopper. black. Wings are subhyaline and These maggots then consume the iridescent and the veins black. grasshopper and the wasp egg. {192, p. 116; 28i, pp. 78-79) AF and HAB: Not given. Discussion.—This parasite is (220, p. 10; 228, pp. 615-619; extremely active and definitely 265, p. 270; 287, p. 78; 368, pp. reduces grasshopper populations 203-20i,211) (265, p. 270; 368, pp. 203-20Í, Description.—Eggs, larvae, and 211). pupae unknown. Adult females are 10 to 11 mm. Tachysphex tenuipunctus Fox long and males are 7 to 8 mm. 1893: 525 General color i^ a bright f errugin- Synonymy.—Tachysphex gran- eous. The body and legs have a ulosus Mickel (225, p. 953). more or less obvious silver ciner- Recorded distribution.—South- eous pile. Front of the male is western Canada and western coarsely granulated, whereas that United States (225, p. 953). of the female is fine. The vertex (Map 63.) and clypeus of the male contain Host species.—Oedaleonotus very close punctures. Antennae enigma (Scudder), Aulocara el- are rather stout and long; the liotti Thomas, Melanoplus spp., second segment is about twice and an unknown Hymenoptera the length of the third. Posterior species (123, p. Í90;228, pp. 615- part of the face is distinctly 619; 389). 116 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

host's prothorax and mesothorax in preparation for oviposition. During this time it often feeds ^ on the body fluids and the regur- gitated gastric juices of the host. It then straddles the host and ; drags it back to the burrow. The wasp pulls the grasshopper into ' its burrow and remains therein ' for several minutes while it de- '_ posits an egg on the ventral sur- face between the prothoracic legs. ; MAP 63.—Recorded distribution of The wasp then fills the entrance Tachysphex tenuipunctus Fox. with small grains of sand, packing it periodically. After hatching, > Incidence of parasitism.—Not the larva usually inserts only its given. head within the body cavity of the Life history and habits.—SO: paralyzed grasshopper. Larval v Eggs are deposited in the summer. feeding usually causes the grass- Larvae occur in the summer and hopper's death by the second day fall. Pupae can be found from after hatching, but the larva con- -î fall to spring. Adults are in the tinues to feed for an additional field from May to June. LOD: 8 days. Newton (228) reported Eggs require 2 to 3 days. Larvae possibly four larval instars. When 8 to 10 days. Pupae 2 weeks ; pre- mature, the larva constructs a pupae 1 week. Adults not given. hard cocoon from sand grains and OWS: Cocoons. G/Y: One. SA: cement in which it remains until " Nymphs. N/H : Not given. MOA : late April or early May of the fol- . Stings and paralyzes prey on lowing year. Adult emergence us- which young will mature. AF: ually coincides with the immature Hemolymph of the host. HAB : stages of 0. enigma. (228, pp. Sunny forested areas and transi- 615-619) tional zones. Description.—Eggs are approx- A gravid wasp chooses an area imately 2 mm. long, pale gray, of soil and digs a small diagnoal and sausage shaped, tapering burrow 114 to 2 inches deep, after slightly to the distad pole. The which it walks around the en- egg is placed transversely along trance several times, apparently the ventral prothorax and meso- taking its bearings. It then flies thorax of the grasshopper nymph, off to hunt for a suitable host. with the anterior part cemented When located, it attacks, stings, adjacent to the membrane, which paralyzes, and finally softens by surrounds the procoxa. {228, chewing the ventral surface of the p. 216). Larvae (full grown) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 117 are 14 mm. long, lack legs, and Tachysphex terminatus (Smith) are chocolate brown with a bluish (Larrada) 1856: 291 cast. They are shiny and appear Synonymy.—Larrada termi- plump. When the larvae are feed- nata Smith, Larra minor Pro- ing, the body is usually curved vancher, Larra terminata Patton about the ventral neck of the host. (225, p. 953). (228, p. 216) Cocoons are from Recorded distribution.—From 10 to 15 mm. long (average 12 British Columbia east through mm. ) by 6 mm. wide and are oval. Ontario, south through Georgia, The cocoon shell is constructed of west through Arizona, and north sand, which is cemented together. into British Columbia (225, p. Within the newly constructed co- 953). (Map 64.) coon the mature larva is organish Host species.—Syrbula admir- yellow, which fades to a light yel- abilis Uhl, Melanoplus nymphs, low with age. {228, p. 217) Chortophaga viridifasciata (De- Prepupae are light yellow, which Geer), and several species of is maintained through the early slant-face pigmy locusts and other pupal stage. (228, p. 217) Acrididae (123, p. Jt90; 225, p. Adult females are 9 to 11 mm. 953; 25A, pp. 157-260; 368, pp. long and generally black. Front, 201, 211). face, and clypeus have a sparse Incidence of parasitism.—None silvery pubescence, whereas the located. thorax is bare. The space between Life history and habits.—SO: the eyes at the top is equal to Eggs, larvae, and pupae unknown. about the length of the second and Adults not given. LOD : Unknown. third antennal joints combined. OWS, G/Y, and SA: Not given. Mesonotum in front of the scutel- N/H: One parasite per one to lum is very finely and closely nine hosts. punctured, whereas the scutellum is slightly impressed. The meta- pleura are leatherlike, with the posterior section slightly striated. First three segments of the abdo- men (sometimes only one-half of the third) are bright red. The py- gidial area contains sparse but strong punctures and is about 21/2 times longer than it is wide at the base. Wings are subhyaline and iridescent. (115, p. 525) Discussion.—The burrow of this species is constructed prior MAP 64.—Recorded distribution of to obtaining the host. Tachysphex terminatus (Smith). 118 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

MOA : The life history and hab- absolutely, he did observe that one its of this species are very similar young larva on a brown grasshop- to those of the other Tachysphex per was itself brown, whereas species mentioned here in that the one on green food was green. Most female excavates a burrow with of the victims deposited within one or more cells, in which var- the cell are killed before the egg ious insect prey are stored to sup- is deposited ; some have the head ply food for the developing egg. entirely removed. Those remain- The burrow is constructed by dog- ing alive are so weak and feeble like digging. The ground is at- from the sting that they are un- tacked with the forelegs and the able to respond except to extreme soil and particles are flung under stimulation and only barely so. the body and behind it. As the bur- The egg is deposited in the area row increases in depth, the wasp of the legs between the second will back out of the hole and and third pair on the ventral side, sweep out the accumulated debris and feeding commences at that from in front of the door. Sandy point. (172, pp. 317-322; 173, areas are usually preferred, al- pp. Jf36-JÍ53\ 25Jf, pp. 157-260; though other areas may be util- 256, p. 10; 368, pp. 201, 211) ized. Burrows within which the Description.—Eggs unknown. female rests are dug at night. In Larvae in diapause are 9 mm. the morning she seeks a host, long. The integument is smooth stings it, and returns with it to dorsally, with minute spines cov- the burrow opening. The wasp ering parts of the venter. The ori- crawls into the burrow, pulling fice of the spiracles is large and the host in behind her. From five the atrium lined with weak ridges. to nine insects are usually used to The head is virtually unpig- provision a cell, although one mented except the anterior ten- large insect may be utilized. Once torial arms, the hypostomal thick- the prey has been stashed, an egg enings, mandibles, and palpi are is deposited on it and the wasp light brown. Antennal orbitals exits to seal the mouth of the bur- are nearly circular, and the head row. Although the female wasp contains very prominent setae. will attack any intruder during There are single lateral setae at this operation, immediately upon the base of the mandibles and five completion she deserts her work. teeth at the apex. Palpi are stout AF: Frequents flowers of Ceano- and slender. {96, pp. 118-119). thus. HAB: Not given. Pupae unknown. Rau (25^) suggested a possible Adult males are 6 mm. long, relationship between the color of adult females 9 to 10 mm. Gen- the developing wasp larva and erally the female is black with that of its prey. Although he the midpart of the mandibles and stated that this cannot be declared the scape beneath at the apex ARr;ROPOD AND NEMATODE PARASITES IN AMERICA 119 reddish. Tarsi are reddish testa- ceous apically, and the last two segments of the abdomen are a bright red. Front is highly gran- ulated and the vertex is strongly punctured. The first joint of the antenna is nearly one-third shorter than the second. The up- per surface of the metathorax is granulate and distinctly striated. Legs are not strongly spined. Py- gidial area is sparsely punctured. Wings are subhyaline and irides- MAP 65.—Recorded distribution of cent with the veins testaceous. Tachysphex texanus (Cresson). The male of this species is similar to that of T. fusus. The legs are granuled and that of the female is less strongly spinulose and the fine. In the male the antenna is front is somewhat more closely short and stout and the first seg- and finely punctured. The clypeus ment is scarcely one-half as long is dentiform. {299, p. 291) as the second. The dorsulum is punctured, whereas the meta- Tachysphex texanus (Cresson) thorax above is coarsely granular (barrada) 1872: 214 and finely reticulated with the Synonymy.—Larrada texana sides and posterior parts dis- Cresson, Larra texana Patton tinctly striated. Legs are feebly (225, p. 953). spinulose. Color of both male and Recorded distribution.—South- female is similar, with the ab- ern California, Utah, and Texas domen of both covered with a sil- (225, p. 953). (Map 65.) very pile. In the female the sec- Host species.—Oedipodinae and ond antennal segment is four flies (123, p. A90; 368, p. 211). times that of the first. Scutellum Incidence of parasitism.—None and dorsulum are covered with located. punctures; the former is slightly Life history and habits.—SO: impressed. The abdomen is also Eggs, larvae, and pupae unknown. punctured and bright red. The Adults not given. LOD : Unknown. midpart of the mandibles, as well OWS, G/Y, SA, N/H, and MOA : as the spot on the scape at the Not given. AF : Unknown. HAB : apex beneath, and the tarsi are an Desert. (368, p. 211) obscure red. Both the head in Description.—Eggs, larvae, and front and the thorax are sparsely pupae unknown. Adult males are covered with a silvery pubescence, 7 mm. long, females 9 to 10 mm. as is the abdomen, which is cov- The front of the male is coarsely ered with a silvery pile. Wings 120 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE are subhyaline with the veins antennae are black; the fourth black. (75, p. 2H) one has a lateral sericeous lunule. ~ When viewed from above, the ^ Tachytes adbominalis (Say) anal segments have a red reflec- (Larra) 1823: 77 tion. Feet are black. (75, p. 217; Synonymy.—Larra abdomi- in, pp. 2U, 2k8-2k9; 192, pp. i nalis Say (225, p. 9^7). 165-166; 284, p. 77) Recorded distribution.—Colo- rado, Texas, and Arkansas (225, Tachytes elongatus elongatus p. 947). (Map 66.) Cresson ''< Host species.—Acridinae and 1872: 215 Tettigoniidae (123, p. 490; 368, Synonymy.—None (225, p. ^ p. 211). 945). Incidence of parasitism.—Not Recorded distribution.—Brit- given. ish Columbia southeast into Mary- Life history and habits.—SO: land, south into Florida, west into Eggs, larvae, and pupae unknown. California, and north into British Adults not given. LOD : Unknown. Columbia (225, p. 945). (Map OWS and G/Y: Not given. SA: 67.) Nymphs. N/H, MOA, AF, and Host species.—Melanoplus spp., HAB : Not given. (200, p. 2^3) Ageneotettix spp., and Héspero- í Description.—Eggs, larvae, and tettix spp. Melanoplus spp. is pupae unknown. Adults are 14 to preferred. M. lakinus (Scudder). 18 mm. long and black in general. (200, pp. 241-245; 368, pp. 194- The thorax is cinereous and the 197) first, second, and third segments Incidence of parasitism.—Not of the abdomen are red. The front given. is silvery as are the hairs on the Life history and habits.—SO: feet. Terminal segments of the Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS, G/Y, SA, and N/H: Not given. MOA: Paralyzes prey on which young will mature. AF and < HAB : Not given. Burrows of T. elongatus elonga- tus are approximately 12 inches deep and the entrance is sur- rounded by a mound of soil. The burrow leads to numerous cells, which are each stocked with one or several specimens of short- MAP 66.—Recorded distribution of horned grasshoppers and on which Tachytes ahdomirMlis (Say). an egg is deposited. Williams ARTHROPOD AND NEMATODE PARASITES IN AMERICA 121

same grasshopper. He also re- : ^m^y ported one unidentified dipterous maggot in the cells where wasp larvae had developed. (98, pp. 286-287; 255, p. 260; 368, pp. 194-197) Description.—Eggs, larvae, and pupae unknown. Adult males are 14 to 18 mm. long and black. Head and thorax are covered with a short silkylike pubescence, which is silverish under certain lights. MAP 67.—Recorded distribution of The face and the clypeus have a Tachytes elongatus elongatus Cresson. dense pale-golden pubescence. Mandibles are black at the tips, (368) described two nests, one yellowish at the base, and red- containing 20 cells with a total of dish in between. Scape of the an- 56 prey and the second with 16 tennae is short and has long pale cells and 38 prey. Burrows are hairs. The posterior margin of generally constructed in sandy the prothorax is silvery. Meso- areas, although Williams did re- thorax and scutellum are closely port the soil around one was hard. and feebly punctured and are Because of the confusion as to clothed with short, erect, silky- the status of the two species it is type pubescence. The metathorax uncertain which species Evans lacks distinct punctures but is (98) was describing when he covered with a whitish pubes- wrote of Tachytes distinctus cence. Abdomen is a deep velvetry Smith. He reported that the bur- black with the basal segments rows contained several adults or bearing long white pubescence. last-instar njrmphs of Melanoplus The apex of segments 1 to 4 has lakinus. The burrows descended a band of dense, silvery silky pile, at an 80° angle with the surface whereas segments 5 and 6 are en- to a depth of 42 cm., which is tirely black. The apical segment somewhat longer than the 10 to is bright silvery. Legs are brown- 12 inches described by Williams ish yellow with a silvery silky pile. and Rau (255). At a depth of 35 Coxae, trochanters, and femora cm. Evans located three cells, except the tips are black. Wings each of which was at the end of a are yellow hyaline and paler to- long side burrow; these cells ward the apex. (75, pp.215-216) measured 12 by 20 mm. He found Discussion.—There is some some cells containing one larva confusion as to the status of this per grasshopper, whereas in oth- species and its relationship to T. ers several larvae utilized the distinctus. 122 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Tachytes exornatus Fox strongly rounded out medially and 1893: 501 supports one large blunt lateral Synonymy.—None (225, p. tooth. The dorsulum and scutel- H5). lum are indistinctly punctured Recorded distribution.—South- and the thorax is clothed with a ern California, Arizona, New grayish-golden pubescence. There Mexico, and Texas (225, p. 945). is an impressed medial line on the (Map 68.) metanotum and a transverse band Host species.—Conalcaea spp. on the abdominal segments above (170, pp. 75-79; 200, p. 2^3). and beneath. Tibiae and tarsi are Incidence of parasitism.—None spinose with the longer spur of located. the hindtibiae shorter than the Life history and habits.—SO: first hind tarsal segment. Scape, Eggs, larvae, and pupae unknown. tegulae, legs (except coxae), tro- Adults not given. LOD : Unknown. chanter, forefemora, midfemora, OWS, G/Y, SA, N/H, MOA, AF, and a transverse band on the ab- and HAB: Not given. (100, p. dominal segments above and be- 332) neath are reddish. Wings are yel- Description.—Eggs, larvae, and low. (115, p. 501) pupae unknown. Adult males are Discussion.—It appears to pre- 17 mm. long and black. The third fer grasshoppers of noneconomic joints of the antennae are equal importance and is usually found in to or possibly somewhat longer dry warm places. than the fourth, and the basal joints of the flagellum are Tachytes fulviventris fulviventris rounded. The front of the clypeus Cresson is covered with a golden pubes- 1865: 466 cence, and the anterior margin is Synonymy.—Tachytes caelebs Patton (225, p. 9i6)- Recorded distribution.—West- ern United States except south- west border area (225, p. 946). (Map 69.) Host species.—Alpha crenulata (123, p. 490; 368, pp. 197, 211). Incidence of parasitism.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown.

MAP 68.—Recorded distribution of OWS, G/Y, SA, and N/H: Not Tachytes exornatus Pox. given. MOA: Unknown. AF: Not ARTHROPOD AND NEMATODE PARASITES IN AMERICA 123

Tachytes obductus Fox 1892: 250 Synonymy.—None (225, p. 9i8). Recorded distribution.—Ne- braska, Kansas, Missouri, Tennes- see, and Texas (IH, pp. 2UU, 250 ; 225, p. 948; 368, p. 198). (Map 70.) Host species.— Tetrigidae (pigmy locusts) (122, p. i90; 365, p. 198). MAP 69.—Recorded distribution of Incidence of parasitism.—None Tachytes fulviventris fulviventris located. Cresson. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. given. HAB: Dry plains. (100, OWS and G/Y: Not given. SA: p. 332; 200, p. 2^3; 368, pp. 197, Nymphs. N/H : Not given. MOA : 211) The pigmy locusts are easily sub- Description.—Eggs, larvae, and dued and transported in night. pupae unknown. Adult females AF: Not given. HAB: Muddy are about 16 mm. long and basi- shores of the Solomon River in cally black. Antennae are black. Kansas. (200, p. 2k3; 251, pp. The face, cheeks, thorax, and most 150-151 ; 252, p. 27; 368, p. 198) of the legs are densely covered Description.—Eggs, larvae, and vfith. a golden silky pubescence, pupae unknown. Adult females which reflects in certain lights. are 11 mm. long. The head, tho- Mandibles are silvery at the base. Abdomen is short, broad, oval, and dark brown. Apical margins of the five basal segments contain a golden silky band, which is more obvious in certain lights. The three apical segments are black- ish; the terminal one is reddish brown as seen from above. Wings are hyaline with honey-yellow veins. Legs are black, tibiae and tarsi spinose, brownish, and densely covered with a golden silky pile. {25, p. 298 ; 75, p. 2^7 ; MAP 70.—Recorded distribution of llJf, pp. 237, 2U3; 302) Tachytes obductus Fox. 124 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE rax, and first abdominal segments are covered with a sparse, pale pubescence. Trunk and especially the ocellar region are impressed. The third antennal segment is about equal in length to either the fourth or fifth segment. The ver- tex and the dorsum along vs^ith the metanotum are very finely and closely punctured ; the last is depressed on each side with a faint, though distinct, medial im- pressed line. The scutellum is not MAP 71.—Recorded distribution of impressed. Abdomen is black and Tachytes pepticus (Say). very finely and closely punctured above, whereas beneath are large scattered punctures on segments OWS and G/Y: Not given. SA: 3 to 6. On the face, clypeus, Immature stages. N/H: MOA, ' cheeks, dorsal part of the abdo- AF, and HAB: Not given. (100, men, and most of the legs and pp. 323, 331 ; 252, p. 26) thorax is a bright silvery pile. Description.—Eggs, larvae, and '■ Tegulae veins, and spines on the pupae unknown. Adult males are legs are testaceous. Mandibles are about 13 mm. long, the females ferrugineous in the middle. Wings slightly larger. The body is black are subhyaline with a dark cloud with a silvery reflection, espe- in the marginal cell. {IH, PP- cially noticeable on the head and 2U, 250) feet. The palpi are piceous yellow ' and the head and base of the > Tachytes pépticas (Say) mandibles of the male reflect gold. • (Lyrops) 1837: 371 The female is silvery on the pos- Synonymy.—Lyrops pepticus terior margin as are the sutures Say, Tachytes pennsylvanieus of the thorax and scutellum. Ter- Banks (225, p. H6). gum segments on the posterior ; Recorded distribution.—East- margins reflect silver. The anal ern United States (225, p. 9^6). segment of the female has a ■ (Map 71.) bright golden reflection and that Host species.—Melanoplus spp. of the male is silver. Tarsi of both (100, pp. 323, 331). sexes are black with a silver re- Incidence of parasitism.—None flection; the last tarsal joints are located. red. Front coxae lack spines and ■ Life history and habits.—SO : the femora are without grooves. Eggs, larvae, and pupae unknown. (16, pp. 18-19; m, pp. 236-237, Adults not given. LOD : Unknown. 2Í2-2A3; 293, p. 371) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 125

Tachytes rufofasciatus Cresson for hunting. HAB: Not given. 1872: 199 (64, pp. 22-24; 100, p. 332; 200, j Synonymy.—None (225, p. p. 243; 368, pp. 199, 211) Description.—Eggs, larvae, and Recorded distribution.—Ore- pupae unknown. Adult males are gon east into North Carolina, 11 to 12 mm. long and black in south into Florida, west into general. The face, clypeus, and California, and north into Oregon posterior margins of the protho- (225, p. H6). (Map 72). rax and mesothorax are silvery. Host species.—Melanoplus spp. The abdomen is black except the (64-, pp. 22-24; 123, p. U90; 368, apical segment, which is also pp. 199, 211). silvery. A narrow band extending Incidence of parasitism.—None from the first to the basal margin located. of the second abdominal segment Life history and habits.—SO: is reddish fulvous. The apical Eggs, larvae, and pupae unknown. margins of the remaining seg- Adults not given. LOD : Unknown. ments are a pale testaceous. Man- OWS and G/Y: Not given. SA: dibles are yellow as is the tegula. All stages. N/H: Not given. Legs are yellow with the coxae MOA: The wasp pounces on its black. Tibiae are silvery. Wings prey and stings it in the thoracic are subhyaline and a yellow-honey region. It drags the larger prey color. (75, p. 199; 114, pp. 244, or flies with the smaller pley to 247-248) its burrow, pulling it inside. The task appears arduous and is some- Tachytes sayi Banks times made more so by the dis- 1942: 421-422 - turbing presence of parasitic flies. AF: Uses Russian-thistle plant Synonymy.—None (225, p. 946). Recorded distribution.—Wash- ington east into Nebraska, south through Colorado and central California, and north into Wash- ington (225, p. 946). (Map 73.) Host species.—Melanoplus la- kinus (Scudder) (100, pp. 323, 331). Incidence of parasitism.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. MAP 72.—Recorded distribution of Adults not given. LOD : Unknown. Tachytes rufofasciatus Cresson. OWS, G/Y, SA, N/H, MOA, AF, 126 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE and HAB : Not given. (100, pp. 323, 333) Description.—Eggs, larvae, and pupae unknown. Adult males are 10 to 12 mm. long and similar to T. pepticus, except that they can be separated by the presence of white pubescence covering the base of the mandibles and the presence of silvery hair on the face. Tenth and eleventh joints of the antenane are not widened MAP 73.—Recorded distribution of and the twelfth is not narrow. The Tachytes sayi Banks. phallus of T. sayi is more slender than in T. pepticus, the parameres slender near the tip but more are slightly curved with truncate narrow than in T. pepticus. (17, tips, and the penis is slightly pp. Í21-Í22)

Acariña At least 12 specific names of of the wing. Attachment generally mites are recorded in the litera- occurs under the wing pads, on the ture as parasitic on grasshoppers wings, or any place from which in America north of Mexico. Six they cannot be dislodged by the of these were either described by grasshopper (fig. 14). Once en- C. V. Riley or mentioned in his gorged with the grasshopper's publications. However, only one hemolymph, the larva drops to the or two species appear to be valid. ground and constructs a chamber Adults of the red grasshopper in the ground, within which it mite (Eutrombidium locustarum enters a prenymphal pupal stage. (Walsh)) begin seeking grass- It later emerges as an eight-legged hopper eggs during the first warm nymph and begins seeking grass- days of spring. When an egg pod hopper eggs. Upon completion of is found, the mite engorges itself this feeding it constructs a pre- on the contents. Within 2 or 3 imaginal pupal cell within the soil weeks after feeding, the female and remains there until emerging deposits up to 4,000 eggs in a as an adult. The life cycle may be chamber in the soil. The Isrvae, completed in 2 months in some which have three pairs of legs, areas or may require a full year. scurry to the surface and seek The adults and the eight-legged grasshoppers of nearly any instar nymphs cause more damage to the on which to attach themselves by potential grasshopper population piercing the integument or a vein than do the young nymphs, which ARTHROPOD AND NEMATODE PARASITES IN AMERICA 127

PN-2732 FIGURE 14.—Mites on grasshopper wing. are externally parasitic on the Say, T. scabrum Say, Astoma active grasshopper. The type and locustarum Walsh, T. locustarum amount of damage by the nymphs Riley, Ottonia locustarum (Riley), are still not fully understood. The T. trigonum Hermann, 0. trigona grasshopper host does not suc- Canestrini, T. holovericuum Oude- cumb to the parasitic attacks man, Allotrombidium spp., Micro- directly, but large numbers per trombidium locustarum Ewing host may cause a weakened condi- (Walsh), Italicum Oudeman (152, tion that results in death from pp. 2-28; 385). another source. It is also possible Recorded distribution.—United that mites may be a vector for States (385). (Map 74.) diseases, although there is little Host species.—Melanoplus san- evidence to support this. guinipes (Fabricius), M. bivit- tatus (Say), M. differentialis Eutrombidium locustarum Thomas, M. confusus Scudder, M. (Walsh) femurrubrum (DeGeer), Syrbula 1866: 126 admirabilis (Uhler), Brachystola Synonymy.—Astoma gryllaria magna (Girard), Phoetaliotes ne- Riley, Atoma gryllaria Lebaron, brascensis (Thomas), M. regalis Eutrombidium trigonum (Her- Dodge, Arphia xanthoptera (Bur- mann), Trombidium sericeum meister), M. gladstoni Scudder, 128 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE days at 80° F. Larvae need 5 to 8 days of engorging and total 90 days. Pupae live 20 to 25 days. Diapause takes 186 days. Adults live a year. OWS: Nymphs and adults. G/Y: One every 2 years. SA : All stages. N/H : All females require seven to eight eggs, all males 2.7, and nymphs 2.11. MOA : Adults attack grasshopper eggs; nymphs attack grasshopper nymphs and adults. AF: Grass- MAP 74.—Recorded distribution of hopper eggs. HAB : In the soil and Eutrombidium locustarum (Walsh). on host. Each produces one to six egg masses (2.76 average) with Hesperotettix viridis pratensis up to 600 eggs per mass over a Scudder, trifasciatus period of 13 days. Cool tempera- (Say), ture stimulates egg production. (Haldeman), (150; 152; 297; 385) (Scudder), Campylacantha oliva- Description.—Eggs, larvae, and cea (Scudder), Encoptolophus pupae unknown. Adults not given sordidus sordidus (Burmeister), because of difficult taxonomy. and rugosus (Scudder) Discussion.—The taxonomy of (262; 26i; 385). this species (complex ?) is still Incidence of parasitism.—As unresolved, but for the present high as 60 to 80 percent with purpose all American references some species of Melanoplus (385). are considered as applying to this Life history and habits.—SO: Eggs are deposited in the spring. name. C. V. Riley reported large Larvae occur all year. Prenymphs destruction of eggs by this spe- appear in the fall. Pupae are in cies, but the effect of the parasitic the spring. Adults can be found nymphal stages on the host has all year. LOD: Eggs require 19 not been fully determined.

HYPERPARASITOroS Hymenoptera Each of the grasshopper para- matodes, insects (generally from sites, parasitoids, and predators the orders Coleóptera, Díptera, described in this bulletin has its and Hymenoptera), and mites. own controlling agents. These in- Most of the insect predators fol- clude birds, small mammals, ne- low a similar pattern, usually not ARTHROPOD AND NEMATODE PARASITES IN AMERICA 129

preferring their catch but taking Host species.—Oscinis spp., it because it is available at the Sarcophaga d'herminieri (Des- proper time and place. voidy), S. arapacus (Walker), Like the parasitoid, the hyper- Siphona irritans (Linnaeus), parasitoid is an insect that is Agromyza platyptera var. jucunda parasitic within a parasitic insect Wulp, Tritoxa flexa (Wiede- for some of its life and free living mann), Muscina assimilis (Fal- during the remainder. Generally len), Ucila spp., Hylemia antiqua infection occurs when the primary (Meigen), and H. cilicrura (Ron- parasite is in its free-living state, dani). Thirteen A. auripes insects although there are some excep- were reared from about 80 dip- tions as noted earlier. As treated terous larvae that emerged from here the secondary insect is a an unknown number of Camnula parasite of grasshoppers. pellucida (Scudder) in Chester, The life history of the hyper- Calif. (123, p. 500; 300, p. 60; parasitoid is by necessity, closely 308; 316, p. 501 ; 327, p. 88; 389) alined to that of its host and hence Incidence of parasitism.—Not to that of its host's host. This spe- given. cialization has become so refined Life history and habits.—SO: in some species that the hyper- Eggs, larvae, and pupae unknown. parasitoid utilizes the host's host Adults not given. LOD : Unknown. as a place for its larvae to locate OWS, G/Y, SA, N/H, MOA, AF, and to infect its host. Such is the and HAB : Not given. case with the hyperparasitoid Description.—Eggs, larvae, and Perilampus hylanus Say as de- pupae unknown. Adult males are scribed later under its life history. 2.4 mm. long and a shining bril- Only a few of the better known liant black. Mandibles, base of the hyperparasitoids are included antennae, feet, and the first ab- here.

Braconidae Aphaereta auripes (Provancher) (Tñchesía) 1881: 203 Synonymy.—Trichesia auripes Provancher, Trinaría pilicornis Provancher, Aphaereta muscae Ashmead, A. calif ornica Ash- mead, A. oscinidis Ashmead (225, p. U8). Recorded distribution.—Que-

- bee and United States (225, p. MAP 75.—Recorded distribution of U8). (Map 75.) Aphaereta auripes (Provancher). 130 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE dominai segment are yellow. An- tennae are longer than the body and brown with a yellow base. Scales are white. Wings are hya- line and iridescent with yellow veins. Abdomen is shorter than the head and thorax combined and is narrower at the base. The first segment is striated and yellow and the remaining segments are a shining brilliant black. Extremi- ties are compressed; tarsi are MAP 76.—Recorded distribution of barely visible. The female is sim- Aphaereta sarcophagae Gahan. ilar to the male except the ab- domen, which is narrow at the base and enlarged toward the end, Unknown. (117, pp. 158-159; where it suddenly terminates in 389) a point. The ovipositor is approx- Description.—Eggs, larvae, and imately the same length as the pupae unknown. Adult females ^ abdomen with a supply of long are about 2 mm. long and a pol- hair, which is not too thick. ished black. Antennal scape, (250, pp. 203-20A) pedicel, palpi, mandibles, legs, and ^ the first abdominal segment have Aphaereta sarcophagae Gahan a hard brownish-yellow covering. Antennae are 23-jointed in the 1914: 158-159 female and 22-jointed in the male. Synonymy.—None (225, p. The head is about twice as wide U8). as long and the occiput is concave , Recorded distribution.—Cali- and polished. There is no notice- fornia and Kansas (225, p. 1A8). able median groove on the vertex •- (Map 76.) except possibly a trace between Host species.—Blaesoxipha kel- the ocelli. The face is smooth with lyi (Aldrich) and Sarcophaga a few indistinct scattered punc- tures. These punctures are dis- helicis Townsend (117, pp. 158- tinct on the clypeus. Mesoscutum 159; 326, p .89; 389). is smooth and polished without * Incidence of parasitism.—Not furrows. The abdomen is about as given. long as the thorax and segments Life history and habits.—SO beyond the first are smooth and and LOD: Unknown. OWS and polished. The abdomen is spatu- G/Y: Not given. SA: Pupae. late in the male. The ovipositor N/H: Eight to 12 MOA: Un- is about as long as the abdomen. known. AF: Not given. HAB: The wings are subhyaline and ARTHROPOD AND NEMATODE PARASITES IN AMERICA 131 contain a rather broad dark- does all in its power to escape the y brown stigma. {117, pp. 158- attack but eventually becomes 159) tired and somewhat passive. The V Discussion.—Since California wasp then jumps on the crevix or if and Kansas are widely separated just behind the prothoracic shield both geographically and clima- and apparently injects an egg into tologically, the distribution is the membrane. Inside the grass- j. probably much wider. hopper the egg hatches and the larva of the wasp seeks out that Chalcidae of the fly. It penetrates the larva and once inside becomes dormant Brachymeria coloradensis until after the fly has emerged i (Cresson) from its host and pupated. Then j (Chalcis) 1872: 60 the wasp completes its develop- Synonymy.—Chalcis femorata ment, killing the fly in its pupal Dalman, C. coloradensis Cresson, stage. AF: A wide variety of C. dalmanni Thomson {225, p. range flowers. HAB: Located in 586). dry areas where the following Recorded distribution.—Sas- grasshoppers are found : Melano- katchewan, Ontario, Idaho, Mon- plus angustipennis Dodge, M. tana, California, Illinois, Colo- flavidus Scudder, M. femurru- rado, Kansas, Texas and east brum (DeGeer), M. bivittatus through North Carolina, South (Say), M. sanguinipes (Fabri- Carolina, and Florida {225, p. cius), Ageneotettix deorum 586). (Map 77.) (Scudder), Schistocerca alutacea Host species.—Blaesoxipha kel- (Harris), and Mermiria neomexi- lyi (Aldrich) and Ceracia dentata cana (Scudder). (40, pp. 251- (Coquillett) {123, pp. 462-500; 254 -,41, p. 208 ; 152, p. 36 ; 164, p. 152, p. 36). \ Incidence of parasitism.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults appear in July and August. LOD : Unknown. OWS : Not given. G/Y: Probably more than one. SA: Grasshoppers infected with fly larvae. N/H : Not given. MOA : Spencer {316) reported that when the wasp is ready to oviposit, it chooses a grasshopper infected with a sarcophagid fly larva. The MAP 77.—Recorded distribution of grasshopper kicks, jumps, and Brachymeria coloradensis (Cresson). 132 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

UU2 ; SOO, V, 61 ; 806, pp. 28-82 ; U.S.S.R., and Asia. (225, p. 587 ; 807, pp. 258-259; 808, pp. 187- 267). (Map 78.) 199 \ 816, p. 501; 886) Host species.—Synthesiomyia Description.—Eggs, larvae, and nudiseta Wulp, Calliphore eolo- pupae unknown. Adult males are radensis Hough, Callitroga maeel- about 5.0 mm. long and black. The laria (F.), Phoenieia mexieana integument is densely punctured (Macquart), P. serieata (Mei- and covered with short golden gen), Phormia regina (Meigen), pubescence. The thorax is regular Blaesoxipha impar (Aldrich), and has dilated punctures. Apex Sarcophaga earnarea (L.), S. of the scutellum is emarginated haemorrhoidalis (Fallen), and S. and the tegula yellowish white. plenthopyga Wiedemann (225, p. Legs are rusty red brown. Coxae 587; 267). are black. The hindfemora of the Life history and habits.—SO: female are red with an apical Eggs are deposited from March to yellow spot, whereas in the male November. Larvae also occur they are black with the apex yel- from March to November. Pupae low, red, or both. Numerous small can be found any time from March black teeth are on the femora. to November. Adults also occur Abdomen is short, ovate, smooth, during the same time. LOD : Eggs' polished on the apical segments, total time to maturity is 8 to 12 and covered with a pale golden days. Larvae, pupae, and adults, pubescence. The malar ridge is unknown. OWS : At the last larval or the pupal stage. G/Y: Eight. directed backward. Wings are SA: Larvae. N/H: Not given. hyaline. (75, p. 60) MOA: Roberts (269) described Discussion.—Since most of the the attack of these parasites on sarcophagids are identified by certain fly larvae as occurring in adult characters and B. coloraden- one of two ways. Prior to oviposi- sis prevents host adult emergence, tion, in the first method, the fe- knowledge of its range is limited male wasp thoroughly examines mainly to Blaesoxipha kellyi. It the prospective host larvae by the has also been reported from Cera- use of its antennae. If satisfied, cia dentata. (168, p. 86) the wasp may then face away from the host and jab its ovi- Brachymeria fonscolombei positor into the host's body and (Dufour) insert eggs. The second method is that of grasping the host with all (Chaléis) 1841: 12 of its legs and forcing the ovi- Synonymy.—Chaléis fonseo- positor into the hemocoel. Both lombei Dufour (225, p. 587). of these methods require that the Recorded distribution.—South- host larvae be exposed to the ern United States. Also Europe, wasp. This would extremely limit ARTHROPOD AND NEMATODE PARASITES IN AMERICA 133 nodules in small chitinized circles. Mandibles are wide, flattened, and triangular. {23A, pp. 297-299) Prepupae larvae differ from pre- ceding stages by having the tho- racic segments larger, and the body is surrounded by sheaths from the skin of the last larval stage. The head alone appears as before. Pupae : The pupal stage is a continuation of the development of the prepupal. (23i, p. 299) MAP 78.—Recorded distribution of Adults are 41/2 to 6V2 mm. long ^ Brachymeria fonscolombei (Dufour). and a dull black. The head and thorax are punctured and cov- ered with a reclining greyish the time in which hosts could be pubescence. Antennae are inserted t parasitized, since larvae emerg- in pits near the middle of the ing from a grasshopper host face. The abdomen is pointed pos- generally burrow immediately teriorly, smooth, shining, and into the soil. However, Blaesoxi- covered with a light grayish pub- pha impar and Sarcophaga haem- escence behind and on the sides. orrhoidalis are also scavengers The posterior femora are reddish and may remain exposed for some with blackish bases and are en- time. AF: Unknown. HAB: Not larged and oval with about 10 given. (269, pp. 1-21) "teeth" on the inner edge. Pos- Description.—Eggs not given. terior tibiae are arched, also ^ Larvae (second instar) are 1.4 reddish, but with white extremi- mm. long by 0.3 mm. wide. They ties. Anterior femora and mid- '- are elongated ovoid with the ven- femora are black with white ex- ^ tral side convex and the dorsal tremities, and all tarsi are red concave. Segmentation is weak with black claws and empodium but evident. Very short small is black. Wings are small, clear, spines cover the body. The head and strongly punctured. is longer than wide, sclerotized Discussion.—Two of the hosts, but white, and the antennae are Blaesoxipha impar and Sarco- reduced. Mandibles are narrow. phaga haemorrhoidalis, are of The fifth-instar larvae are 5 mm. minor importance as grasshopper long by 2.5 mm. wide, short, oval, parasites. The third grasshopper and arched. The body is white, parasite, Sarcophaga carnarea, glossy, and distinctly divided into does not occur in the United 13 segments. The head is almost States. However, Blaesoxipha an- spherical. Antennae appear as gustifrons (Aldrich) has been 134 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE confused with S. carnarea in Can- 36; 281, p. 83; 307, p. 256; 316, ada and records may refer to it. p. 501). Roberts (269) reported that the Incidence of parasitism.—Not number of eggs produced by each given. However, it was collected female far exceeds 177. Unferti- at a high rate from fly larvae lized females produce viable eggs, during the summer of 1954 in but only male offspring result. Canada (316, p. 501). The period of development for Life history and habits.—SO : unfertilized eggs was 19.5 days. Eggs, larvae, and pupae unknown. Pupation requires 10 to 12 days. Adults not given. LOD : Unknown. The maximum number of genera- OWS and G/Y: Not given. SA: tions per year is believed to be Larvae. N/H: Not given. MOA: eight, with winter passed in hiber- Burks (4^1) indicated that al- nation in the last larval or pre- though the life history and habits pupal stage within a host pupa. of this species are not as well (88, p. 12) studied as those of B. coloraden- sis, he believed the two species to Brachymeria tegularis be closely related in their habits (Cresson) with only slight variation. AF: (Chaléis) 1872:60 In Canada it frequents "every Synonymy.—Chalcis tegularis clump of flowers on the ranges." Cresson (225, p. 587). HAB : In Illinois it is found as- Recorded distribution.—Brit- sociated with ordinary grassland ish Columbia, Wyoming, Texas, and field-crop grasshoppers. (H, Missouri, and Illinois (152, p. 36; p. 208; 281, p. 83; 316, p. 501) 225, p, 587). (Map 79.) Description.—Eggs, larvae, and Host species.—Acemyia den- pupae unknown. Adult females tata Coquillett (U, p. 208; 152, p. are about 5 mm. long and black. Labrum and base of the scape beneath are yellow, and the sides of the face are covered with dense silvery pubescence. The apex of the flagellum is reddish brown. Thorax is large and immaculate; it contains dilated punctures. Metathorax is coarsely reticu- late. Abdomen is short, ovate, smooth, shining, and immaculate; the sides and the apex are covered with pale pubescence. Legs are lemon yellow. The posterior coxae

MAP 79.—Recorded distribution of above, the apical half of the fe- Brachymeria tegularis (Cresson). mora within, and a row of minute ARTHROPOD AND NEMATODE PARASITES IN AMERICA 135 teeth are black. The posterior Ct^-VL.^K^| tibiae are black with a pale-yellow stripe behind, whereas the pos- terior tarsi are dull reddish brown. Wings are hyaline. {75, p. 60)

Perilampidae Perilampus hyalinus Say 1828: 79 Synonymy.—Perilampus acicu- latus Provancher (225, p. 518). MAP 80.—Recorded distribution of Recorded distribution.—Sas- Perilampus hyalinus Say. katchewan, Manitoba, Ontario, Quebec, and United States (225, Life history and habits.—SO : p. 518). (Map 80.) Eggs, larvae, and pupae unknown. Host species.—Blaesoxipha col- Adults not given. LOD: Eggs oradensis (Aldrich), B. hunteri hatch in 5 to 15 days. Larvae de- (Hough), B. kellyi (Aldrich). B. velop in 17 to 18 days. Pupae reversa (Aldrich), B. atlanis develop in 10 days. Adults un- (Aldrich) (Sarcophagidae), Ac- known. OWS: As larvae in the emya tibialis Coquillett, Ceracia host. G/Y: Several. SA: Larvae dentata (Coquillett) (Tachinidae), within their host. N/H: Not Acridomyia canadensis Snyder given. (Anthomyiidae), and more than MOA : About 250 eggs are laid 23 other insect species (123, pp. by a single female on the food ^72, 500; 16A; 389). plants of the host's host, or in Incidence of parasitism.—The this case the grasshopper. Free- effect of the hyperparasite on the living planidia emerge from the grasshopper parasite is often very eggs and attempt to find a grass- pronounced in localized areas. In hopper within the following 2 a valley 34 miles east of Hardin, weeks. The larvae enter the Mont., in 1967, 67 percent of the hemocoel and remain until the grasshopper parasites Blaesoxi- grasshopper is parasitized by the pha kellyi and B. reversa were proper species of fly. When the themselves parasitized, and in larvae of the parasitic fly is lo- 1968, 51 percent of the same spe- cated, it is entered by the planidia cies were eliminated by this wasp. of P. hyalinus. The planidia then In Missouri in 1959 P. hyalinus cease development while the fly parasitized 8.7 percent of Ceracia larva continues through three in- dentata from three Melanoplus stars within the grasshopper host. species. (152, p. 36) When mature, the fly larva exits 136 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE from its host, burrows into the long-ovate, and dark brown. They soil, and eventually pupates. At appear highly organized and have this point, the larva of P. hya- well-defined mouth parts, strong linus migrates outside the fly- curved mandibles, and a conspic- larva but remains within the fly- uous trachéal system. The first pupal case as an ectoparasite. It segment forms a distinct head and arranges itself so that its head is armed with several curved is in the same attitude as that of hooks followed by 12 well-differ- the host. Both larvae continue de- entiated segments. The dorsal part velopment, but the wasp larva is covered by a series of heavy eventually consumes much of the chitinized overlapping plates, fly larva. Development of the which nearly encircle the body. wasp within the pupal case gen- One to two pairs of bristles are erally extends over a longer pe- on the dorsum of most of these riod of time than normal fly de- segments, and ventrally there are velopment and emergence of the several backward projecting wasp is often 4 days to a week spines believed to aid in locomo- behind that of the normal host. tion. The head is heavily armored Although many planidia may be on the anterior margin with two found in a grasshopper or in a powerful hooks, which project dipterous host, only one per host laterally. In addition, two strong completes development. If the host hooks are located in about the has several generations per year, middle, curving backward at the so does P, hyalinus. tips. Posterior to these are several AF: Adult female wasps fre- semitransparent round spots that quently feed on flowers of the may be sensory in function. On group Compositae by puncturing the last segment of the posterior the epidermis of the leaf with end is a pair of unjointed stylets their ovipositor and ingesting the that are attached at about mid- sap that exudes from the wound. segment and are about twice the On occasion some frequent aphid- length of the segment itself. infested foliage and apparently After growth of the planidia feed on the honey dew. HAB. Not stage becomes rapid, the larva ap- given. (55, pp, 18, 215-220; pears very different. Mandibles do 126, p. 223 ; 30J^, pp. 8-55 ; 305, pp. not seem to enlarge. The mouth 6Í-68 ; 323, pp. 160, 282, 299, 321 ) parts are situated in a triangular Description.—Eggs of Perilam- basinlike depression, which is pus in the ovum tubes have been bounded on two sides and parts of described as elongate, oval shaped, the third by a strong chitinous not stalked, and whitish (304^, pp. rim. In addition to the mandibles 8-55). are two bulblike appendages just Larvae (first instar) or plan- beneath. Above the oral opening idia are about 0.3 mm. long, ob- are two large rounded elevations. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 137 which are probably the larval an- triangular, and very convex above tennae. The segments forming the and beneath. The scutellum is head are separated by a distinct much elongated and slightly neck and are bent beneath with emarginated. Tarsi are yellow the face downward. The first two with the anterior tibia a honey thoracic segments are each pro- yellow. The most obvious char- vided laterally with a raised acteristics of this wasp are its tubercle at about the middle part geniculate antennae and metallic of the segment. Beneath, the larva blue green color. Also the integu- is smooth and free from constric- ment appears to have been fash- tions although the segmental ioned with a ball peen hammer, structures are visible. Dorsally, especially in the thoracic region. segmentation is very evident be- (192, p. 382 \ 288, p, 79) cause of deep constrictions be- Discussion.—Because of its un- tween the segments. {SOi, pp. 8- usual method of contacting its 55) host, P. hyalinus is found in a Pupae resemble very closely the wide range of grasshoppers. Its adults. They are short and robust. presence appears to cause no dis- The abdomen has five to six trans- comfort to these insects. Adults, verse ridges, one for each seg- for example, of Melanoplus bivit- ment. Young pupae are totally tatus (Say) generally do not allow white or yellowish white, but as development of parasites within they age the eyes become reddish them and usually cause the death and the thorax becomes black, of this parasite with an eventual later changing to metallic blue melanization. However, first-in- and blue green. The pupal case is star planidia of P. hyalinus ap- generally transparent. At the parently cause no physiological completion of maturation the head reaction with the grasshopper and and abdomen attain the same me- can be found still in the first- tallic hue as the thorax. During instar planidia stage within this later stages of development, legs, species, apparently quite healthy antennae, and mandibles are and active. Only after one of the movable within their sheaths, and dipterous parasites has attempted twitching and movement can be to parasitize a grasshopper host seen in exposed pupae. Shortly will the physiology of the grass- after the wasp has attained the hopper cause the death of the complete metallic coloring, it wasp larva. sheds the pupal skin and emerges First-instar larvae of this wasp as an adult. {SOJ^y pp, 8-55 ; S89) are small enough that even under Adults are less than 0.2 inch low-power microscopes they may long, metallic green, and punc- be often missed. Their appearance tured. They have hyaline wings. is unusual in that they do not re- The abdomen is very short, wide. semble any of the dipterous para- 138 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE sites of grasshoppers. Instead their general appearance is that of a thysanurous or possibly a coleopterous larva except the head. Higher magnification re- veals the presence of rather scle- rotized areas as can be found in many Coleóptera. Movement is of an up-and-down nature as op- posed to side to side, and the rest- ing position is humped similar to that of a shrimp. (389) MAP 81.—Recorded distribution of Pteromalidae Eupteromalus sarcophaga Gahan. Eupteromalus sarcophaga male is brownish yellow. Apex of - Gahan the scutellum is more faintly r 1914: 162-163 sculptured than that of the re- Synonymy.—None (225, p. mainder, somewhat shining, and 554). with shallow reticulate punctures. Recorded distribution.—Kan- The abdomen of the male is sas and Michigan (225, p. 554). scarcely longer than it is broad, (Map 81.) with the first segment wider at Host species.—Blaesoxipha kel- the apex than the base of the sec- lyi (Aldrich) (325, p. 67). ond. Segments beyond the first Incidence of parasitism.—Not fire scarcely longer than the first. given. Coxae, trochanter, and femora Life history and habits.—SO: are black, the tibiae of variable Eggs, larvae, and pupae unknown. brownish to black, and the bases ' Adults not given. LOD : Unknown. and apices yellow. Except the api- G/Y, SA, N/H, MOA, AF, and cal joint, the tarsi are yellow to HAB : Not given. reddish yellow. Wings are hyaliiie Description.—Eggs, larvae, and and the stigmal vein is nearly as pupae unknown. Adult females long as the marginal. Venation of are 1.9 mm. long and dark green the wings is pale. (117, pp. 162- to brassy green or yellow. The 163) scape and legs except the coxae are reddish yellow and the apical Nasonia vitripennis (Walker) tar sal joints are brown. The head (Pteromalus) 1836: 492 and thorax are covered with close Synonymy.—Pteromalus vitri- thimblelike punctures. The abdo- pennis Walker, P. abnormis Bohe- men is smooth and conic-ovate. man, Mormoniella previcornis The antennal flagellum of the Ashmead, Nasomia previcornis male is black and that of the fe- Ashmead (225, p. 559). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 139 Recorded distribution.—Most by a chitinized ring. Mandibles of Canada and the United States are curved and pointed but broad including Alaska {225, p. 559). at the base and articulate. An- (Map 82.) tennae are very small. The respi- Host species.—Musca domes- ratory system contains lateral tica Linnaeus, Phormia regina trunks with four pairs of open (Meigen), and many sarcophagids spiracles, one on the thorax and U7, pp. 495-501 ; 87, pp. 151, 153, three in the abdominal region. 155, 158; 119, pp. 119-132; 123, The second instar is similar to the p. 500; 316, p. 501 ; 323, pp. 291, first. (55, pp. 121, 12U, 126-129 ; 301,332). 95, pp. A00~W3). Pupae un- Incidence of parasitism.—Not known. given. Adult females are 2.12 mm. Life history and habits.—SO: long and shining bronze to black. Eggs, larvae, and pupae unknown. Antennae are dark, clávate, thin, Adults not given. LOD: Eggs and divided. The second joint is hatch in 4 days. Larvae : First in- dark yellowish brown and the star requires 1 day, second instar first is yellow brown. Head is 2 days, and third instar 3 days. green black and has reddish-black Pupae and adults unknown. OWS ocelli. The body is short and and G/Y : Not given. SA : Pupae. broad, and the thorax is as wide N/H : Up to 36. MOA : This para- as the head. The thorax is ovate, site is able to seek its prey and the metathorax short, and the ab- oviposit in total darkness. Spen- domen ovate and shining. Feet cer (316) indicated that when the are yellow bronze and the femora eggs have been deposited within dark. Coxae are bronze and the a pupal case, an odor is left that midfeet, hindfeet, tibiae, and prevents attraction of additional apex of the tarsi are golden yel- females. The larvae eat and ma- ture on the contents of the host and all exit from a single host. AF and HAB: Not given. (47, pp. 495-501 ; 80, pp. 365-369 ; 87, pp. 151, 153, 155, 158; 90, pp. 88- 112;119, pp.119-132;120, pp. 9- 28; 268, pp. 307-309, 312; 316, p. 501 ; 323, pp. 291, 301, 332) Description.—Eggs unknown. Larvae are extremely fragile. The thorax is broad and tapers pos- teriorly. There are 13 posterior cephalic segments. The head is MAP 82.—Recorded distribution of greatly reduced and surrounded Nasonia vitripennis (Walker). 140 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE low. Wings are clear whitish with the stigma small and unicolor. {8Jt8, p. A92) Discussion.—There are many reports on laboratory research of this species, including its histol- ogy and internal anatomy.

Sphecidae Oxybelus quadrinotatus Say 1824: 338 Synonymy.—Oxybelus impa- MAP 83.—Recorded distribution of Oxybelus quadrinotatus Say. tiens Smith, 0. interruptus Cres- son, 0. brodiei Provancher (225, Description.—Eggs, larvae, and p. 1033). pupae unknown. Adults are 6 to 7 Recorded distribution.—South- mm. long and basically black. The ern Canada and the United States antennal attachment is toward the (225, p. 1033). (Map 83.) tip and beneath on the head. Tho- Host species.—Musca domes- rax is immaculate. Metathorax tica Linnaeus, Phyra leucostoma and scutellum each bear three Wiedemann, Sarcophaga rapax raised spines ; two superior spines Walker, Blaesoxipha hunteri of the former are whitish at the Hough, and Hylernya cilcrura tip and decurved; the inferior (Rondani) (123, p. 500; 169, p. spine is larger and black. Tergum 67; 257, p. 101). is polished. Tarsi are testaceous Incidence of parasitism.—Not at the tip and the anterior tibiae given. are testaceous on the inner side. Life history and habits.—SO: Wings are hyaline and the veins Eggs, larvae, and pupae unknown. brown. (289, p. 338) Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Stizoides unicinctus (Say) Adults. N/H: Not given. MOA: (Stizus) 1823: 77 Prey is usually impaled upon the Synonymy.—Stizus unicinctus ovipositor where it remains while Say (225, p. 99^). the burrow is being constructed. Recorded distribution.—Mich- AF: Nectar. HAB: It nests in igan through Arkansas and Texas, loose cinder or sand-type soil, dig- west through California, north ging shallow burrows 2 to 3 inches through Washington, and east long at a 45° angle with the sur- into Michigan (225, p. 994-). face. (55, p. 332; 169, p. 67; (Map 84.) 235, pp. 7U-75; 257, p. 101; 322, Host species.—Utilizes the p. 313) grasshoppers taken by Sphex ARTHROPOD AND NEMATODE PARASITES IN AMERICA 141 atria Lepeletier {123, pp. AGO, 500; 256, p. 11; 257, pp. U, 180- 193; 369, p. 230). Incidence of parasitism.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Eggs. N/H: Not given. MOA: Observations by Rau (256; 257) and Williams {369) indicate that this wasp waits until after a nest MAP 84.—Recorded distribution of of Sphex atria has been stocked Stizoides unicinctus (Say). with a grasshopper and sealed. It then reexcavates the nest, de- structed by Priononyx, which still stroys the egg of S. atria, and de- had the prey, a specimen of Aulo- posits one of its own. It closes the cara, near the burrow. When burrow entrance with no attention Priononyx returned and discov- to neatness, camouflage, or com- ered S. unicinctus, it left the prey pletion. Therefore the nest is and burrow to the latter. AF and often destroyed later by other in- HAB: Not given. {257, pp. i, sects such as ants, which are able 180-193; 369, p. 230) to enter the partially closed en- Description.—Eggs, larvae, and pupae unknown. Adult males are trance. In this way S. unicinctus about 12.7 mm. long, broad, and does more harm to the predators black. The abdomen is polished of grasshoppers than it does to with a rufous band above on the the grasshopper population it- basal half of the second segment. self. Williams {369) reported an The anal spines are prominent and incidence where S. unicinctus the wings are dark violaceous. merely entered a burrow con- {192,pp.A,165;28Jt,p.77)

Díptera Calliphoridae Recorded distribution.—South- Senotainia ruhriventris central and southeastern Canada Macquart and the United States {320, p. 1846: 295 939). (Map 85.) Synonymy.—Miltogramma de- Host species.—Sphex pubidor- cisa Townsend (4, P- 4-4-7; 320, p. sus (Costa) {97, pp. 177-186; 939). 123, p. 500). 142 TECHNICAL BULLETIN NO. 14^0, U. S. DEPT. OF AGRICULTURE There is a tuft of white hair on each side. The fifth segment is black and also contains white tufts of hair. Feet are black. Haltères are white and wings hyaline. {212, p. 295) Taxigramma heteroneura (Meigen) (Miltogramma) 1830: 367 Synonymy.—Miltogramma het- eroneura Meigen, Heteropterina MAP 85.—Recorded distribution of nasoni Coquillett (320, p. 9^0). Senotainia rubriventris Macquart. Recorded distribution.—South- ern Canada and the United States Incidence of parasitism.—Not except the mid-Atlantic and given. Southeastern States (320, p. 9W). Life history and habits.—SO: (Map 86.) Eggs, larvae, and pupae unknown. Host species.—Tachysphex ten- Adults not given. LOD : Eggs re- uipunctus Fox (123, p. 500; 228, quire about 1 day to hatch. Larvae pp. 615-619). develop in 4 days. Pupae not Incidence of parasitism.—Not given. Adults unknown. OWS, given. G/Y, SA, and N/H: Not given. Life history and habits.—SO: MOA: S. rubriventris female Eggs not given. Larvae and pupae utilizes the prey of Sphex pubi- unknown. Adults not given. LOD: dorsus as well as its egg by de- Unknown. OWS, G/Y, SA, N/H, positing its own eggs before clo- MOA, AF, and HAB : Not given. sure of the cell. Both the wasp egg and grasshopper egg are utilized. {97, pp. 177-186). AF and HAB : Not given. Description.—Eggs, larvae, and pupae unknown. Adults are 5 to 6 mm. long and basically gray to black. Face of the female is white and front is white to yellowish with black streaks. Antennae are yellow with the apex of the second segment black. Thorax is ash gray with a black pile. The first seg- ment of the abdomen is black ; the MAP 86.—Recorded distribution of second, third, and fourth are red. Taxigramma heteroneura (Meigen). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 143 Description.—Eggs, larvae, and The abdomen is punctured and pupae unknown. Adults are 2.65 whitish behind. Bristles are dense mm. long and basically gray. An- behind and are directed poste- tennae and legs are orange and the riorly. {218, p. 367) feet brown. The frons is brownish.

PREDATORS

A predator is an that insects jumping or on the wing. utilizes several or more In the case of these flies the prey as a source of food or nutrition may be equal in size to the pred- during its lifetime. Insect pred- ator or larger, but the strong fly- ators may completely devour the ing ability of the robber fly seems insect body or the egg as the cic- to make the collection, subduing, indelids or carabids, or they may and evacuation of body contents require only the contents of the a simple task. hemocoel as the asilids and spid- Predators may have both ad- ers. In either situation the prey vantages and disadvantages as forfeits its life. control agents. Whereas a pred- Occasionally a predator prefers ator population may remain high a definite prey, as the Stenopogon and relatively constant because species, which concentrate on of the availability of many types grasshoppers. More often, how- of usable food, its efforts are gen- ever, as shown by most of the Ef- erally not concentrated toward feria group, the diet includes the elimination of one particular most flying insects in addition to prey species or group of species. grasshoppers. Beetles are gener- Egg predators probably have ally not preferred, and yet flies of more effect on grasshopper pop- the same genus or even the same ulations than do predators of species are relished. nymphs and adults. Some species The predaceous nature may that are predaceous during part take many forms. Larvae of of their life are pests during an- asilids, therevids, carabids, and other part. In the case of the bombyliids while seeking food ac- meloids, the adult beetles are of cidently locate and utilize grass- economic importance since they hopper egg pods. Adults and lar- attack garden and rangeland vae of the cicindelidae often lie in crops. Therevid, asilid, and cara- wait and attempt to ambush their bid larvae, although destructive prey. Robber fly (Asilidae) adults upon the egg pods of grasshop- generally need the attraction of pers, also prey on various soft- movement to stimulate their at- bodied insects. This leaves the tack and usually take only those bombyliids, or bee flies, which at- 144 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE tack only grasshopper egg pods, their larval and adult stages, and which appear to be as the most spiders utilize prey during all but dependable. their egg stage. The meloids, Generally a predator collects carabids, bombyliids, and chloro- prey over a considerable period pids spend much of their life in of its life. The asilids and cicinde- the larval stage feeding on their lids are predatory during both prey.

Coleóptera Carabîdae eggs and soft-bodied insects. Usu- ally only the body contents of The ground beetles are predace- the prey are consumed or, as in ous in both their larval and adult the case of the eggs, the yolk. stages. Adult beetles are nocturnal Strong piercing mandibles can in nature and shun the daylight mash all the eggs in the egg pod, by hiding in cracks, crevices, after which the contents are in- holes, burrows, and under stones, gested, or they can pierce and debris, and loose bark of trees. crush soft-bodied insects and Most are ground dwellers, al- make the exuded body fluids avail- though a few species of the genera able for consumption. The adult, Calosoma and Lebia are aboreal on the other hand, can move much in habit. faster and either chases or am- Adults are long legged, strong bushes its prey. The adults are mandibled, and usually dull in not considered as important pred- color. Shortly after mating, adults ators as are the larvae. deposit eggs in small holes. Lar- The only attempt to assess the vae are generally long and the effect of this group in the litera- body is nearly equal in width ture was by Parker and Wakeland throughout. Mandibles are sharp (236), They combined informa- and projecting, and the caudal tion from the former ARS Plant end usually bears a pair of coni- Pest Control Division with their cal bristly appendages. Larval own and other egg-survey data. and adult forms occupy very sim- The average percentage of egg ilar habitats and in many ways pods destroyed by act similarly. larvae in 25 States ranged from a The larvae pupate in prepared cells within the soil and may be high of 17.7 percent in Washing- a foot or more under the surface. ton in 1938 (dropped to 2.3 per- A generation usually requires 1 cent in 1939) to 0 in Mississippi full year and often the adult beetle for both 1938 and 1939. From 16 lives an additional year. established plots in seven States Young larvae feed on insect they found that the average an- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 145 nual destruction of grasshopper egg pods by ground beetles was 2.89 percent, and the most com- mon species were Amara obesa (Say) and A. hesperia Casey. The highest percentage of egg pods destroyed by ground beetle larvae per type of land was idle land (9.55 percent) in north-central Montana, rangeland (14.44 per- cent) in North Dakota, and in le- gumes (6.6 percent) in South Da- kota. Ground beetles in Minnesota MAP 87.—Recorded distribution of destroyed more egg pods than bee Agonoderus comma (Fabricius). flies (Bombyliidae) or blister beetles (Meloidae). Incidence of prédation.—Not Shotwell (298) reported that in given. the fall of 1938 from an 11-State Life history and habits.—SO: survey ground beetle larvae were Eggs, larvae, and pupae unknown. present at the rate of 0.06 per Adults not given. LOD : Unknown. square foot. L. P. Rockwood (in OWS and G/Y: Not given. SA: 236, p. 27) observed the digging Eggs : N/H, MOA, AF, and HAB : and feeding habits of A. obesa in Not given. (262, p. 289) Camnula pellucida (Scudder) egg Description.—Eggs, larvae, and beds. Nevertheless there is some pupae unknown. Adults are 5.5 to question of the eflfectiveness of 7.5 mm. long and 2.0 to 2.6 mm. carabids in seeking grasshopper wide. The body is rather large, egg pods as a main source of food. stout, and oblong. Head and un- dersurface are black with the Agonoderus comma pronotum and elytra yellowish (Fabricius) brown to reddish brown. There is (Carabus) 1801: 207 a large black spot on the disk of Synonymy.—Carabus comma the pronotum. The antennae are Fabricius, Feronia tallies Say, stout and extend behind the tho- Acupalpus pallies Say, Agono- racic base. The thorax is wider derus dorsalis Jaques (195, p. 75). than it is long and is widest near its apical third. Sides are strongly Recorded distribution.—East- rounded and become oblique and ern United States (H, p. 295 ; 86, feebly arcuate basally. Elytra are pp. 61, 125-126; 159, p. 52). deeply striated. The hindcoxae (Map 87.) are continguous and the middle Host species.—"Locusts" (123, coxal cavities are entirely en- p. 4-90; 159, p. 52; 26U, p. 289). closed by the sterna alone. (^^, 146 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE V. 295; 86, pp. 61, 125-126; 10k, The head is less than half as wide p. 207; 159, p. 52) as the prothorax ; both are sinuate - but the head is slightly more so. , Amara hesperia Casey Striae are somewhat fine. (k5, 1918: 298 p. 298) Synonymy.—None {195, p. 61). Discussion.—Parker and Wake- Recorded distribution.—Ore- land (236, p. 3) reported it was gon, California, Nevada, Utah, among the most common species "^ southern Colorado, and New Mex- of grasshopper egg-pod predators , ico U5, pp. 298, 312; U6, p. 200; found during their tests in Wash- 195, p. 61). (Map 88.) ington. *" Host species.—Grasshoppers (236, p. 3). Amara impuncticollis (Say) Incidence of prédation.—Not (Feronia) 1823: 36 given. Synonymy.—Feronia impuncti- Life history and habits.—SO : collis Say, (genus ?) truvuakus ^ Eggs, larvae, and pupae unknown. Dejohn, (genus ?) anthracina Adults not given. LOD : Unknown. Haldeman (195, p. 61). OWS and G/Y: Not given. SA: Recorded distribution.—South- . Eggs. N/H, MOA, AF, and HAB : ern Canada, Alaska, United States Not given. (236, p. 3) east of the Rocky Mountains, Ore- Description.—Eggs, larvae, and gon, and California (159, p. 37; >. pupae unknown. Adults are 7 to 9 195, p. 61). (Map 89.) mm. in length and a brassy Host species.—"Grasshopper" bronze color. Antennae are rather (123, p. k90; 236, p. 27; 328). long. Elytra are similar to the an- Incidence of prédation.—Not terior parts of the body and more given. ' than one-half longer than wide. Life history and habits.—SO; Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Eggs. N/H, MOA, AF, and HAB : Not given. (236, p. 27) ' Description.—Eggs, larvae, and pupae unknown. Adults are 7 to 10 mm. long, dark bronze to black- ish bronze with the underside black, oval and somewhat convex. The head is slightly narrower than the thorax at its apex. Antennae are brown and the three basal MAP 88.—Recorded distribution of joints red. Palpi are black or red- Amara hesperia Casey. dish black. Elytra are striated ARTHROPOD AND NEMATODE PARASITES IN AMERICA 147

ish Columbia and the United States (195, p. 60; 236, pp. 3, 27; 336,p. 35; 364, pp.35, AO). (Map 90.) Host species.—Camnula pellw- cida (Scudder) and grasshoppers (123, p. 490; 236, pp. 3, 27; 264, pp. 290-292; 330, pp. 178, 182; 336, p. 35; 364, pp. 35, 40). Incidence of prédation.—Not given. However, it is believed that when eggs are available, the inci- dence is rather high. MAP 89.—Recorded distribution of Life history and habits.—SO: Amara impuncticollis (Say). Eggs are deposited in the summer. Larvae occur in the fall, winter, but without dorsal punctures. The and spring. Pupae unknown. body beneath is black and impunc- Adults not given. LOD : Unknown. tured; the margins are not de- OWS: Larvae. G/Y: Not given. pressed. Feet are piceous black; SA: Eggs. N/H and MOA: Un- the hindtibiae of the males have known. AF: Not given. HAB: a pubescence on the inner side. During the day adult beetles may Apical spur on the front tibia is be found under stones, dirt clods, simple. Femora are piceous, tibiae or grass. (77; 133, pp. 49, 51- and tarsi are rufous or red, and 52; 236, pp. 3, 27; 264, pp. 290- the midtarsi and hindtarsi have 292; 300, p. 60; 336, p. 35; 364, the basal joint grooved on the pp. 35, 40) outer side. Head is not constricted Description.—Eggs unknown. behind the eyes. Mandibles lack a bristle-bearing puncture in the groove on the outer side, but there are two punctures above the eye with a single heavy bristle in each. One or more bristly hairs are found above each eye. (133, pp. U9, 51-52; 159, p. 37; 193, pp. 463, 542; 287, p. 36)

Amara obesa (Say) 1823: 37 Synonymy.—Percosia patricia LeConte (195, p. 60). MAP 90.—Recorded distribution of Recorded distribution.—Brit- Amara obesa (Say). 148 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Larvae average 14 mm. long and dish brown. {193, pp. UßU, 5U2; are dull white. Head and protho- 262, pp. 35, hO ; 287, p. 37) rax as viewed from above are Anisodactylus calif or nicus orange yellow. Body is widest at DeJean the middle and tapers suddenly from joint 10. Joints 2 to 12 each 1829: 148 contain a dorsal transverse corne- Synonymy.—None {195, p. 72) ous plate, which is deeper brown- Recorded distribution.—Cali- ish yellow. On each side of all fornia {195, p. 72). (Map 91.) plates are two distinct impres- Host species.—Grasshoppers sions. The head is flattened and {236, p. 27). depressed. Mandibles are moder- Incidence of prédation.—Not ately stout with a small middle given. tooth. Antennae are four-jointed; Life history and habits.—SO the third is bent inwardly and the and LOD: Unknown. OWS and fourth outward. The second joint G/Y : Not given. SA : Eggs. N/H : of the maxillae is slightly shorter Unknown. MOA : Larvae chew on than the basal joints. Palpi are egs pods. AF: Not given. HAB: two-jointed. Prothoracic joint is Not located. {236, p. 27) rounded and faintly striated at Description.—Eggs, larvae, and the front and hind borders. The pupae unknown. Adults: Dejean legs are pale and contain an "X"- described this species by compar- shaped black mark on the outside ing it with Anisodactylus binota- of the coxae. Tarsi are distinctly tus DeJean and indicating the dif- two-clawed and nearly as long as ference. A. californicus is about the tibia. The anal proleg is hori- 101/2 mm. long by 4 mm. wide and zontal and rather obliquely trun- colored similarly to A. binotatus. cate beneath. {26A, pp.290-292) The tarsi are blackish brown. Pupae are unarmed, elongate, and the tips of the tarsi are broad and distinctly bifid (26A, pp. 290- 292). Adults are about 10 mm. long and black. Underneath is piceous black with a reddish tinge, and the antennae, mouth, and feet are reddish to reddish-brown. The thorax is slightly margined and quite punctured at the base. The dorsal line is slightly depressed and the basal line is distinct.

Elytra with striae are very dis- MAP 91.—Recorded distribution of tinctly punctured. Feet are red- Anisodactylus californicus DeJean. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 149 There are two reddish spots be- tween the eyes and a blackish spot above the yellow testaceous an- tennae. Eyes are less projecting than in A. binotatus, and the punctuation of the integument is less. The eighth and ninth ab- dominal segments lack bristles and deep spots. (8^, p. H.8) Discussion.—It was reared by C. C. Wilson from larvae and pu- pae found in or near grasshopper egg pods. It is a possible predator. MAP 92.—Recorded distribution of {236, p. 27) Anisodactylus nivalus Horn.

Anisodactylus nivalus Horn pale tawny yellow; it has a large 1880: 172 blackish macula. There are some V Synonymy.—None. fine scattered punctures on the Recorded distribution.—Cali- abdomen at the base of all but the fornia and Utah (4.4., pp. 203- last segment. Femora are black- 204). (Map 92.) ish. Tibiae and tarsi are testace- Host species.—Grasshoppers ous. iU, pp. 203-204) {236, p. 27). Incidence of prédation.—Not Callisthenes latipennis Horn given. 1870: 69-70 Life history and habits.—SO: Synonymy.—Callisthenes opi- Eggs, larvae, and pupae unknown. mvs Casey {195, p. 45). Adults not given. LOD : Unknown. Recorded distribution.—Cali- OWS and G/Y: Not given. SA: fornia {195, p. 45). (Map 93.) Egg pods. N/H, MOA, AF, and Host species.—Oedaleonotus HAB : Not given. enigma Scudder and Sticthippus Description.—Eggs, larvae, and californicus Scudder {330, p. 183). pupae unknown. Adult females Incidence of prédation.—Not are about 100 mm. long, oblong- given. oval, and black with a pale dia- Life history and habits.—SO: phanous color on the elytra and Eggs, larvae, and pupae unknown. margins of the thorax. The head Adults not given. LOD : Unknown. is short and two-thirds as wide OWS and G/Y: Not given. SA: as the thorax. Eyes are promi- Nymphs. N/H: Unknown. MOA: nent. Antennae are pale black. Attack nymphs and eggs. AF : In- Elytra are oblong-oval and one- sects. HAB : Not given. {42, pp. third wider than the prothorax. 156-160; 330, p. 183) The surface is "opaculate" and Description.—Eggs, larvae, and 150 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Host species.—Melanoplus spretus (Walsh) and the army- worm Cirphis unipuncta Haworth (123,p.490;264,p.314). Incidence of prédation.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults are active from May to September. LOD : Eggs unknown Larvae: First instar 3 days, sec- ond instar 4 days, and third instar MAP 93.—Recorded distribution of unknown. Pupae and adults un- Callisthenes latipennis Horn. known. OWS: Adults. G/Y: One generation lasts 2 to 3 years. SA : pupae unknown. Adults are 15 to Eggs. N/H: Unknown. MOA: 18 mm. long and deep black. Head Larvae encounter food. A/F : Not is small and one-half as wide as located. HAB : None given. Adults the prothorax. Antennae are long hibernate from September to May and slender, longer than the width and live 2 to 3 years. They are of the thorax. Prothorax is one- difficult to rear in captivity. (39, half as long as it is wide and two- pp. 21-25; 160, p. 170; 344, p. thirds to three-fifths narrower 18) than the elytra. The elytra are ob- Description.—Eggs when long-oval. The foveae are small freshly laid are white to faint yel- and cupreous. Body surface is lowish and average 5.4 mm. by 2.5 shining, smooth, and convex with mm. They are slightly elliptical sparse punctures. Tarsi are slen- and taper toward one end but der. (43, p. 69) Discussion.—It can apparently reduce the nymphal population in California. Calosoma externum (Say) (Carabus) 1823: 150 Synonymy.—Calosoma longi- penne DeJean, Carabus externum Say (195, p. 45). Recorded distribution.—East- ern United States with the high- est population in the South (39, pp. 21-25; 160, p. 26; 195, p. 45). MAP 94.—Recorded distribution of (Map 94.) Calosoma externum (Say). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 151 vary greatly in form, ranging Calosoma scrutator (Fabricius) from long, almost cylindrical, and (Carabus) 1775: 239 tapering slightly toward both Synonymy.—Carabus scrutator ends to short, oval, and kidney Fabricius (195, p. i5). shaped. (39, pp. 21-25) Recorded distribution.—South- Larvae (first instar) are short, central and southeastern Canada stout, and approximately 7.7 mm. and the United States (9, p. 101; long. They are black above with 39, pp. 26-38; 86, pp. 65-66; 159, the ventral plates dark to grayish p. 26). (Map 95.) brown. Caudal appendages are Host species.—"Grasshoppers," straight, slender, and support nu- armyworms, tent caterpillars, and merous short spines. Second in- canker worms (39, pp. 26-38 ; 123, stars are also short, very broad, p. 4-90; 264-, p. 3U; 328, p. 16). and approximately 13 mm. long. Incidence of prédation.—Not Color is similar to that of the first given. instar and the caudal appendages Life history and habits.—SO: are very long, the same length, or Eggs, larvae, and pupae unknown. slightly larger than in the first Adults not given. LOD : Eggs 5 to instar. Ends of the tergite pro- 7 days average. Larvae: First in- ject well laterad; the posterior star 3 to 4 days, second instar 3 angles of the same are obtuse and to 10 days, and third instar 14 to project prominently backward. 15 days. Prepupae 1 week. Pupae Third-instar larvae are robust 13 days. Adults 2 to 3 years from and average about 20.5 mm. long. egg to end of adult stage. OWS : They are dull black on the dorsal Not given. G/Y: A generation part and the ventral plates are lasts 2 to 3 years. SA : Eggs. N/H : dark brown to grayish brown. Unknown. MOA: Larvae encoun- Caudal appendages are long, slen- ter eggs. AF : Not located. HAB : der, and spread widely apart at the tips. {39, pp. 21-25) Pu- pae unknown. Adults are 28 to 32 mm. long and deep black with a purplish margin. The elytra are black with a blue border. Both the thorax and the elytra have blue side margins. The margins of the elytra are par- allel for three-quarters of their length and the lateral edges are curved at their base. Elytra are striate with conspicuous definite punctures. (159, p. 26; 192, p. MAP 95.—Recorded distribution of 97; 197, p. lAO; 285, p. 150) Calosoma scrutator (Fabricius). 152 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Often found in large numbers and reddish hair near the tip. The are able to climb trees. (9, p. thorax is impunctured and the 101 ; 39, pp. 26-88 \ 86, pp. 65-66 disk is blue to dark purple with 9i, pp. 519, 521, 585 \ 159, p. 26 margins of gold or red. Elytra 160, pp. 170, 192 \ 26U, P^ 8U are bright green with very slight 8U, p. 19) cupreous reflections. Abdomen is Description.—Eggs are white green or red and the legs are blue. with a faint yellow tinge, nearly The male is distinguished by a elliptical, and taper slightly to- dense brush of hairs on the inner ward one end. They are approxi- surface of the curved middle tibia mately 5 mm. long. {89, pp. 26- near the apex. There are 15 striae, 88). Larvae are robust, very and transverse lines are very nu- dark brown to black, with light- merous. The fourth, eighth, and brown ventral plates. They aver- twelfth interstitial lines each con- age 9.5 mm. long. Caudal append- tain several distinct impressed ages are of medium length and punctures. External margins are rather stout. Second-instar larvae a reddish-cupreous color. {86, are similar to the first in shape pp. 65-66 ; 102, p. 289 ; 159, p. 26 ; and color and have an average 198, p. Jf91) length of 16.9 mm. Caudal ap- Discussion.—Not enough infor- pendages are also short, stout, mation is available. This species and straight. Third-instar larvae prefers caterpillars. Since it can are robust, dark brown, and black. climb trees in search of them, it They average 25.6 mm. in length. has been called searcher or cater- Caudal appendages are straight pillar hunter. and very stout near the base. Ven- tral plates are light brown, and Calosoma tepidum LeConte there is a reddish-brown patch at 1848: 199 the base of the caudal appendages in the second and third instars. Synonymy.—Calosoma caelator {89, pp. 26-88) Casey, C. indigens Casey, C. ir- Pupae when young are creamy reguläre Walker {195, p. i6). white and 20 mm. long. This color Recorded distribution.—Brit- later changes to slight violet ex- ish Columbia and Pacific Coast cept the abdomen, which remains States east into Idaho and south creamy white. {89, pp. 26-88) into Arizona {185, p. 199; 195, Adults are 27 to 32 mm. long, p. i6 ; 8Í9, p. 8i2). (Map 96.) metallic, and highly iridescent Host species.—Possibly preda- throughout. The underside of the ceous on grasshopper eggs {886, body is green varied with a p. 85). golden-cupreous color. Head is Incidence of prédation.—Not impunctured and black with the given. orbits above gold. Antennae have Life history and habits.—SO ARTHROPOD AND NEMATODE PARASITES IN AMERICA 153

Calosoma ivilcoxi LeConte 1848: 446 Synonymy.—None (195, p. 4.5). Recorded distribution.—South- ern Canada and the United States (only specific Canadian reference was that of southern Ontario) (39, pp. 38-U; 159, p. 26). (Map 97.) Host species.—Melanoplus spretus (Walsh), canker worms, certain caterpillars, and others MAP 96.—Recorded distribution of (123, p. Í90; 26U, p. 3U; 328, p. Calosoma tepidum LeConte. 17). Incidence of prédation.—Not given. and LOD: Unknown. OWS and Life history and habits.—SO: G/Y : Not given. SA : Eggs. N/H : Eggs, larvae, and pupae not given. Unknown. MOA: Larvae encoun- Adults are first seen from May to ter eggs. AF : Not located. HAB : June. LOD: Eggs 3 to 12 days. Not given. (336, p. 35) Larvae: First instar 7 days, sec- Description.—Eggs, larvae, and ond instar 5 days, and third in- pwpae unknown. Adults are ap- star 14 days. Prepupae 4 to 5 days. proximately 25 mm. long and gen- Pupae 2 weeks. Adults not given. erally black. Underneath is black. OWS : Adults. G/Y : One genera- Head is rugulose in front and con- tion takes 3 years. SA: Eggs. tains an impression and a retuse N/H: Unknown. MOA: Larvae border on each side. Thorax is encounter eggs. AF : Not located. more finely rugulose than the head with an impressed medial line. Elytra are less finely rugulose than the head, each containing three diskal lines of gilded points and with a submarginal line of smaller points. Since this species does not contain striated lines on the elytra, it is separated from C. calidum Fabricius and C. frigi- dum Kirby. (185, p. 199; 3^9, p. 3i2) Discussion.—It is only a pos- sible predator of grasshopper MAP 97.—Recorded distribution of eggs. Calosoma wilcoxi LeConte. 154 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

HAB: Not given. Eggs are laid ner surface of the curved middle in the soil. Adults of this species tibiae. The pronotum has a trans- apparently climb trees in search verse impressed median line, the of food, although it is believed sides are broadly curved, and the that the larvae are ground dwell- base is narrower than the apex. ers. (9, p. 101; 39, pp. 38-U; Elytra contain fine punctured 86, pp. 65-66; 159, p. 26; 26U, p. striae. This species is very simi- 3H) lar to C. scrutator but is distin- Description.—Eggs are white guished by its smaller size and the with a faint yellowish tinge and somewhat narrower thorax. (9, average 3.9 mm. long. They are p. 101; 86, pp. 65-66; 159, p. 26; elliptical and sometimes kidney 185, p. U6) shaped but often taper toward one end. {39, pp. 38-H) Larvae Elaphrus ruscarius Say (first instar) are somewhat stout, 1834: 417 brown above with light-brown Synonymy.—Elaphrus ameri- ventral plates. They average 7.8 canus DeJean, E. riparius Say mm. long. Caudal appendages are {195, p. i6). of medium length and straight. Recorded distribution.—South- Second-instar larvae are stouter western and south-central Canada than the first stage, slightly and western and central United lighter in color, and approxi- States {195, p. A6). (Map 98.) mately 12.9 mm. long. The caudal Host species.—"Grasshoppers" appendages are short and stout. {264,p.31i). Third-instar larvae are robust Incidence of prédation.—Not and chestnut brown above with given. ventral amber plates. A reddish- Life history and habits.—SO brown patch is at the base of the and LOD: Unknown. OWS and caudal appendages of the second and third instars. Third instars are 22 mm. long. {39, pp. 38-U) A. ^AftÇ^:;] Pupae are approximately 16.5 mm. long by 7.5 mm. wide (39, pp. 38-U). Adults are 17 to 20 mm. long and blue or purple with lateral margins of gold, green, or red. Elytra are metallic green or blue green with green, gold, or red margins. Undersurface has varied metallic colors. Abdomen is green and red. Male is distinguished by MAP 98.—Recorded distribution of a dense brush of hairs on the in- Elaphrus ruscarius Say. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 155 G/Y: Not given. SA: Eggs. N/H: Not given. MOA: Larvae en- counter eggs. AF and HAB : Not given. {26^, p. Sl^) Description.—Eggs, larvae, and pupae unknown. Adults are 8 mm. or more long and dark brownish green, which is paler beneath and polished. Punctures are very crowded and numerous. Anten- nae and labrum are black and the palpi above are piceous but slightly paler beneath. Thorax is MAP 99.—Recorded distribution of broadest just anterior to the Harpalus caliginosus (Fabricius). middle but narrower than the elytra. There is a transversely in- 123, p. 490; 216, p. 135; 217, p. dented curved line anterior to the 31U) middle and a longitudinal abbre- Incidence of prédation.—Not viated one posteriorly. Elytra are given. equally punctured with about 20 Life history and habits.—SO: large dilated spots and two to Eggs, larvae, and pupae unknown. three equidistant, elevated, and Adults, not given. LOD: Un- elongated bronzed spots on each known. OWS and G/Y : Not given. side near the suture ; the one most SA : Eggs of grasshoppers. N/H : anterior is the largest. The post- Unknown. MOA: Larvae and pectus is green and tinged on each adults attack eggs. AF : Also seeds side with a cupreous color. Feet of ragweed. HAB: Not given. are green. Femora and tibiae are (159, p. 50; 216, p. 135; 252, p. piceous at the base. The venter is U6; 26U, p. 31U; 323, p. 231) green and the disk impunctured. Description.—Eggs, larvae, and Segments are brassy at the tip. pupae unknown. Adults are 18 to (292, p. U17) 26 mm. long and are about the largest species of this genus in Harpalus caliginosas (Fabricius) the United States. The body is (Carabus) 1798: 57 elongate to oblong and basically Synonymy.—Carabus politus black. Elytra are deeply striated, Melsheimer, C. caliginosus Fabri- and the thorax contains numerous cius {195, p. 70). minute punctures. Antennae and Recorded distribution.—South- palpi are reddish brown. Head is ern Canada and the United States smooth with two slightly im- {86, pp. 116-117). (Map 99.) pressed points anteriorly and Host species.—Grasshoppers there is an indented puncture on and cutworms (86, pp. 116-117; each side of the front. Labrum is 156 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE slightly emarginate at the tip and unarmed in the sinus. Thorax is %^^^^»SSNV broad and smooth with minute punctures. Legs are black; an- terior tarsi and midtarsi are blackish red. There are two rows of small scales beneath on the front tarsi. Midcoxal cavities are entirely enclosed by the sterna alone and the epimeron does not touch the coxae on the outer side. The first four segments of the hindtarsi decrease evenly and MAP 100.—Recorded distribution of Harpalus desertus LeConte. rapidly in length; the first seg- ment is longer than the fifth. (9, p. 140; 86, pp. 116-117; 103, p. Description.—Eggs, larvae, and 572; 159, p. 50; 193, p. 454) pupae unknown. Adults are 6 to Discussion.—Information is in- 7 mm. long, oblong, and shining sufficient to evaluate. This species black. Antennae, palpi, and feet is commonly attracted to light. are rusty brown. Elytra toward When irritated, it gives off a very the apex are obtuse and rounded ; pungent odor. (159, p. 50) they are covered with deeply im- pressed striae. Both the thorax Harpalus desertus LeConte and elytra are impunctured. Tho- 1859: 3 rax is narrow with the sides Synonymy.—(Genus ?) ochro- rounded normally; the anterior pus Kirby {195, p. 71). angles are blunt and also rounded Recorded distribution.—Wy- toward the base. {189, p. 3) oming, Colorado, and New Mexico Discussion.—Lavigne and {183, p. 7; 189, p. 3; 195, p. 71). Pfadt {183, p. 7) noted that this (Map 100.) species was rather abundant Host species.—"Grasshoppers" where grasshoppers were present {183, p.7). and would eat grasshopper eggs Incidence of prédation.—Not fed to it in the laboratory. It was given. not directly observed feeding on Life history and habits.—SO: grasshopper eggs in the field. Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. Harpalus f rater nus LeConte OWS and G/Y: Not given. SA: 1865: 103 Eggs. N/H: Unknown. MOA: Synonymy.—Harpalus occiden- Larvae encounter and feed on talis Chaudoir {195, p. 71). eggs. AF and HAB: Not given. Recorded distribution.—Brit- {183, p. 7) ish Columbia south through Wy- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 157 oming {195, p. 71; 336, p. 35). hopper egg pods. Lavigne and (Map 101.) Pfadt (183, p. 7) found that Host species.—"Grasshoppers" beetles in captivity fed readily on (183, p. 7; 331, p. 21 ; 336, p. 35). grasshopper eggs. Natural inci- Incidence of prédation.—Not dence data were not located. given. Life history and habits.—SO: Harpalus pennsylvanicus Not given. LOD : Unknown. OWS (DeGeer) and G/Y: Not given. SA: Eggs. 1752: 108 N/H: Unknown. MOA: En- Synonymy.—Harpalîis bicolor counter eggs as they search for Fabricius, H. fannus DeJean (195, food. AF and HAB : Not given. p. 70). Description.—Eggs, larvae, and Recorded distribution.—Most pupae not given. Adults : LeConte of Canada and the United States (188) described this species as (86, p. 118; 159, p. 50). (Map being closely allied to H. funestus 102.) LeConte and H. oblitus LeConte. Host species—Grasshoppers, The main differences are the tho- various insects, and caterpillars rax near the basal foveae is feebly (264., p. 314; 323, p. 231). and sparsely punctured, the elytra Incidence of prédation.—Not are more than 2i/^ times longer given. than the thorax and nearly equally Life history and habits.—SO: shining in both sexes, and the Eggs, larvae, and pupae unknown. sutural spine of the female is Adults not given. LOD : Unknown. distinct. (190, p. 103) OWS and G/Y: Not given. SA: Discussion.—Treherne and Nymphs. N/H: Unknown. MOA: Buckell (336, p. 35) suggested Adults. AF and HAB : Not given. that it is predaceous on grass- Description.—Eggs, larvae, and

MAP 101.—Recorded distribution of MAP 102.—Recorded distribution of Harpalus fratemus LeConte. Harpalus pennsylvanicus (DeGeer). 158 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE pupae unknown. Adults are 13 to 16 mm. long and black to reddish brown. Undersurface is also red- dish brown. The antennae and legs are pale yellow. Shape is elongate-oblong, robust, and con- vex. Striae of the elytra are mod- erately deep. Head is moderately large and the eyes are moderate. Prothorax is rounded at the sides and more converging anteriorly; the apex is much narrower than MAP 103.—Recorded distribution of the base. Head is a dusky reddish Harpalus somnulentis DeJean. brown and the labrum slightly darker. Head contains but one Host species.—Possibly grass- bristle-bearing puncture above the hoppers (336, p. 35). eye. Punctures may contain one or Incidence of prédation.—Not more bristles each. Mandibles are located. black at the tip. Thorax is also Life history and habits.—SO: reddish brown and as broad as Eggs, larvae, and pupae unknown. the elytra. Elytra are deeply stri- Adults not located. LOD: Un- ated and convex. Middle coxal known. MOA : Not given. AF : Not cavities are entirely enclosed by located. HAB : Not given. the sterna alone, and the epi- Description.—Eggs, larvae, and meron does not touch the coxa on pupae unknown. Adults' descrip- its outer side. There are two rows tion is unavailable. of small scales on the front tarsi Discussion.—Treherne and beneath, and the front tibia is Buckell (336, p. 35) reported this normal. {81, p. 108; 86, p. 118; species as a possible predator of 159, p. 50) grasshopper egg pods. Discussion.—This species is attracted toward lights. Not Pasimachus californiens Chaudoir enough information is available for an accurate evaluation as a 1850: 437 grasshopper predator. Synonymy.—(Genus ?) trans- versus Casey, (Genus ?) aco- Harpalus somnulentis DeJean manus Casey, (Genus ?) validus 1829: 333 LeConte, (Genus ?) corpulentus Synonymy.—None. LeConte, (Genus ?) cephatates Recorded distribution.— Casey, (Genus ?) obesus Casey Alaska, British Columbia, and (195, p. i7). California (W, p. 333; 336, p. 39). Recorded distribution.—Utah (Map 103.) Arizona, Colorado, New Mexico, ARTHROPOD AND NEMATODE PARASITES IN AMERICA 159 and Texas {195, p. ^7). (Map 104.) "!'^ ^^«fc5'x>^ Host species.—Grasshopper egg pods (264). Incidence of prédation Not given. Life history and habits.—SO and LOD: Unknown. OWS and G/Y: Not given. SA: Egg pods. N/H: Unknown. MOA and AF: Not given. HAB: Not located. (i8, p. 437) Description—Eggs, larvae, and MAP 105.—Recorded distribution of pupae unknown. Adults' descrip- Pasimachus elongatus LeConte. tion is unavailable. Discussion.—Only Riley's Host species.—"Grasshoppers" (264) prédation record was lo- (123, p. 490; 264, p. 314). cated. Incidence of prédation.—Not given. Pasimachus elongatus LeConte Life history and habits.—SO: 1848: 174 Eggs, larvae, and pupae unknown. Adults not given. LOD, OWS, Synonymy.—None (195, p. 47). G/Y, SA, and N/H: Unknown. Recorded distribution.—South- MOA and AF : Not given. HAB : central and southwestern Canada Dry open woodlands or prairies and north-central and eastern that contain sparse vegetation. United States (197; 264, p. 314). (Map 105.) (185, p. 174; 216, p. 136) Description.—Eggs, larvae, and pupae unknown. Adults are 28 mm. long and black. The thorax and elytra are margined with purple. Elytra are perfectly smooth and glabrous. Antennae are black and obscure at the tip. The head contains two impressed lines anteriorly and there is a longitudinal impressed line on the thorax. First joint of the antenna is black. Mandibles are as long as the head and strongly dentate in the middle ; tooth in the left one is MAP 104.—Recorded distribution of double. Outer margins of elytra Pasimachus californicus Chaudoir. have a line of elevated granules. 160 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Tarsi are blackish brown. (185, Description.—Eggs, larvae, and p. 174; 193, p. U9) (Fig. 15.) pupae unknown. Adults are 32 Discussion.—Not enough infor- mm. long, black, smooth, and mation is available for evaluation. shining. The marginal groove of the elytra is purple. Pairs of punc- Pasimachus punctatus Haldeman tures can usually be seen on the 1843: 298-299 elytra. The mandibles are deeply Synonymy.—(Genus ?) sinii- striated. The male has a rufous atus Casey (195, p. 47). brush on the posterior tibiae. Recorded distribution.—Cen- (127, pp. 298-299 ; 159, p. 28 ; 197, tral and southern United States p. 181) (159, p. 28). (Map 106.) Discussion.—The only préda- Host species.—Grasshopper tion records located was that of egg pods (264, V- 314). Riley (264, p. 314). Incidence of prédation.—Not given. Pterostichus occidentalis Life history and habits.—SO: (DeJean) Eggs, larvae, and pupae unknown. 1829: 231 Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA. Synonymy.—Platysma occiden- Eggs. N/H : Unknown. MOA, AF, tale DeJean (131, p. 115). and HAB : Not given. Recorded distribution.—Brit-

PN-2733 FIGURE 15.—Adult of Pasimachus elongatus LeConte. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 161 ish Columbia, Oregon, and Cali- fornia (131, p. 115). (Map 107.) Host species.—Grasshopper egg pods {123, p. A91 ; 237, p. 27). Incidence of prédation.—Not located. Life history and habits.—SO and LOD: Unknown. OWS and G/Y: Not given. SA: Eggs. N/H: Unknown. MOA and AF: Not given. HAB : Not located. (123, p. 491 ; 237, p. 27) Description.—Eggs, larvae, and MAP 107.—Recorded distribution of pupae unknown. Adults are 9 to Pterostichus occidentalis (DeJean). 12 mm. long and black. The pro- notum is more than three-fourths Buckell, but these could not be as long as wide and the entire located. basal region is ruggedly punc- tured. The elji;ra are alutaceous Selenophorus pedicularias or skinlike, especially in the fe- DeJean male. Antennae are pale for the 1829: 100 first three joints. The elytra are Synonymy.—Selenophorus striated and scarcely punctured. aereus LeConte, S. puellus Putz (84, p. 231 ; 131, p. 115) (193, p. 71). Discussion.—Greathead (123, Recorded distribution.—Wy- pp. 342, 491) referred to préda- oming, Missouri, Indiana, Vir- tion records of Treherne and ginia, Louisiana, and Florida (183, p. 7). (Map 108.) Host species.—Grasshoppers (183, p. 7). Incidence of prédation.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Eggs. N/H: Unknown. MOA: Tested only in the laboratory. AF and HAB : Not given. (183, p. 7) Description.—Eggs, larvae, and pupae unknown. Adults are 5 to MAP 106.—Recorded distribution of 6 mm. long, ovate, and dark Pasimachus punctatus Haldeman. bronze. The thorax is squarish. 162 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Members of this family gen- erally inhabit such exposed areas as paths, roadways, or sandy flats. Shortly after mating, the females excavate small cavities in the soil, usually less than one-half inch in depth, and singly deposit small translucent, ovate eggs. Young first-instar larvae have a large heavily sclerotized head and pro- notum. Their mandibles are ex- ceedingly large, powerful, and MAP 108.—Recorded distribution of bent slightly upward. The legs Selenophorus pedicularius DeJean. have long sharp claws. On the dorsum of the fifth abdominal The sides of the posterior part segment is a pair of f orwardly di- are poorly impressed and the rected hooks. The body is gen- posterior angles rounded. The erally S-shaped and adapted to elytra are short and rounded. The existence in a vertical burrow. bases of the antennae are testa- These burrows are usually con- ceous. Integument is punctured. structed in sandy soil and of a (8i, p. 100) diameter that allows the S-shaped Discussion.—Lavigne and larva to rigidly hold its position Pfadt (183, p. 7) reported that when attempting to subdue prey. these insects were abundant Winter is usually passed in the where there were grasshoppers larval stage. Just prior to pupa- and that they fed readily on grass- tion the larva seals the entrance hopper eggs in the laboratory. and forms a special pupation chamber either at the bottom of Cicindelidae the burrow or to one side. The diet of the adult is consid- The tiger beetles are aggressive erably more varied than that of predators both as larvae and as the larva since the adult moves adults. The larvae generally have more freely and thus has a wider a large sclerotized head with choice of prey. The rapid running strong mandibles and a pair of ability of the adult combined with hooks on the dorsum of the fifth its strength and sharp mandibles abdominal segment. The adults makes most insects subject to are characterized by their bril- attack. In addition to this method liant coloring, unique markings, of attack, some individuals wait swift running ability, and long, in ambush. The attack is aggres- slender, tapering mandibles (fig. sive and quick, and the prey is 16). usually torn apart. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 163

PN-2734 FIGURE 16.—Adult of Cicindela sp.

Tiger beetles do not limit their spp., plus a large number of insect diet to any particular group of species listed for both the adult insects and are able to utilize and larval stages (123, p. ^91 ; nearly anything that is available. 264, pp. 315-317-,330, p. 177; 331, This characteristic may detract V. 25). from their usefulness as grass- Incidence of prédation.—Not hopper predators. given. Life history and habits.—SO: Amblycheila cylindriformis Eggs, larvae, and pupae unknown. (Say) Adults not given. LOD : Unknown. (Manticora) 1823: 139 OWS, G/Y, and SA: Not given. Synonymy.—Manticora cylin- N/H: Unknown. MOA: The lar- driformis Say, Amblycheila pico- vae trap their prey, dragging it lominii Reiche {193, pp. 89, 521). down into their holes where it is Recorded distribution.—Mon- usually devoured. AF: Insects. tana, Wyoming, Utah, Colorado, HAB : Adults usually remain hid- Kansas, New Mexico, Texas, Okla- ing below ground unless the soil homa, and Arkansas (128, p. 70; becomes extremely dry. Larvae 159, p. 16). (Map 109.) can usually be found in burrows Host species.—Melanoplus very close together, often not spretus (Walsh), Brachystola more than II/2 inches apart. Thus 164 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE ous tubercles, each bearing a hair. Antennae are situated imme- diately behind the base of the mandibles and have four seg- ments, the second of which is nearly twice as long as the first. Arising from their upper surface are moderately long hairs. Man- dibles are long, slender, and mod- erately curved with a triangular tooth slightly behind the middle. The prothorax is slightly broader MAP 109.—Recorded distribution of than the head, nearly semicir- Amblycheila cylindriformis (Say). cular, and broadly emarginated anteriorly. The mesothorax is a colony is usually in a very small much narrower than the protho- area. The burrows are generally rax, trapezoidal, and narrow pos- on the brow of a cliff, whereas teriorly. The metathorax is as others are on plains away from broad at the apex as the base of the bluff. (94, pp. 531-532 ; 128, the mesothorax and gradually pp. 70-71 ; 159, p. 169; 163, p. 253 ; broadens posteriorly. (lU, pp. 264,pp. 315-317; 3U, p. 7) 29-31; 3U, p. 7) Pupae un- Description.—Eggs, unknown. known. Larvae before maturity are about Adults are slightly longer than 12.5 mm. long and 6 to 7.5 mm. 1 inch and dark chestnut brown to wide. Generally they are yellow- blackish with pale-brown elytra. ish. The head is white and the The head is large and the eyes prothoracic suture chestnut are small. The mandibles are very brown. They are cylindrical and strongly toothed. The thorax is slightly flattened. The head and narrow behind but not elevated prothorax are bent at a right and darker than the elytra, which angle to the body. The tip of the are irregularly punctured. Many abdomen beyond the fifth abdom- of the punctures are preceded by inal segment is gradually arcuate. a slightly elevated point. (,159, Viewed laterally the body is p. 16; 193, pp. 89, 521; 285, p. nearly of equal thickness except 139) the swelling of the fifth abdominal Discussion.—This species is segment. The upper surface of the predatory on many species if in- head is broader than it is long sects. Although a few of them are and the front margin is trilobed. rejected, none seem to be pre- Surrounding the eyes are numer- ferred. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 165

Cicindela circumpicta spots are sharply oblique. The LaFerté-Sénectère thorax is roundish and shining. 1841: 39 The elytra are punctured. There Synonymy.—Cicindela collaris are teeth entirely on the inner LaFerté-Sénecterè, C. johnsoni anterior margin of the elytra. Fitch (195, p. A2). (159, p. 23; 175, pp. 39-kO; 350, Recorded distribution.—South- pp. 51-53) ern Wyoming and southern South Discussion.—Evidence is insuf- Dakota south through Arizona ficient for an evaluation. and Texas {195; p. k2\ 26k, p. Cicindela formosa Say 31k; 350, pp. 51-53). (Map 110.) 1817: 19 Host species.—Melanoplus spretus (Walsh) (262, p. 31k). Synonymy.—Cicindela luxurî- Incidence of prédation.—Not osa Casey, C. manitoha Leng, C. given. generosa DeJean, C. latecincta Le- Life history and habits.—SO: Conte (195, p. kO). Eggs, larvae, and pupae unknown. Recorded distribution.—South- Adults not given. LOD. Unknovs^n. central and southeastern Canada OWS, G/Y, SA, N/H, MOA, and and the United States (195, p. kO ; AF : Not given. HAB : Unknown. 350, pp. 15, 36-39). (Map 111.) Description.—Eggs, larvae, and Host species.—Melanoplus pupae unknovni. Adults are 4 to spretus (Walsh) (262, p. 31k). 5 mm. long, subcylindrical, and Incidence of prédation.—Not greenish bronze. Apex of the lún- given. ula is rounded and the median Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. yk OWS and G/Y: Not given. SA: Nymphs, maybe adults. N/H and MOA: Not given. AF: Insects. ^a^^-^Ml HAB : Near rivers, bare spots in sandy areas, and sparse vegeta- .'.írr¿— tion; in blown areas. Hamilton '^¿m (128, pp. 18, k6-k7) and Wallis I (350, pp. 15, 36-39) reported that larvae of C. formosa were col- lected in sand dunes along various KC rivers where the sand was coarse and finely mixed. Burrows have MAP 110.—Recorded distribution of Cicindela circumpicta LeFerté-Sén- vertical pits about 22 inches deep. ectère. Description.—Eggs, unknown. 166 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE hook. (128, pp. 18, Í6-Í7) Pu- pae unknown. Adults are 16 to 18 mm. long. They are a brilliant red cupreous color above with a three-branched broad white margin on the elytra. Underparts are metallic blue or green or purple blue, and the edge of the elytra is green. The femora are blue and the tibiae green. There is much hair on the side of the head, sides of the thorax, ab- MAP HI.—Recorded distribution of domen, trochanters, and femur, on Cicindela formosa Say. the front, and generally beneath. Labrum is large and two-toothed. Larvae (mature) are 24 to 26 mm. The thorax is broader than it is long. The head and pronotum are long. {159, p. 22; 192, p. 35; 282, chestnut brown with lighter areas p. 19) in the color pattern. Setae on the Discussion.—This is a very dorsal aspect are long, stout, and beautiful species and resembles transparent to white. Other setae somewhat Cicindela hirticollis are brown. On the caudal part of Say. the frons is a U-shaped ridge with three setae. The proximal segment Cicindela fulgida Say of the antennae is slightly shorter 1823: 141 than the second with the third Synonymy.—Cicindela paro- one-half and the distal one-fourth wana Wickham, C. wallisi Calder the length of the second. The max- {195, p. Al). illary palpi are three-segmented. Recorded distribution.—South- On the pronotum are cephalolat- ern Canada and the United States eral angles extending almost to {159, p. 23; 195, Ul ; 285, p. HI ; the mesal part. The lateral mar- 350, pp. 36-39). (Map 112). gins are roughened. The primary Host species.—Grasshoppers setae are not large and secondary and many insect species. (26i, p. setae are absent. The chitinized 3U). areas are distinct on the abdomen. Incidence of prédation.—Not The secondary setae are about given. one-half the length of the primary Life history and habits.—SO: although not numerous. The me- Eggs, larvae, and pupae unknown. dian hooks contain two setae ; the Adults not given. LOD : Unknown. inner hooks have four setae, OWS and G/Y: Not given. SA: which are spinelike projections Nymphs and adults. N/H: Un- about one-third the length of the known. MOA: Adults ambush ARTHROPOD AND NEMATODE PARASITES IN AMERICA 167 prey. AF: Insects. HAB: Saline Cicindela pulchra Say or alkaline soils with sparse veg- 1823: 142 etation. It has been reported from both areas near mountains and Synonymy :—None. rivers. (350, pp. 36-39) Recorded distribution.—North Description.—Eggs, larvae, and Dakota southeast into Arkansas pupae unknown. Adults are less (128, pp. 18,51). (Map 113). than 13 mm. long and brilliant Host species.—Melanoplus red cupreous, which appears to be spretMs (Walsh) (193, p. 91 ; 26A, highly polished. The anterior area p. 3U). of the head is hairy. It varies from Incidence of prédation.—Not green to blue and on each side. given. The exterior base of the mandibles Life history and habits.—SO: and the labrum are white and the Eggs, larvae, and pupae unknown. antennae black at the tip. Blue Adults not given. LOD : Unknown. lines on the thorax are impressed, OWS and G/Y: Not given. SA: and dense punctures are on the Nymphs and adults. N/H: Un- elytra. On the elytra are two known. MOA: Adults pounce on lunules and an intermediate re- prey. AF : Insects. HAB : Areas fracted band behind the middle. bordering rivers and near moun- The dilated lunule at the tip is tains. (128, pp. 18, 51) white. The under part of the body Description.—Eggs unknown. is hairy and green. {159, p. 23; Larvae (mature) are 21 to 23 mm. 285, p. HI) long. The head and pronotum are Discussion.—Not enough infor- dark purple with dark-green re- mation is available for an accu- flections. Setae on the dorsum of rate evaluation. the head and pronotum are long, stout, and white, whereas other setae are brown. On the U-shaped ridge on the caudal part of the frons are three setae. The proxi- mal segment of the antennae is shorter than the second and the distal segment is one-third as broad as the second. Maxillary palpi are three-segmented. Pri- mary and secondary setae are small on the pronotum. The lat- eral angles of the pronotum ex- tend anteriorly as far as the f ' .-U^i-i mesal part of the lateral margins.

MAP 112.—Recorded distribution of Chitinized areas of the abdomen Cicindela fulgida Say. are distinct, although the second- 168 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Casey, C. prolixa Casei/, C. fonti- naria Casey. Recorded distribution.—South- ern Canada and the United States (86, pp. 51-53, 55, 58; 363, p. 152). (Map 114.) Host species.—Melanoplus spretus (Walsh) (26J^, p. ^31). Incidence of prédation.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. MAP 113.—Recorded distribution of Adults not given. LOD : Unknown. Cicindela pulchra Say. OWS and G/Y: Not given. SA: Nymphs and adults. N/H: Un- known. MOA: Adults pounce on ary setae are short and few. prey. AF: Insects. HAB: Not Median and inner hooks have two given. (26i, p. A31 ; 363, p. 152) setae. (128, pp. 18, 51; 285, p. Description.—Eggs, larvae, and H2) Pupae unknown. pupae unknown. Adults are 13 to Adults are about 18 mm. long 14 mm. long, obscure cupreous be- and greenish purpurescent. Elytra neath, and varied with blue, pur- are red cupreous and highly pol- ple, and white markings on the ished; the external margins are elytra. Head is cupreous, obscure, purple and have two white dots. with two blue lines between the Front is very hairy. The terminal eyes. Antenna are brown and the segments of the antennae are base is cupreous. Labrum is white black. Labrum is short. Mandibles with a single prominent tooth and bear teeth and are white; the ex- tremity and inner surfaces are black. There is a large double purple-cupreous spot on the disk of the thorax. Elytra have nu- merous large punctures, which are more profound toward the base. Feet are hairy. (193, p. 91 ; 285, p. U2) Cicindela punctulata Oliver 1790: 27 Synonymy.—Cicindela micans Fabricius, C. obscura Melsheimer, MAP 114.—Recorded distribution of C. tenisoni Gistel, C. boulderensis Cicindela punctulata Oliver. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 169 six submarginal punctures. Palpi (86, pp. 51-5Í; 350, pp. 14, i-3- and labials are white; the last Jf6). (Map 115.) joints are green. Thorax is the Host species.—Melanoplus same color as the head. It has spretus (Walsh) (123, p. 491; impressed lines and blue lateral 264, p. 354). margin. Elytra are the color of Incidence of prédation.—Not the thorax and irregularly punc- given. tured with green. There are five Life history and habits.—SO : small white dots on the elytra, of Eggs, larvae, and pupae unknown. which three are on the disk ar- Adult populations peak first in ranged in an oblique line, one near June, then August and Septem- the humérus, the second central, ber. LOD: Unknown. OWS and and the third near the suture. G/Y : Not given. SA : Nymphs and Fourth and fifth dots are marginal adults. N/H: Unknown. MOA: situated opposite the two preced- Adults pounce on prey. AF: In- ing ones. Venter of the abdomen sects. HAB : On meadow path- is varied with purple and blue and ways and in grass along road- the tail is a bronzed color. Le- ways. (86, pp. 51-54; 197, pp. Conte (193) reported that the 91-92) variations are confined to changes Description.—Eggs, larvae, and in the number and magnitude of pupae unknown. Adults are 14 to the spots on the elytra. The an- 16 mm. long. The upper surfaces terior marginal dot is always mi- are generally reddish cupreous nute, and all the others are fre- and the underside bluish green. quently lacking. (86, pp. 51-53, Sutures of the eljrtra are green, 55-58; 102, p. 2U; 193, p. A31; and the elytral markings consist 197, p. 93) of a short median band and a dot Discussion.—This is a common near the apex ; the ends are tipped species but it is not important as a predator of grasshoppers. Cicindela purpurea Oliver 1790: 14 Synonymy.—Cicindela margin- alis Fabricius, C ramosa Melshei- mer, C. supera Dahl, C. gramínea Schaupp, C. spreta LeConte, C. auduboni LeConte, C. auguralis Casey, C. pugetana Casey, C. lim- balis Casey, C. inducta Casey, C. nigerrima Casey. (195, p. ^0) Recorded distribution.—South- MAP 115.—Recorded distribution of ern Canada and the United States Cicindela purpurea Oliver. 170 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE with cream color. The head is red i!l-: ■ cupreous, hairy, and with green edges. Two distinct green lines between the eyes originate at the base of the antennae and proceed toward the vertex. Antennae are green at the base and brown on the tip. Clypeus is blue and the labrum is white. Labrum contains three teeth ; the edge is black with about eight marginal punctures. Mandibles are black within and at the tip. Palpi are green. Thorax MAP 116.—Recorded distribution of is granulate rugose and contains Cicindela repanda DeJean a cupreous disk. Elytra are oli- vaceous green to brilliant cupre- Host species.—Rocky Mountain ous red with a bright green mar- locust and a wide variety of in- gin. A reclivate band near the sects including flies and other middle originates at the green beetles (123, p. 419; 264, p. 314; margin and terminates some dis- 331, pp. 25, 31-32). tance from the suture. There is Incidence of prédation.—Not also a transverse line at the tip given. and the intermediate submarginal Life history and habits.—SO : dot is white. Trochanters are pur- Eggs are deposited in May and ple and tibiae are covered with June. Larvae occur from June to long strong hair behind. (86, pp. spring. Pupae are present in the 51-54-; 193, pp. 429, 523; 197, pp. spring. Adults emerge in May and 91-92; 230, p. H; 283, pp. 419- can be found throughout the year. 420; 350, pp. 14, 43-46; 363, pp. LOD : Eggs not given. Larvae, pu- 151-152). pae, and adults unknown. OWS: As third-instar larvae. G/Y : One Cicindela repanda DeJean generation takes 2 years. SA : Not 1825: 74 given. N/H: Unknown. MOA: Synonymy.—Cicindela hirticol- While hunting, adults of this tiger lis Gould, C. baltimorensis Le- beetle stand very high or when Conte, C. unijuncta Casey (195, ambushing will crouch in a de- p. 40). pression in the soil and await the Recorded distribution.—South- arrival of the prey. They attack, ern Canada and the United States puncture, and toss their prey east of the Rocky Mountains (86, around. The beetle then carries or pp. 51-54; 159, p. 22; 195, p. 93; drags its prey for a short distance 221, pp. 338-343; 328, pp. 16-20; before beginning to feed. Small 363, p. 152). (Map 116.) insects are chewed and swallowed, ARTHROPOD AND NEMATODE PARASITES IN AMERICA 171 whereas larger ones are broken imately 0.8 by 0.5 mm., short, oval, into pieces. AF: Insects. HAB: and pearly white to straw yellow. Wet sandy soil, wet muddy soil, The chorion is corrugated with a moist clay-type soil, and soil with triangular pattern and there is a considerable humus; also flat, ridge at one pole above which the sparsely vegetated areas in the egg is flattened. Eggs are gener- immediate area of travel, such as ally covered with adhesive fila- dirt roadways and paths through ments. (223, pp. 338-3Í3) grassy areas and margins sur- Larvae (mature) are 16 to 18 rounding small ponds, lakes, and mm. long and both the head and streams. pronotum are dark coppery bronze Eggs are laid in small chambers with a light-green reflection. All constructed in the soil about 8 to setae are brown except those on 10 cm. deep. They are generally at the dorsal aspect of the head and right angles to the sloping surface. pronotum, which are white. Setae First-instar larvae hatch shortly on the dorsal aspect of the head thereafter and develop within the are long, stout, and prominent. soil habitat to the third instar, There is a U-shaped ridge on the which overwinters. Adults appear caudal part of the frons between in the field the following May but two setae. The proximal and sec- do not become sexually mature un- ond segments of the antennae are til May of the second year. The subequal in length. Maxillary eggs are laid, the adult dies, and palpi are three-segmented and the the life cycle is complete. proximal segments of the labial Moore {221) noted that this palpi have three spinelike projec- species digs its mandibles into the tions on the ventral distal margin soil to obtain moisture, but when and two setae on each side of these left with dry sand and a water spines. The cephalolateral angles dish, it will die of apparent thirst. of the pronotum extend almost as When disturbed, these beetles far forward as the mesanotum; squat and remain motionless un- the lateral margins are keeled or til the intruder has moved. If the carinate. Chitinized areas of the intruder approaches within 4 to larvae are distinct. (128, p. 17; 6 inches, the springs lU, pp. 35-37) Pupae unknown. several inches into the air before Adults are 12 to 13 mm. long, it begins its flight. Although C. elongate-ovate, robust, subconvex, repanda is a strong flier, it very and brownish bronze with a cop- seldom travels long distances un- pery or strong greenish reflection. less provoked to do so. (86, pp, The under parts are usually thinly 51-5Í; 128, p. 17; 159, p. 22\ 228, covered with long white hairs and pp. 338-3JfS', 252, p. 37; 350, pp. the markings, dorsally are con- 16-20; 363, p, 152) nected and enter the interior of Description.—Eggs are approx- the wing cover. The humeral 172 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE lunule is C-shaped; the middle unicolor DeJean, C. nigrior band rectangularly bent and con- Schaupp, C. modesta DeJean, C. - nected with the marginal line obscura Say (195, p. Ji.1). . nearly reaches the lumulus. The Recorded distribution.—South- pronotum is subquadrate and dis- ern Canada and the United States "^ tinctly narrowed posteriorly. The (159, p. 21; 350, pp. H, 33-36). , disk is finely and transversely ru- (Map 117.) gose. Outer margin of the elytra Host species.—Grasshoppers ' of the female is not angulate. The (26A,p.3H). thorax is cylindrical with no mar- Incidence of prédation.—Not gin. Both male and female have given. flying wings and the elytra are Life history and habits.—SO: slightly flattened. Eyes are large. Eggs, larvae, and pupae unknown. The third coxae touch each other. Adults not given. LOD : Unknown. (83, p. 70; 86, pp. 51-5Í; 159, p. OWS, G/Y, SA, Not given. N/H : ' 22; 197, p. 93; 283, pp. A01-A26; Unknown. MOA : Not given. AF : 363, p.152) Insects. HAB : Sandy blownout Discussion.—This insect seems areas where the sand is loose and -' to be limited in vision. Quick vegetation is sparse. Its habitat movements or change in light in- is generally that associated with tensities 6 to 8 feet away will C. formosa. Although its habitat ' cause the beetle to take defensive extends deep into Canada, it is . action, but apparently it is only more common in the southern re- able to discern insects and other gions. (350, pp. lU, 33-36) objects correctly within 4 to 6 Description.—Eggs, larvae, and . inches. It makes up for this, how- pupae unknown. Adults may vary ever, in its quick running ability widely in color. Therefore they - and decisive method of attack. Al- have been divided into several . though this beetle will attack grasshoppers, and is abundant over a large geographical area, it is doubtful that it has much ef- fect on reducing populations. Cicindela scutellaris Say 1823: 140 Synonymy.—Cicindela lecontei Haldeman, C. criddlei Casey, C. rugifrons DeJean, C. denticulata Harrington, C. oberleitneri Gistel, C. varians Harrington, C. mutans Harrington, C. variata Harring- MAP 117.—Recorded distribution of ton, C. Carolina E. D. Harris, C. Cicindela scutellaris Say. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 173 varieties, including C. unicolor DeJean, C. modesta DeJean, C. nigrior Schaupp, and C. rugifrons De Jean. They are 11 to 13 mm. long and generally green or some- times blue on the head and tho- rax. The elytra are coppery or greenish with distinct or con- nected marginal dots. Antennae are black at the tips, whereas the labrum and mandibles above the base are white. The elytra are abruptly rounded at the tip and MAP 118.—Recorded distribution of Cicindela sexguttata Fabricius. punctured. The brilliant reddish- brassy region of the scutellum ex- tends from the middle of the base Incidence of prédation.—Not to beyond the middle of the suture given. and beneath is generally blue with Life history and habits.—SO: a violet tinge. Wallis {350) stated Eggs, larvae, and pupae unknown. that in the male the labrum is Adults not given. LOD : Unknovsm. white and the front hairy, OWS, G/Y, and SA: Not given. whereas in the female the labrum N/H: Unknown. MOA: Adults. is black and there are few hairs AF: Insects. HAB: Shady paths in clusters near the front inner in forests or pastures, on summits edge of each eye. {159, p. 21 ; of rocky knolls, in conifers, on 193, pp. 89-90; 285, p. 1^0; 350, green moss, sand, or gravel, and pp. H, 33-36) on any seemingly barren area. {350, pp. 13, 30-33) Description.—Eggs, larvae, and Cicindela sexguttata Fabricius pupae unknown. Adults are 13 to 1775: 241 15 mm. long and polished greenish Synonymy.—Cicindela guttata blue. Three marginal white dots Emmons, C. thalassima DeJean, are on each elytra. The four basal C. varians Ljungh, C. qvndrigut- joints of the antennae are green and the remainder is blackish tata Davis {195, p. Jfl). brown. Labrum is white edged Recorded distribution.—Mani- with brown. Whereas the man- toba, Ontario, Quebec, and United dibles are white above and the States {86, p. 52; 159, p. 20; 195, tips black. Palpi are green. Eyes p. Al). (Map 118.) are brown. The forepart of the Host species.—Melanoplus body is green tinged with blue and spretus (Walsh) {26i, p. 314.; cupreous. Hairs are few and 331, pp. 23-25). short. The venter of the abdomen 174 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE is bluish green and the segments are bronzed and edged in purple. The hindmargin of the green ely- tra is rounded and obscurely ser- rated. Each elytron is marked with three dots ; the first is in the middle of the margin, the second at the posterior curve, and the third is transverse and terminal. (86, p. 52; 102, p. 2U0 ; 128, pp. 17, 28; 159, p. 20; 193, pp. Í25-426, 523; 197, p. 88; 283, p. Ui; 350, pp. 13, 30-33; 363, p. 151) MAP 119.—Recorded distribution of Cicindela splendida Hentz. Cicindela splendida Hentz 1830: 254 green. The elji;ra are densely and Synonymy.—Cicindela discus finely roughened, whereas the Klug, C. transversa Leng, C. den- pronotum is distinctly narrowed verensis Casey, C. gramínea posteriorly and the surface Casey, C. conquisita Casey, C. densely and transversely oreada Casey, C. ludoviciana Leng wrinkled. Margins of the elytra (195, p. iO). are green and the remainder is Recorded distribution.—United a bright metallic bronze. There States (86, pp. 5Jt, 56-57; 195, p. are three distinct whitish marks, W). (Map 119.) one at the humérus, one at the Host species.—Grasshoppers basal fourth, and the third at (26^, p 315). the apex. There are also two Incidence of prédation.—Not bands on the elji;ra; the broad given. median band is recurved and di- Life history and habits.—SO: rected obliquely posterior toward Eggs, larvae, and pupae unknown the suture, and the second band Adults not given. LOD : Unknown. is curved toward the apex. (86, OWS; G/Y, and SA: Not given. pp. 5U, 56-57; 131^, p. 25k; 197, N/H: Unknown. MOA: Adults p. 92; 363, p. 151) and larvae search for prey. AF: Insects. HAB : Barren dry clay or Cicindela tranquebarica Herbst sand. Adults swarm in April in 1806: 178 North Carolina. (13J^, p. 25A) Synonymy.—Cicindela vulgaris Description.—Eggs, larvae, and Say, C. obliquata DeJean, C. tur- pupae unknown. Adults are 12 to bulenta Casey, C. wichitana Casey 14 mm. long, elongate-ovate, sub- (195, p. JfO). convex, and rather robust. They Recorded distribution.—South- are bright metallic green to bluish ern Canada and the United ARTHROPOD AND NEMATODE PARASITES IN AMERICA 175 states {36S, p. 152). (Map merus, which is sometimes inter- 120). rupted on the margin. The abdom- Host species.—Melanoplus inal tergites are greenish blue. spretus (Walsh) {26i, p. 3H). The venter is blue with purple Incidence of prédation.—Not shading. (135, p. 178; 193, pp. given. U22, 522 ; 197, pp. 192-93 ; 283, pp. Life history and habits.—SO U09-U10\ 363, p. 152) and LOD : Unknown. OWS, G/Y, and SA: Not given. N/H: Un- knovs^n. MOA: Adults search and Cantharidae ambush. AF: Insects. HAB: Not Adult soldier beetles are elon- given. {26Jf,p. 3H;363,p.l52) gate and soft bodied. They often Description.—Eggs, larvae, and frequent flowers. Many species pupae unknown. Adults are 16 to are predaceous and the others 18 mm. long. The head is black- phytophagous. Larvae are gen- ish to obscure cupreous ; the base erally predaceous on other insects. above is green and the front is One species, Cantharis bilineatus covered with cinereous hair. (Say), is predaceous on grass- ^ Palpi are green as are the first hopper egg pods. Several addi- four joints of the antennae; re- tional species also have been re- maining antennal joints are ported as predaceous on the egg ■ black. Mandibles are white at the pods of grasshoppers, but their base. Feet are green and the identification is apparently in er- thigh is a brassy red above. The ror since the species in question do elytra are cupreous brown or not inhabit North America. ^ obscure black with minute punc- Eggs are deposited in the soil in tures. An external lunule on each mass, generally from 100 to 200. "~ elytron originates on the hu- They hatch in 1 to 2 weeks, and the larvae develop and pupate in the soil. They usually overwinter E^'-^vy as larvae in the soil, and there are one to two generations per year. Cantharis bilineatus (Say) (Telephorus) 1823: 182 Synonymy.—Telephorus biline- atus Say {195, p. m). Recorded distribution.—East- ern United States (86, pp. 259, 26A). (Map 121.) Host species.—"The Rocky p MAP 120.—Recorded distribution of Mountain Locust" {26JÍ, pp. 302- Cicindela tranquebarica Herbst. 303). 176 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

segments are four times as long as the first. Labrum is not visible, but the mandibles are furcate with a strong tooth slightly be- yond the middle of the inner edge. Maxillary palpi are three-jointed; the second is twice as long as the , first. Labial palpi are two-jointed ; the second is rudimentary. Two large sunken suboval brown spots I À^^-M with pale centers are on each side of the dorsum of segments 2 and MAP 121.—Recorded distribution of 3 of the thorax. They are con- Cantharis hilineatus (Say). tinued on the abdomen less dis- tinctly as a slightly sunken pale Incidence of prédation.—Not line with still another line parallel given. with it on the sides. Between these Life history and habits.—SO: pale lines on all segments but the Eggs, larvae, and pupae unknown. last can be seen a dark beadlike Adults not given. LOD : Unknown. tubercle resembling spiracles. OWS and G/Y: Not given. SA: Anal joint bears a proleg. Legs Egg pods. N/H : Unknown. MOA : are rather short, pilose, and ter- Larvae encounter food. AF and minate in a sharp claw. (26Ji., HAB : Not given. {26A, pp. 302- pp. 302, 303; 3U, p. 19) 303; 323, p. 238) Pupae unknown. Adults are 6 Description.—Eggs unknown. to 8 mm. long and dull reddish Larva body is 12-segmented ; the yellow. The elytra, back of the first segment is the longest, the head, and spots on the thorax, fourth is the shortest, and the antennae except the basal joint, remainder are of about equal and palpi are black. There are two length. The body is flattened and parallel, abbreviated, dilated black tapers slightly toward each end. lines on the thorax and the exte- It is a deep rich brown above and rior basal margin is pale. The pale below. The head is flattened thorax is widely margined and and divided by a transverse sparsely punctured. Tibia and suture. The basal part is velvety tarsus are black. (86, pp. 259, sooty black and the anterior end 26^; 159, p. 130; 193, p. 118; 285, is polished and forms three well- p.182) denned lobes. Eyes are prominent and located laterally immediately Meloidae anterior to the occipital suture. Antennae are two-jointed and in- Blister beetles are well known serted in deep sockets ; the second both for their medicinal proper- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 177 ties and for their destructiveness Blister beetles undergo hyper- to commercial crops. In medicine, metamorphosis, which is slightly the powder obtained from pulver- different from that common to izing the dried body of the adult other beetles. This includes the beetle is made into a paste, which usual egg, larval, pupal, and adult produces blisters when coming in stages, but development is ex- contact with the human skin. Al- tended by the addition of two though the species best known in nonf eeding larval stages, one rest- this group, the Spanishfly (Lytta ing and the other ambulatory. vesicatoria (L.)) is European, the Adults that are predaceous on American species also possess the grasshopper eggs first appear in same blistering quality. Some the field from April to June and products of the beetles are taken copulate within hours of emer- internally as a stimulant or di- gence, generally on the food plant uretic and in the past were also or on the soil. Shortly thereafter used to make love potions. the females construct burrows in Adult beetles are considered the soil where oviposition occurs. important pests of both forage As the beetle leaves the oviposi- and garden crops. The foliage of tion chamber, the entrance is potatoes and tomatoes is pre- closed with bits of soil that are ferred. Certain weeds are at- packed with the abdomen. tacked as well as some ornamen- One to two hundred eggs are tals. The adults are gregarious. deposited in sticky masses within They attack gardens and crops in the earthen chambers. Several of large or small swarms, which these egg masses may be de- concentrate their feeding in a posited, each in its own individual small area. From time to time earthen chamber over a period of some or all of the group leave a days or weeks. After a short in- particular feeding area and travel cubation period, first-instar thy- a considerable distance to another sanuriform larvae emerge and location. begin seeking grasshopper or The larvae do not feed on plants other insect eggs. The second but are predaceous. The 26 spe- instar is caraboid but still am- cies included here have been re- bulatory. The third and fourth ported in the literature as attack- instars are scarabaeoid in form ing grasshopper eggs and have and still mobile although much been considered as control agents. more plump. It is known that cricket eggs are After completing the feeding sometimes included in the diet of stages, the larva generally bur- the various species, but the full rows downward 1 or 2 inches into range of food material is not the soil where it constructs a known and it is assumed that chamber about twice its length other eggs may also be eaten. and molts into the sixth-instar 178 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(coarctate) stage. In some species equal to that which would have this is considered the fifth instar. been consumed by the grasshopers The coarctate larva is a resting, they destroyed. Therefore blister nonfeeding form, which is highly beetles are of questionable value resistant to desiccation and tem- as biological control agents. perature change and is often the overwintering stage. At its com- Epicauta alhidae (Say) pletion, the larva molts to the seventh instar, burrows upward {Lytta) 1823: 305 to within an inch or two of the Synonymy.—Lytta albida Say, soil surface, constructs a second Macrobans albida (Say), L. lutei- chamber similar to that of the cornis LeConte (195, p, 158). first, and pupates. Common name.—It has been In some species and under ideal called the two-spotted blister conditions the sixth and seventh beetle (195, p. 158). larval stages may be bypassed. If Recorded distribution.—Colo- so, the second generation in that rado, New Mexico, Kansas, Texas, year is attempted although often Oklahoma, Arkansas, and Louis- eggs are not laid or development iana (85, pp. UOS-UOU ; 159, p.l50\ is not completed before winter. 195, p. 158). (Map 122.) With Epicauta lemniscata (Fab- Host species.—Grasshopper ricius), there may be three gen- egg pods and soft-bodied insects erations a year in Louisiana. (123, p. Í92; 236, p. 26). Other species require 2 years to Incidence of prédation.—Not complete a generation. However, given. one generation a year is generally Life history and habits.—SO: the rule for those predaceous on Eggs, larvae, and pupae unknown. grasshoppers. Adults not given. LOD : Unknown. Primary larvae are abundant in OWS and G/Y: Not given. SA: the field from July to September Egg pods of grasshoppers and when grasshoppers are oviposit- larvae of other insects. N/H : Un- ing. Upon locating a grasshopper known. MOA: Larvae encounter egg pod, the larva enters and be- egg pods when searching for food. gins feeding. Usually one egg pod AF: Range grasses, sugarbeets, is sufficient to rear one larva. and other garden crops. HAB: Larvae are antagonistic, and if Range and gardens. (160, p. two are feeding in the same egg 181 ; 222, p. 5) pod, either one or both are killed. Description.—Eggs unknown. The beneficial aspect of the Larvae (immature) are about 3.3 larva is offset by the destructive mm. long and yellowish. Head and habit of the adult. One wonders if thorax are yellow brown, and ab- the beetles do not consume a dominal segments 7 to 9, along quantity of vegetation at least with the lateral margin of the ARTHROPOD AND NEMATODE PARASITES IN AMERICA 179

They are black to gray or yellow- ish, and the body is densely cov- ered with pale hair. The entire upper surface is rather finely and densely punctured. Front of the head, mouth parts, antennae, and palpi are often pale yellow brown to orange brown. There are hu- meral spots and two nearly par- allel lines on the thorax. Antennae are of moderate length, extend back to about the basal fourth of MAP 122.—Recorded distribution of the elytra, and are filiform; the Epicauta albidae (Say). first segment is much enlarged. At least six segments of the an- sixth segment, are brown. The tennae are darkened. Legs, except body surface is covered with fine the tarsi, are slightly orange lines. The head is longer than the brown. Tarsal claws are cleft to prothorax and mesothorax com- the base. Protibia and mesotibia bined, and the lateral margins are have the outer spurs shorter than slightly emarginate. Antennae are the inner. Wings are usually moderately robust; the second present. {85, pp. UOS-UOU; 1U2, segments are more than twice as pp. 89-90; 159, p. 150; 192, p. 6; long as the third. At the apex of 193; p. 170; 285, p. 305) (Fig. the second segment are one short 17.) and two long setae. Mandibles are Discussion.—There is no evi- moderately robust and bear 13 dence of larval food preference. teeth, nine of which can be seen Adults are sometimes a serious in outline. The thorax is about pest on sugarbeets and other gar- one-fourth as long as the body den crops. and the prothorax about as long as the mesothorax and the meta- Epicauta callosa LeConte thorax combined. Ten setae in 1866: 158 marginal rows are located on ab- Synonymy.—None (193, p. dominal tergites 1 to 8, and spine- 156). like evaginations are on tergites Common name.—It has been 1 to 5. There are seven lanceolate called the sunflower blister beetle. setae on the anterior femora. Recorded distribution.—West- (21-i, pp. Jf5, 55-56) Pupae un- ern United States except region known. west of the Rocky Mountains (5^, Adults are between 14 to 20 pp. 38Í-385; 193, p. 156; 220, p. mm. long; one description indi- 6; 339, pp. 16,29). (Map 123.) cates an upper limit of 28 mm. Host species.—Melanoplus 180 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

'X Vl

PN-273B i FIGURE 17.—Adult of Epicauta albidae (Say). sanguinipes (Fabricius) and sev- Adults not given. LOD : Unknown. eral other rangeland grasshoppers OWS and G/Y: Not given. SA: ■ (117, p. 20; 172, p. U92; 212, pp. Egg pods. N/H : Unknown. MOA : 61-62 ; 23Jt, p. 26 ; 355, pp. A3-U). Larvae encounter eggs while Incidence of prédation.—Not searching. AF: Sunflower plants. given. HAB : Idle land where sunflowers Life history and habits.—SO: are growing. (117, p. 20 ; 220, p. Eggs, larvae, and pupae unknown. 6) Description.—Eggs unknown. ^ Larvae (immature) are 1.6 mm. long and generally yellow. Head, prothorax, metathorax, and ab- , dominai segments 5 to 8 are brown. The body surface is cov- ' ered with a flne network of lines. Head is as long as the prothorax . and mesothorax combined. An- tennae are moderately robust ; the second segment is twice as long as > the third. One short and two long setae are on the apex of the second MAP 123.—Recorded distribution of Epicauta callosa LeConte. segment. The terminal seta is 11/^ ARTHROPOD AND NEMATODE PARASITES IN AMERICA 181 times as long as the second seg- wide. (86, pp. 38A-385; H2, p. ment. Mandibles are very robust 99 ; 191, p. 158 ; 3A2, pp. 16, 29) and bear five teeth, all of which Discussion.—This is one of the are visible in outline when viewed few meloids that is beneficial as from above. The thorax is one- a larva and not economically de- third as long as the body and the structive as an adult, since it con- prothorax as long as the meso- fines its feeding to sunflowers and thorax and metathorax combined. has not been noted attacking There are 10 setae in a marginal cultivated or range crops. row on tergites 1 to 8 of the ab- domen and spinelike evaginations Epicauta cinérea (Forster) at the base of the marginal setae (Macrobasis) 1771: 62 on tergites 1 to 5. The sternum Synonymy.—Macrobasis cine- of segments 1 to 7 is weakly rea Forster, M. clemátides Wood- sclerotized, whereas segments 8 house, Lytta fimbreata (Thun- and 9 are well sclerotized. There berg), Epicauta fissilabris are six lanceolate setae on the an- LeConte, L. marginata Fabricius, terior femora. {2H., pp. 61-62) E. marginata (Fabricius) (185, p. Pupae unknown. 158; 390). Adult beetles are 7 to 13 mm. Recorded distribution.—Most long, usually 9 to 10 mm. The of Canada and the United States body is black with a dense pubes- east of the Rocky Mountains {H-9, cence of yellow ash gray, that pp. 37-U2, U6-U9; 21U, pp. U, obscures the surface. The elytra Jt8-A9). (Map 124.) lack any dark basal markings. Host species.—Melanoplus The entire upper surface is finely differentialis (Thomas), M. fem- punctured, whereas the head and urrubrum (DeGeer), M. sanguin- pronotum are punctured minutely. ipes (Fabricius), and M. spretus Head is about one-third wider than it is long and the sides are rounded to the eyes. The pronotum is slightly wider than it is long and is nearly as wide at the base as it is at the head. On the pro- notal disk is a distinct median groove. All tibial spurs are short and flattened internally. The first tarsal segments lack pads. An- tennae are barely past the base of the elytra and are of uniform thickness. The second segment is about one-half the length of the MAP 124.—Recorded distribution of first and is a little longer than Epicauta cinérea (Forster). 182 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(Walsh) {15) 12U, V. k92\ U9, segments are twice as long as the VV. S7-U2, U6-U9\ 222, p. H; 26JÍ, third. On the apex of the second p. 298; 272, p. 265; SSO, p. 198; antennal segment are one short SSI, p. 16; SS6, p. S5). and two long setae. Mandibles are Incidence of prédation.—Not very slender and bear 17 teeth, given. 14 of which can be seen in outline. Life history and habits.—SO: The thorax is less than one-third Eggs are deposited in late August as long as the body and the pro- or early September. Larvae are in thorax as long as the mesothorax the ground from September to the and metathorax combined. Along spring. Pupae occur in the spring. the length of the promesonotum Adults emerge in early August. and mesonotum is a line of dehis- LOD : Eggs require 2 weeks. Lar- cence, which is absent from the val duration of the first five stages metanotum. There are 10 setae in ranges from 15 to 196 days. Pu- a marginal row on abdominal pal stage lasts an average of 14^2 tergites 1 to 8, and spinelike eva- days at TO"" F. Adults unknown. ginations are weakly developed at OWS: Feeding stages and sixth- the base of the marginal setae on instar larvae (coarctate). G/Y: tergites 1 to 6. Sternum with seg- Not given. SA : Eggs. N/H : Un- ments 1 to 6 is weakly sclerotized, known. MOA: Larvae encounter whereas segments 8 and 9 are well egg pods. AF : Includes Symphori- sclerotized. There are seven lan- carpos, groundcherry, tomatoes, ceolate setae on the anterior potatoes, cabbages, pumpkins, femura. Coarctate larvae are beets, beans, lupines, clover, aster, 7.45 mm. long by 4.08 mm. wide. sunflower, and pigweed. HAB: This stage is more robust than the Gardens. Each female lays from same stage of most species of the 117 to 324 eggs per mass at the genus and it is a darker reddish bottom of earthen egg cavities, brown. {21Í, pp, UU, A8-Í9 ; 2U5, averaging 247. (H9, pp. S7-Í2, p. 81 ; SU, p. J^S) U6-U9; 160, p. 180; 222, p. lU) Pupae are 12 mm. by 4 mm. Description.—Eggs are 1.208 and similar superficially to the mm. long and light yellow. Larvae pupae of other species in this (young) are 2.35 mm. long, bas- genus, especially E, immaculata ically yellow, and slightly darker (Say) {220, p. H), Adults are 8 on the head and margins of the to 18 mm. long, elongate, and prothorax. The body is covered robust. They are black basically with a network of fine lines. Head and completely covered with a is slightly shorter than the pro- gray pubescence. Antennae are thorax and mesothorax combined filiform ; the first segments are of and the lateral margins are moderate length but never more slightly emarginated. Antennae than one-half the length of the are moderately robust ; the second third. Mandibles are short, not ARTHROPOD AND NEMATODE PARASITES IN AMERICA 183

reaching beyond the labrum. Head and thorax are densely and finely punctured. The elytra are finely granulate and punctate. They are of normal length and the sides are almost parallel. Tarsal claws are cleft at the base. Wings are usu- ally present. (113, p. 62; H2, pp. 96, 102; 159, p. 152) Discussion.—The larvae of this species are beneficial, but the adults are very destructive to garden plants, especially potatoes. MAP 125.—Recorded distribution of Epicauta conferta (Say). Epicauta confería (Say) (Henous) 1823: 281 given. OWS: Fifth-instar larvae Synonymy.—Meloae conferta and coarctate stage. G/Y: One ~ Say, Henous techanus Haldeman, generation requires 2 to 4 years. H. conferta Say, H. confertus SA: Eggs. N/H: Unknown. (Say), H. confertus LeConte (195, MOA: Larvae. AF: Includes ■ p. 159) flowers and fruits of alfalfa, to- Common name.—It has been matoes, and several cucurbits. called the squash blister beetle HAB : Gardens, meadowlands, (1Í8, pp. 93-99). woods. Recorded distribution.—From About 35 to 224 eggs are laid Montana and North Dakota south in masses at the bottom of hori- to Mexico (85, p. 380; 118, p. 19; zontal, narrow, tubular cavities H8, pp. 93-99; H9, pp. 20-27; about an inch below the soil sur- 18A, p. 159; 339, pp. 15, 25). face. After oviposition, the cavity (Map 125.) is plugged and sealed with soil. Host species.—Melanoplus bi- Young larvae hatch and remain vittatus (Say) and M. differen- in the egg cavity for a day or so tialis (Thomas) (123, p. A92; H9, before they burrow out and dis- pp. 20-27; 21U, pp. U3, A5-A6). perse over the soil surface. Under Life history and habits.—SO: favorable conditions they are able Eggs are laid in June and July. to survive about 20 days without Larvae can be found all year. Pu- food. Apparently the discovery of pae occur in the spring. Adults egg pods is accidental, since the emerge in May. LOD: Eggs re- larvae crawl along the soil surface quire 16 to 26 days. Larval de- and drill into it or into cracks and velopment may total up to 2 to 3 undersurface debris periodically. years; first instar 2 to 7 days. The pod is usually encountered Pupae 10 to 15 days. Adults not along its side, and the larva enters 184 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE at this point. Larvae feed by- and imparts a reddish-orange cast puncturing the egg chorion with to the eggs. {118, p. 19) the sickle-shaped mandibles and Larvae: Horsfall {H8) re- thus cause the fluid contents of ported that the morphological the egg to exude into the mouth. description of this species is sim- One to two eggs are required ilar to that of the black blister for completion of first-instar de- beetle {E. pennsylvanica (De- velopment, which is followed by- Geer) ) except E. confería larvae four additional feeding instars. are much larger. He added that Larvae go through a seventh in- the larvae are reddish brown and star in the spring before pupating the dorsum of the prothorax and unless unfavorable conditions abdominal segments 6, 7, and 8 is cause them to have a late coarc- darker brown. MacSwain {2H) tate stage and a late seventh in- pointed out that superficially E. star, which then cause the adult confería can be confused with stage to be postponed until the members of either the E. funebris following year. The coarctate Horn or E. vittata (Fabricius) stage is resistant to cold and des- group. He noted that the third iccation and under unfavorable segment of the maxillae palpus conditions may allow the larvae is only partly expanded at the to survive for 2 to 3 years before outer basal margin and that there further development takes place. are two distinct size categories of Seventh-instar larvae are some- larvae, a common characteristic times able to revert back to the of the genus Lytta. Lengths of the sixth or coarctate stage when sub- two larval categories are 2.4 and jected to unusually dry conditions. 3.3 mm. Both are light amber with When conditions again become a dark-brown prothorax, and the favorable, regular development head and abdominal segments 5 continues. to 8 are darker brown. Pupae are always found in the The body surface is covered soil in nearly vertical positions with a fine network of lines and with the head upward. (118, p. the head is about as long as the 19; 148, pp. 93-99; 1Í9, pp. 20- prothorax and mesothorax com- 27) bined. Antennae are robust; the Description.—Eggs vary from second segment is 21/2 times as 1.75 mm. to 2.44 mm. in length long as the third. On the apex of the second segment are one short and are orange yellow. Covering and two long setae. Mandibles are the eggs is a reddish, watery liq- robust and contain 12 teeth, nine uid, which is sufficiently sticky to or 10 of which can be seen in keep them in masses, even when outline. The thorax is about one- handled. This liquid is usually third as long as the body, and the seen as droplets on the surface prothorax is longer than the meso- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 185 thorax and metathorax combined. Epicauta fabricii (LeConte) There are 12 to 14 setae in a mar- (Lytta cinérea Fabricius) ginal row on tergites 1 to 8 of the 1798: 119 abdomen. At the base of the mar- Synonymy.—Lyatta cinérea ginal setae on tergites 1 to 5 is a Fabricius (original description L. spinelike invagination that is cinérea Fabricius, but preoccupied moderately developed. Segments in Epicauta), L. fabricii LeConte, 1 to 7 of the sternum are weakly Macrobasis fabricii (LeConte), L. sclerotized and segments 8 and 9 debilis LeConte, M. unicolor are well sclerotized. There are (Kirby) (86, p. 298 ; 1^9, pp. 27- nine lanceolate setae on the an- 32; 195, p. 158). terior femora. {1A8, pp. 93-99; Common name.—Ashgray blis- 21 i, pp. A3, Jf5-A6) ter beetle {H.9, pp. 27-32). Pupae are whitish at first but Recorded distribution.—United become darker with age. This States except Pacific Coast States, darkening begins with a blacken- Idaho, and Nevada (85, pp. 4-07- ing of the eyes. 408; 86, p. 298; 195, p. 158; 358, Adult males are 13 mm. and pp. 46-47). (Map 126.) females 20 mm. long. The body Host species.—Melanoplus bi- is black and opaque. A large vittatus (Say), M. differentialis orange spot is on each side of the (Thomas), M. femurrubrum (De- abdomen. The integument is punc- Geer), and M. sanguinipes (Fab- tured heavily and uniformly and ricius) (149, pp. 27-32; 214, pp. is covered with short black hairs. 53-54 ; 236, p. 26 ; 276, pp. 14-18). Punctures on the head are sepa- Incidence of prédation.—Not rated, on the clypeus distinct, and given. on the labrum remote. Antennae Life history and habits.—SO: are regular. The thorax is as wide Eggs are deposited through the as the head and emarginated at the base. The elytra are rough and shorter than the abdomen; they contain minute elevated points and numerous short, black hairs. {85, p. 380; H9, pp. k3, A5-A6; 193, p. 167; 285, p. 281; 3il, pp. ilO-411 ; 3Jf2, pp. 15, 25) Discussion.—Horsfall (1^8, pp. 93-99) estimated that each larva requires at least 37 grasshopper eggs of M. differentialis to com-

plete development. The adult is an MAP 126.—Recorded distribution of economic pest. Epicauta fabricii (LeConte). 186 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE summer and fall. Larvae occur a long seventh instar. Some gen- from fall to spring ; seventh instar erations therefore require 2 years. in April or May. Pupae occur in Seventh-instar larvae are mobile spring. Adults emerge in May and and burrow into the soil to form again in August and September. a pupal chamber, within which LOD : Eggs require 77 days. Lar- they pupate. (i^P, pp, 27-32 \ vae not given. Pupae 9 to 14 days. 237, p. 26; 276, pp. li-18) Adults not given. OWS: Fifth- Description.—Eggs are 1.286 instar larvae or coarctate stage. mm. long and pale yellow. There G/Y : One generation lasts 1 to 2 is no visible sticky media on the years. SA : Eggs. N/H : Unknown. surface, and they adhere to each MOA : Larvae encounter the prey. other only slightly. Larvae (im- AF : Legume crops, preferably al- mature) are 2.3 mm. long and falfa and soybeans, and Irish light amber. The prothorax is potatoes. Attacks both flowers and usually somewhat darker. A fine leaves. HAB : Not given. network of lines covers the body Although adults first begin to surface. The head is about as long appear in the field by mid-May as the prothorax and mesothorax with a second peak by late August combined and the lateral margins or early September, Horsfall are only slightly emarginated. An- {IJfO) suspected that there is only tennae are robust. The second one generation a year ; the second segment is slightly longer than peak is produced by late emerg- 21/2 times the third. At the apex ence of the first generation. From of the second segment are one 56 to 335 eggs, with an average short and two long setae. Man- of 202, are laid in masses within dibles are moderately robust and tujbular cavities that extend one- have teeth, nine or 10 of which half to 1 inch below the soil sur- can be seen in outline. The first face. Undisturbed soil, as in seven of these teeth are well de- alfalfa fields, is preferred for veloped and the remainder very oviposition. Each female is weak. Third segments of the max- thought to lay one to two egg illary palpi are distinctly less masses in its lifetime of about 40 than twice as long as they are days. Emergence of the larvae broad. The thorax is one-third as coincides with oviposition of the long as the body and the pro- preferred hosts. Larvae of this thorax longer than the mesotho- species do not bypass the sixth- rax and metathorax combined. and seventh-instar stages as does Extending the full length of the E, unicolor (Kirby). Overwinter- pronotum and mesonotum is a ing coarctate larvae begin chang- line of dehiscence, which is only ing to the seventh instar by April slightly indicated on the metano- and May and may or may not have tum. There are 10 setae in a mar- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 187 ginal row on tergites 1 to 8 of the is distinctly longer than the outer. abdomen and spinelike evagina- (85, pp. 407-408; 86, p. 298; IOS, tions at the base of the marginal p. 119; 188, p. 343; 358, pp. 46- setae on tergites 1 to 6. Sternal 47) segments 1 to 6 are weakly sclero- Discussion.—E. fabricii is an- tized and segments 7 to 9 are well other of the grasshopper pred- sclerotized. Seven lanceolate setae ators that is destructive to agri- are on the anterior femora. (21 i, cultural crops during the adult pp. 53-54 ; 245, p. 81 ) Pupae un- stage. known. Adults are 9 to 15 mm. long, Epicauta ferruginea (Say) basically dull black, and heavily (Lytta) 1823: 298 covered with fine white pubes- cence that gives them an ashy or Synonymy.—Lytta ferruginea light-gray appearance. Adults are Say (195, p. 158). elongate and subcylindrical. An- Recorded distribution.—West- tennae, most of the tarsi, and the ern United States except Pacific apices of the tibiae and femora Coast States, Nevada, and parts are not thickly covered with this of North Dakota and Minnesota ashy pubescence. The head is as (85, p. 386-387). (Map 127.) long as it is broad and flat above. Host species.—Grasshoppers Sides of the head converge weakly (123, p. 493; 236, p. 26). to the eyes, and a distinct median Incidence of prédation.—Not line is impressed along the top. given. Second segment of the antennae Life history and habits.—SO: in the male is longer than the Eggs, larvae, and pupae unknown. third and fourth combined and Adults not given. LOD : Unknown. nearly twice as long as broad. In the female the second segment is not quite equal to the third and the fourth together. Antennae in both sexes reach to the basal third of the elytra and taper beyond the middle. The pronotum is nearly as wide as the head and about as wide as long. Scutellum is rather broad. There are two subequal spurs on the protibia ; the anterior one is weakly arcuate. On the mid- tibia the spurs are rather short, tapering, and slightly flattened on MAP 127.—Recorded distribution of the inner surface. The inner spur Epicauta ferruginea (Say). 188 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

OWS and G/Y: Not given. SA: as a larval predator is therefore -^ Eggs. N/H: Unknown. MOA: enhanced. i Larvae encounter food. AF : Blos- soms of thistles (Cirsium), prick- Epicauta funebris Horn lypear (Opuntia polyacantha), 1873: 102 ' and composites. HAB : Not given. (183; 222, p. 5) Synonymy.—None (195, p. '" Description.—Eggs, larvae, and 158). pupae unknown. Adults are 6 to 9 Comomn name.—It has been mm. long, black basically, and called the ebony blister beetle rather densely and roughly cov- (U9, pp. ^1-^6). ered with brownish-yellow or Recorded distribution.—South- , pale-gray pubescence. Antennae, central United States (85, p. 389; tibiae, and tarsi are dark brown 21JÍ, pp. U, U6-U7). (Map 128.) to black and the elytra lack mac- Host species.—Melanoplus bi- ulae. The pronotum is about as vittatus (Say) and M. different- long as broad but not quite as Mis (Thomas) (123, p. 493; H9, broad as the head. Sides are par- pp. U1-U6;21U,PP. U,U6-U7). allel to the apical fourth and then Incidence of prédation.—Not narrowed to the apex. In the male given. ' the protibiae support a single Life history and habits.—SO: - spur ; the outer apical edge of the Eggs are deposited in late sum- tibia is strongly reduced and mer (August-September). Larvae * spurlike. The metatibiae have a occur all through the year, hatch- slender inner spur, which is ing in the fall and spring. Pupae spiniform and flattened on the occur in the spring. Adults appear inner surface. The outer spur is from spring until winter with . very robust with its apex some overwintering. LOD: Eggs obliquely truncate and ovate. An- tennae are robust and thickened beyond the third segment and barely reach the base of the ely- tra. The scape reaches one-fourth across the eye. The second seg- ment is half as long as the first and the last segment as long as the first. (85, pp. 386-387; H2, pp. 95, 98; 159, p. 51 ; 193, p. 167; 285, pp. 298-299 ; 3i2, pp. 16, 27- 28 ; 358, p. ^S) Discussion.—Adults appear not to feed on economically important MAP 128.—Recorded distribution of plants. The value of this species Epicauta funebris Horn. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 189 require 24 to 35 days. Larvae take thorax are yellowish brown to 34 days for the first five larval brown. A fine network of lines is stages. Pupae develop for 12 to on the body surface. The head is 24 days. Adults not located. OWS : as long as the prothorax and meso- Generally coarctate. G/Y: 1 year thorax combined. Lateral margins unless coarctate stage lasts sev- are only slightly emarginate. An- eral years. SA: Eggs. N/H: tennae are moderately robust. The Unknown. MOA: Larvae encoun- second segment is twice as long as ter food. AF : Tomatoes, Irish po- the third. On the apex of the sec- tatoes, clematis, groundcherry, ond segment are one short and wingstem, and cuproseweed. two long setae. Mandibles are ro- HAB : Not located. bust and bear 10 apically rounded Females lay 127 to 316 eggs teeth, six to eight of which can be per mass in tubular cavities. The seen in outline. The third seg- egg stage lasts from 24 to 35 days. ment of the maxillary palpi is ex- Because of the late oviposition, panded at the outer basal margin larval feeding begins later in the and is about twice as long as season than for most of the other broad. Thorax is about one-third blister beetles, but they complete as long as the body, whereas the larval development within a short prothorax is as long as the meso- time. Larvae generally overwinter thorax and metathorax combined. in the sixth instar and this stage There are 10 setae in a marginal may last from several days to row on abdominal tergites 1 to 8. several years, averaging about At the base of the marginal setae 235 days. The seventh iiAstar oc- on tergites 1 to 6 are spinelike curs usually in the spring and evaginations. Segments 1 to 7 of lasts from 6 to 17 days, after the sternum are weakly sclerot- which pupation occurs. Because ized. There are seven lanceolate of the variability and the relation- setae on the anterior femora. ship of the development to cli- {21Jf, pp. U, JÍ6-JÍ7) Pupae un- matic conditions, pupation occurs known. about the same date for all larvae Adults are 16 to 19 mm. long, of this species. {1Í9, pp, 4-1-^6; black, and covered with short, 222, p. 5) sparse pubescence. The entire up- Description.—Eggs are about per surface is punctuated. The 1.58 mm. long and yellow. There is head is slightly wider than long no visible sticky substance on the and a median furrow extends surface of the eggs. The eggs ad- down to the clypeus. Antennae of- here only slightly to each other. ten extend back to the basal quar- {1J,9, pp. U-Í6) ter of the elytra and taper beyond Larvae are about 2.2 mm. long the seventh segment. In the male and yellow. Head and lateral mar- the scape reaches three-fourths of gins of the prothorax and meta- the way across the eye, whereas in 190 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE the female it reaches beyond the hind margin of the eye. In both sexes the scape is longer than the third antennal segment, which is about twice the length of the sec- ond. The pronotum is slightly longer than wide, and the sides are subparallel except at the apex. Scutellum is narrow and acutely triangular. The inner spur of the protibia is distinctly longer than the outer, whereas the spurs of the metatibia are broadened and MAP 129.—Recorded distribution of concave apically. (85, p. 389; Epicauta immaculata (Say). 142, p. 102; 357) Discussion.—Although benefi- seventh instar 15 days. Pupae 10 cial as larvae, the adults are de- to 14 days. Adults not given. structive to agricultural crops. LOD : Unknown. OWS : Coarctate. G/Y: One. SA: Eggs. N/H: Un- Epicauta immaculata (Say) known. MOA: Larvae encounter (Lytta) 1823: 304 food. AF: Garden and forage Synonymy.—Lytta immaculata crops such as Irish potatoes, to- Say, Macrobasis immaculata matoes, peas, beans, onions, cab- (Say), L. articularis Say, L. ful- bages, beets, squash, alfalfa, and vescens LeConte {195, p. 158). sweetclover. HAB : Not given. Common name.—It has been Immediately after emergence called the immaculate blister the adults have an insatiable beetle. appetite. Females lay 25 to 306 Recorded distribution.—Cen- eggs per mass in the bottom of tral and south-central United tubular cavities. These cavities States {85, p. 407; 195, p. 158; are then plugged tightly with soil. 214, pp. 43, 52-53). (Map 129.) Incubation takes 11 to 15 days at Host species.—Melanoplu^ bi- 85° F. in the laboratory and up to vittatus (Say) and M. differen- 80 days in the field where mean tialis (Thomas) (118, pp. 5-10; temperatures are 11° to 18°. Lar- 123, p. 493). vae hatch but remain for several Incidence of prédation.—Not days within the egg cavity before given. burrowing to the surface and dis- Life history and habits.—SO: persing. Those finding egg pods Eggs July and August. Gestation enter them and immediately begin 2 to 3 weeks. Larvae fall to feeding. When the fifth-instar spring: First instar 3 to 5 days, larva has completed feeding, it fifth instar 7 to 13 days, and leaves the egg pod and burrows ARTHROPOD AND NEMATODE PARASITES IN AMERICA 191

downward 3 to 6 inches to con- eight apically rounded teeth, five struct an oval cell in which it of which are visible in outline molts. The sixth instar or coarc- when viewed from above. The tate larva is a resting stage, hard third segment of the maxillary shelled, and incapable of motion. palpi is about twice as long as This stage is extremely resistant broad, and the first segment of to desiccation and protects the the labial palpi is one-half as long larva from several days to several as the second. The thorax is con- years. After molting into the siderably less than one-third the seventh instar, which is mobile, length of the body, whereas the nonfeeding, and resembles the prothorax is about as long as the fifth, it burrows upward to form metathorax and mesothorax com- a chamber 1 to 2 inches below the bined. On the abdomen are about surface of the soil. {118, pp. 5- 10 setae in a marginal row on 10; 160, p, 181 ; 222, pp. 5, 8--9) tergites 1 to 8 and spinelike evagi- Description.—Eggs are about nations at the base of the mar- 1.5 mm. long and 0.5 mm. in diam- ginal setae on tergites 1 to 6. eter, translucent, and yellowish The sternum and segments 1 to 7 white when first laid but become are weakly sclerotized, whereas darker yellow with age. They are segments 8 and 9 are well sclerot- almost cylindrical, elongate-oval, ized. There are seven lanceolate and taper toward the posterior setae on the anterior femora. Legs end. Development of the embryo are three-jointed and support can be seen through the trans- three claws per leg ; the two outer lucent wall of the chorion. {118, ones are spinelike. pp. 5-10) Larvae change greatly in ap- Larvae (first instar) or tri- pearance as they grow older. Sec- ungulins are approximately 3 ond-instar larvae are sluggish, mm. long, light brown to yellow, yellowish-white grubs and cara- and elongate-triangular, tapering bidoid or scaraboid type with con- gradually to the posterior end. The spicuous brown heads. They feed body surface is covered with a and grow rapidly through the next fine network of lines. Head is as four instars, becoming more long as the prothorax and meso- plump and grublike as they do so. thorax combined, and the lateral The sixth-instar or coarctate lar- margins are scarcely rounded. vae are 11 to 14 mm. long, about Antennae are three-segmented 5 mm. wide, and reddish brown. and moderately robust. The sec- Although they are active, their ond segments are twice as long skin is rigid and the legs and as the third. At the apex of the mouth parts are reduced to mere second antenna! segment are one tubercles. Gilbertson and Hors- short and two long setae. Mandi- fall {118) noted that in this stage bles are very robust and bear they resemble "half a peanut 192 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE kernel/' The seventh instar re- impressed, longitudinal line. The sembles the fifth. It is active, ro- pronotum is subquadrate and is bust, grublike, fleshy, much wrin- as long as or slightly longer than kled, and yellowish white. Head is wide. Tibiae, tarsi, and femora horny and yellowish, mandibles are usually black and the tibial are brown, and eyes are minute. spurs elongate, slender, and usu- There are black spots above the ally black and the tibial spurs bases of the antennae. (^^, pp. elongate, slender, and spiniform; 278-279; 118, pp. 5-10 \ 21Í, pp. the outer ones are longer than the JfS, 52-53) inner except those on the pro- Pupae are about 17 mm. long tibiae. Abdominal sternites are by 7 mm. wide, have translucent usually denuded medially. (85, appendages, and are yellowish p, Jf07\ 118, pp. 5-10; lJf2, pp. 89, white when first formed. Wings 93; 193, p. 170; 290, p. 30A) and legs are folded and lie loosely along the side of the bottom of Epicauta lemniscata (Fabricius) the body. Posterior margins of the 1801: 279 prothorax and abdomen support Synonymy.—None (195, p. stiff spines. The pupae become 158). darker as they age ; the coloration Common name.—Threestriped begins at the tips of the append- blister beetle. It has also been ages and extends throughout the called the striped blister beetle body. Several days before emer- and three-lined blister beetle (151, gence the adult begins twitching pp, 32-37). inside the pupal case. (118, pp. Recorded distribution.—^East- 5-10) ern United States except northern Adults are 18 to 25 mm. long. North Dakota and northern Min- The basic black is partly covered nesota (118, pp. 10-H; 159, p. with short, prostrate, cinereous 151; 195, p. 158; 2H, p. 182; hair. The entire body surface is 389), (Map 130.) finely and densely punctuated. Host species.—Melanoplus hi- Head is about one-fourth wider vittatus (Say), M. differentialis than it is long and the sides are (Thomas), Melanoplus spp., and rounded from the vertex to the Chortophaga spp. (118, pp. 10- eyes. Antennae are slender, elon- li; 123, pp. Í58, JÍ93; 21Jf, pp. U, gate, and taper from the fifth seg- 50-51; 236, p. 26). ment onward. Scape of the male Incidence of prédation.—Not reaches halfway beyond the eye given. to the vertex, whereas in the fe- Life history and habits.—SO: male it reaches to almost the hind Eggs are laid in early mid-June. margin of the eye. The second Larvae occur from mid-June to joint of the antennae is as long as mid-September. Pupae appear in the third. The thorax contains an spring. Adults emerge in early ARTHROPOD AND NEMATODE PARASITES IN AMERICA 193 weeks without food. In the early stages larvae become more scara- boid as they feed. Fifth-instar larvae may develop through the sixth and seventh instars, or if temperatures are sufficiently high, they may develop directly into pupae. Therefore winter may be passed by the fifth- or sixth- in- star larvae. At this time larvae leave the egg pods and burrow downward into the soil 1 to 2 MAP 130.—Recorded distribution of inches and pupate during the sum- Epicauta lemniscata (Fabricius). mer. (55, pp. 56^-565; 118, pp. 10-lA; lJf9, pp. 32-37; 160, p. June. LOD : Eggs mature in 10 to 180; 222, p. 5; 236, p. 26; 323, p. 14 days at 80° F. Larvae: First in- 2A2) star takes 2 to 7 days and fifth in- Description.—Eggs are about star 814 days. Pupae require 12 1.4 mm. long, but this varies with to 13 days. Adults not given. the size of the egg mass and the OWS: Fifth and sixth instars. time of oviposition. Eggs laid G/Y : One per year. SA : Egg pods. early in the season or in small N/H: Unknown. MOA: Larvae masses are the largest. {H9, encounter food. AF : Includes soy- pp. 32-37) beans, peppers, tomatoes, pota- Larvae (first instar) are 2.2 toes, cabbages, beans, and alfalfa. mm. long. They are white at first Eat both leaves and succulent but soon change to yellow. The stems, often stripping the plants head, prothorax, and abdominal of all foliage. HAB : Not given. segments 1 to 3 and 7 to 8 are Nearly 150 eggs, on an average, brown. Both the body surfaces are laid per mass in tubular cav- and the median part of the head ities one-fourth inch in diameter are covered with a fine network of located about li/^ inches under lines. The median dorsal parts of the surface of compact soil in the mesothorax, metathorax, and alfalfa fields. Each female aver- abdomen are coarsely covered ages five egg masses. After each with tubercles. The head is as oviposition the cavity openings long as the prothorax and meso- are plugged with moist soil and thorax combined. Antennae are surrounded by a ring of dried soil moderately slender. The second pellets. Several hours after hatch- segments are more than twice as ing the larvae leave the egg cavity long as the third. On the apices in search of grasshopper egg pods. of the second segment of the an- These larvae can survive several tennae are three long setae. Man- 194 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE dibles are moderately robust and very destructive not only because bear 11 apically rounded teeth, it attacks crops of economic im- nine of which can be seen in out- portance but because it is also line. The thorax is less than one- gregarious and thus compounds third as long as the body, and the the damage in a given area. Out- prothorax is as long as the meso- breaks are most frequent in areas thorax and metathorax combined. where alfalfa is widely grown, be- There are 10 setae in a marginal cause this is where grasshoppers row on tergites 1 to 8 of the ab- feed and oviposit and also where domen, and spinelike evaginations a desirable adult food source ex- are well developed at the base of ists. Beetles enter the fields in the marginal setae on tergites 2 swarms and increase in numbers to 5. Sternum and segments 1 to until they become overcrowded or 7 are weakly sclerotized, whereas the field is cut. When this hap- segments 8 to 9 are well sclerot- pens, the swarm leaves. ized. There are seven, rarely eight or nine, lanceolate setae on the Epicauta maculata (Say) anterior femora. {1JÍ9, pp. 32- 37; 21 Jf, pp. U, 50-51 ; 2^5, p. 81) (Lytta) 1823: 340 Pupae are first whitish and be- Synonymy.—Lyatta maculata come darker toward the end of the Say, Epicauta maculata Say, L. pupal stage (1^9, pp. 32-37). conspersa LeConte, Cantharis Adults are 12 to 18 mm. long, punctata Duges, E. media Duges elongate, slender, and subcylin- (195, p. 158). drical. The upper surface is dull Common name.—Spotted blis- clay yellow and the body beneath ter beetle (118, pp. 1^-16, 21). and legs are black. Fine minute Recorded distribution.—From cracks densely cover the upper Idaho into Minnesota, south into surface. E. lemniscata appears Texas, west through Arizona, and similar to E. vittata (Fabricius) north through Montana (85, p. except it has three black stripes 393; 118, pp. IJf-ie, 21; 159, p. on the elytra rather than the two 152; 358, p. 38). (Map 131.) of E. vittata. Antennae are fili- Host species.—Melanoplus san- form, of moderate length, and not guinipes (Fabricius) (118, pp. thickened toward the tip. Mandi- IJf-lG, 21 ; 128, pp. Í58, ^93; 236, bles are short, not reaching be- pp. 3, 26-27). yond the labrum, and obtuse at Incidence of prédation.—Not the tip. Tarsal claws are cleft to given. the base. Wings are usually pres- Life history and habits.—SO: ent and the elytra are of normal Eggs not given. Larvae and pupae length. (85, pp. 398-399 ; 10JÍ, p. unknown. Adults emerge from 279 \ U2, pp. 96,100; 159, p. 151) June to August. LOD: Eggs ma- Discussion.—This species is ture in 10 to 17 days. Larvae: ARTHROPOD AND NEMATODE PARASITES IN AMERICA 195 rax dark yellow and abdominal segments 5 to 9 brown. A fine net- work of lines covers the body sur- face, although the median dorsal part of the head is smooth. The head is shorter than the protho- rax and mesothorax combined. Antennae are slender; the sec- ond segments are nearly three times as long as the third. On the apex of the second segment are one short and two long setae. Man- MAP 131.—Recorded distribution of dibles are slender and bear 12 Epicauta maculata ( Say). apically truncate teeth, nine of which are visible in outline. The First instar 4.1 days, second, thorax is less than one-third the third, and fourth instars 2 days length of the body and the pro- each, fifth instar 13 to 14 days. thorax is as long as the mesotho- Pupae and adults unknown. OWS : rax and metathorax combined. Not given. G/Y: One. SA: Egg- There are 10 setae in a marginal pods. N/H : Unknown. MOA : Lar- row on tergites 1 to 8 of the ab- vae encounter food. AF : Irish po- domen, and spinelike evaginations tatoes, soybeans, alfalfa, sweet- are well developed at the base of clover, and sugarbeets. Feeds on the marginal setae on tergites 1 the tender growth of ñowers. to 6. Sternum and segments 1 to HAB : Not given. 7 are weakly sclerotized, whereas When disturbed the adults drop segments 8 and 9 are well sclero- from their perch and pretend tized. There are seven lanceolate death, remaining absolutely quiet setae on the anterior femora. for several minutes. Eggs are laid {21 A, pp. A6, 56-67; 2i5, p. 81; in masses of 40 to 73 (average 71) 34.4, p. 43) Pupae unknown. and mature in about 13 days. Lar- Adults are 9 to 13 mm. long, vae begin seeking egg pods im- black, and rather densely covered mediately after emergence. Gil- with a pale cinerous-gray pubes- bertson and Horsfall {118) were cence, which lies prostrate. The unable to rear larvae in the lab- arrangement of these prostrate oratory beyond the fifth instar. hairs forms small circular cavities {118, pp. 14.-16, 21; 160, p. 181; in the hair allowing the basal 222, pp. 6, 12-15) black to show through. These Description.—Eggs unknown. spots on the elytra are usually Larvae (first instar) are about 2.6 more distinct than those on the mm. long, basically yellow, with other parts of the body or head. the head, prothorax, and metatho- On the top of the head is an im- 196 TECHNICAL BULLETIN NO. 14«0, U. S. DEPT. OF AGRICULTURE pressed longitudinal line. An- tennae are black and can extend back to the basal one-fifth of the elytra. They taper slightly from the third segment to the end, with the first segment reaching one- half way across the eye in both sexes and the second segment slightly more than one-half as long as the third. Labrum and palpi are glabrous, and the maxil- lary palpi are very dilated at the tip. Sides of the head are feebly MAP 132.—Recorded distribution of rounded to the eyes, which are Epicauta murina (LeConte). elongated. The pronotum is barely longer than wide, and the sides spretus (Walsh) (123, p. U93 ; 261t, are gradually divergent ante- p. 298). riorly. A single, short, recurved Incidence of prédation.—Not '■ spur of the male (two in the fe- given. male) is found on the protibia. Life history and habits.—SO: The metatibial spurs are subequal Eggs not given. Larvae: Seventh ' in length; the outer are slightly instar stage occurs in April. Pu- - more robust than the inner. pae April to May. Adults May to (85, p. 393; H2, pp. 96, 100; 159, June. LOD : Eggs 10 to 18 days. p. 15Í; 192, p. 6; 193, p. 167; Larvae: First five instars require - 285, p. 3U0; 288, p. 101 ; 3Jt2, pp. 12 days, sixth 10 months, seventh 15, 19-21 ; 358, p. 38) 2 weeks. Pupae 10 to 20 days. Discussion.—The beneficial ef- Adults not given. OWS: Larvae, t fects of this species are greatly G/Y: Not given. SA: Egg pods. offset by its harmful ones. N/H: Unknown. MOA: Larvae encounter food. AF and HAB: ^ Epicauta murina (LeConte) Not given. Gilbertson and Horsfall re- (Macrobasis) 1853: 344 ported that under field cage con- r Synonymy.—Macrobasis mu- ditions each female averages 129 ^ rina (LeConte), M. debilis (Le- eggs (77-215) in a single mass Conte) (195, p. 158). in packed moist soil before grass- ' Recorded distribution.—South- hopper eggs are laid. The eggs ern Canada and the United States hatch in 10 to 18 days, and the east of the Rocky Mountains (77, first-instar larvae begin to search * p. 12; 118, p. 19; 159, p. 150). for host eggs. Some first-instar (Map 132.) larvae can survive for 46 days Host species.—Melanoplus without feeding. Thus they are ARTHROPOD AND NEMATODE PARASITES IN AMERICA 197 still present when grasshopper grayish hairs. Antennae are fili- eggs are deposited. Larvae de- form and of moderate length. velop very rapidly once they lo- First antennal segment is elon- cate food. {118, p. 19; 222) gated, and both the first and sec- Description.—Eggs unknown. ond segments of the male an- Larvae are 2.2 mm. long and yel- tennae are thickened. The thorax low brown; the posterior lateral is slightly longer than broad. Tar- m ?ins of the prothorax are pale sal claws are cleft at the base. brown. The body surface is cov- The elytra are of normal length ered by a network of fine lines. and wings are usually present. The head is about as long as the (157, p. 150; 188, p. 344) prothorax and mesothorax com- Discussion.—This species is bined and the lateral margins are probably a very minor predator only slightly emarginate. An- of grasshopper egg pods. tennae are robust. The second seg- Epicauta normalis Werner ments are 21/^ times as long as the third. At the apex of the sec- 1943: 65-66 ond segment are one short and Synonymy.—None located. two long setae. Mandibles are Recorded distribution.—Wash- slender and bear 15 "teeth," 13 ington, Montana, Oregon, Cali- of which can be seen in the out- fornia, Nevada, Wyoming, Utah, line. The first 10 teeth are moder- Colorado, and Ohio (85, pp. 392- ately developed and the apical 393; 183, p. 7; 342, p. 15; 356, teeth are very weak. The thorax pp. 65-66 ; 358, pp. 38-39). (Map is about one-third the length of 133.) the body. The prothorax is longer Host species.—In Wyoming than the mesothorax and meta- from Aulocara elliotti (Thomas) thorax combined, and a line of (183, p. 7). dehiscence extends the full length of the pronotum and mesonotum and is faintly indicated on the metanotum. On the tergites of ab- dominal segments 1 to 8 are 10 setae in a marginal row, and on tergites 1 to 6 are spinelike evagi- nations at the base of the mar- ginal setae. These evaginations are absent at the base of the me- dian transverse row of setae. (214, pp. 43, 54-55) Pupae un- known.

Adults are 8 to 15 mm. long, MAP 133.—Recorded distribution of black, and densely clothed with Epicauta normalis Werner. 198 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Incidence of prédation.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS: Not given. G/Y: Not lo- cated. SA: Egg pods. N/H: Un- known. MOA: Larvae encounter egg pods. AF: Blossoms of Helir- anthus petiolaris Nutt., Cheno- podium spp., Kochia spp., and Salsola spp. (183, p. 7). HAB: Not given. MAP 134.—Recorded distribution of Description.—Eggs, larvae, and Epicauta oblita LeConte. pupae unknown. Adults are very similar to E. maculata in appear- Incidence of prédation.—Not ance. They are black and covered given. with olive-cinereous to cinereous Life history and habits.—SO: pubescence, which is absent in Eggs, larvae, and pupae unknown. various locations, creating numer- Adults not given. LOD : Unknown. ous denuded spots. Maxillary OWS and G/Y: Not given. SA: palpi are small and the same size Egg pods. N/H : Unknown. MOA : in both sexes. Labial palpi are Larvae encounter food. AF and also small. First segment of the HAB : Not given. male anterior tarsi is narrow, Description.—Eggs unknown. slightly curved outward, and lacks Larvae (first instar) are 2.1 mm. the pad of E. maculata on at least long and pale brown except a yel- the basal three-fourths of the lowish-brown head and prothorax. tarsi. (85, pp. 392, 393; 3Í2, p. The lateral margins of the head 15; 356, pp. 65-66; 358, pp. 38- are scarcely rounded and sub- 39) parallel. Antennae are robust ; the second segments are twice as long Epicauta oblita LeConte as the third. Mandibles are very 1851: 162 robust and bear six teeth, five of Synonymy.—None located which are visible in outline when (195, p. 158). viewed from above. There is a Recorded distribution.—Brit- line of dehiscence of all segments ish Columbia and Pacific Coast of the thorax and the first six States (lJf2, pp. 95-97; 187, p. segments of the abdomen. There 162; 195, p. 158; 336, p. 35). are 10 setae in a marginal row on (Map 134.) tergites 1 to 8 and spinelike evagi- Host species.—Grasshoppers nations at the base of marginal (236, p. 26). setae of tergites 1 to 6. Anterior ARTHROPOD AND NEMATODE PARASITES IN AMERICA 199

femora have six lanceolate setae A small midge is attracted to an each. {2H., pp. iS, 6i) Pupae oily secretion that oozes from the unknown. thoracic and femoral-tibial joints Adults are 8 to 10 mm. long, of adult beetles that have fed on elongated, and black. The body is alfalfa and snowberry. Beetles moderately shining and sparsely swarm into fields occupied by clothed with erect black pubes- nymphs of C. pellucida and begin cence. Head and thorax are mod- ovipositing a month prior to erately punctured and the elytra grasshopper oviposition. {236, coarsely punctured. The body be- p. 26; 816, pp. 497-499; 389) neath is more shining than above Description.—Eggs, larvae, and and less punctured. Antennal seg- pupae unknown. Adults are 9 to ments are scarcely one-half times 10 mm. long and basically black. longer than they are thick. (1-^2, They are covered with subopaque pp. 95-97; 187, p. 162) cinereous pubescence. There are Discussion.—A "possible" numerous small denuded spots on grasshopper egg predator. the elytra. The head is sparsely punctured and has a smooth spot Epicauta oregona Horn within and above the insertion of * 1875: 153 each antenna. Antenna are one- i- Synonymy.—None (19i, p. half as long as the body and mod- 158). erately flattened, more so in the Recorded distribution.—From male than in the female. The tho- British Columbia into western rax is slightly narrower than the South Dakota, south along the head. Elytra are slightly wider Rocky Mountains into Colorado behind and sparsely punctured. and New Mexico, south into Cali- The underside of the body and ^^ fornia, and north through the the legs are more shiny than Pacific Coast States {S58, p. 34..). * (Map 135.) Host species.—Camnula pellu- cida (Scudder) (24; 123, p. 498; 236, p. 26; 816, pp. 497-499). p Incidence of prédation.—Not given. Life history and habits.—SO: P Not given. LOD : Unknown. OWS and G/Y: Not given. SA: Egg pods. N/H: Unknown. MOA: Larvae encounter food. AF: Snowberry, alfalfa, iris, lupines, and "vetch" cabbage. HAB: Not MAP 135.—Recorded distribution of given. Epicauta oregona Horn. 200 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE above, sparsely punctured, and covered with pubescence. (148, p. 153; 358, p. 3^) Discussion.—Many grass- hopper egg pods are destroyed in fields infested by larvae of this species. Epicauta pennsylvanica (DeGeer) (Cantharis) 1775: 15 Synonymy.—Cantharis penn- sylvanica DeGeer, Lytta penn- MAP 136.—Recorded distribution of sylvanica (DeGeer), L. penn- Epicauta pennsylvanica (DeGeer). sylvanica LeConte, Epicauta pennsylvanica (DeGeer), Meloe Larvae occur from fall to spring. nigra Woodhouse, L. morio Le- Pupae are present from spring to Conte, E. potosina Duges, (Genus summer. Adults emerge in Au- ?) alrata Fabricius, (Genus ?) gust. LOD : Eggs require 14 to 15 coracina 111. {H9, pp. 3-20; 195, days or may remain as eggs until p. 158) the following spring. Larvae: Common name.—Black blister First instar needs several days of beetle. It has also been called the feeding, second to fifth instars not goldenrod beetle {H9, pp. 3-20) given, and sixth instar needs 6+ Recorded distribution.—United days. Pupae take 11 to 20 days to States except Pacific Coast States, develop. Adults unknown. OWS Idaho, Nevada, and northwest Eggs or larvae. G/Y: One. SA Utah (85, pp. 388-389; 86, pp. Egg pods. N/H : Unknown. MOA 295-297; H9, pp. 3-20; 159, p. Larvae encounter egg pods. AF 151; 3^2, pp. 15, 25; 358). (Map Adults restrict their feeding to 136.) the pollen of goldenrod and small Host species.—Melanoplus wild aster. HAB : Not given. (15, femurruhrum (DeGeer), M. san- p. 10) guinipes (Fabricius), M. spretus Eggs, averaging 165 per mass, (Walsh), and M. differentialis are laid in small tubular cavities. (Thomas) (15, p. 27; 77, p. 12; Oviposition occurs at about 123, pp. 458-Í93; H9, pp. 3-20; weekly intervals. Tubular cavities 236, p. 26; 272, p. 265; 330, pp. are vertical, about one-fourth inch 181-182; 331, p. 77). in diameter and 1 inch deep. Ovi- Incidence of prédation.—Un- position is completed in about 15 known. minutes. The chamber is sealed Life history and habits.—SO: with moist soil. A depression and Eggs are deposited in September. a ring of dried pellets are left ARTHROPOD AND NEMATODE PARASITES IN AMERICA 201 around the plugged entrance. 222, pp. 6,15; 26h. pp. 292, 298; After hatching, the larvae remain 278, p. SSU; S23, p. 2k2) in the egg tube for 1 or 2 days Description.—Eggs are approx- before burrowing to the surface imately 1.3 mm. long and pearly and dispersing. Those locating white to light ivory at first, be- grasshopper egg pods begin feed- coming darker with age. They are ing almost immediately. The oth- nearly spherical or slightly longer ers remain quiet throughout the than wide. The chorion is trans- winter and begin feeding when parent (the developing embryo temperatures rise sufficiently in often visible within) and firm; it the spring. lacks sculpturing. (H9, pp. 3- The larva feeds by puncturing 20) the egg chorion with its mandibles Larvae are 2.1 mm. long, basic- and thus causes the egg yolk to ally yellow, but with the head and ooze into its mouth. Although one prothorax yellow brown. A net- egg contains enough material to work of fine lines covers the body feed one ñrst-instar larva, several surface. The head is about as long eggs are often punctured before as the prothorax and mesothorax this stage is completed. As feed- combined. Antennae are moder- ing continues, the larval body be- ately robust. The second segments comes crescent and engorged. are twice as long as the third. On Within a day or two it molts. Four the apex of the second antennal instars follow in rapid succession, segment are one short and two each stage macerating eggs and long setae. Mandibles are slender ingesting the contents. After the and have 13 apically truncate fifth-instar larva has completed teeth, 10 of which can be seen in its feeding, it leaves the pod and outline. The thorax is less than burrows downward into the soil one-third as long as the body and 1 or 2 inches where it constructs the prothorax as long as the meso- a chamber about twice its length. thorax and metathorax combined. There it molts into either the sixth There are 10 setae in a marginal larval instar or the pupal stage. row on tergites 1 to 8 of the ab- The sixth or coarctate stage is domen. Spinelike evaginations are highly resistant to desiccation and well developed at the base of the temperature change. At comple- marginal setae on terga 1 to 6. tion of this stage the larva molts Sternum and segments 1 to 6 are into the seventh instar and bur- weakly sclerotized. There are rows upward to within an inch seven lanceolate setae on the an- or two of the soil's surface and terior femora. (For a complete constructs another chamber sim- description of the seven instars, ilar to that of the first. It trans- see Horsfall (19^3)^ (2i, p. 278 ; forms into the pupal stage. (15; 21 Jf, pp. Uy U7; 2h5, p. 81; 3U, lJf9, pp. 3-20; 160, p. 81; 208; p. Jf3) 202 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Pupae are about 10 mm. long, ing is limited to pollen, this spe- creamy white at first, and black cies is a desirable predator. From later. Legs are folded loosely on 21 to 27 eggs of M. differentialis the sternum, reaching from the are necessary for one larva to eyes nearly to the anus. Both the complete development. If two body and the legs are sparsely larvae are in the same egg pod, spinate. {IJ^Q, pp. 3-20) one will destroy the other or both Adults are from 7i/^ to 12 mm. will be killed. long, black, and sparsely covered with short pubescence of pros- Epicauta puncticollis (Mannerheim) trate black hairs. Both the head and pronotum are densely cov- 1843: 288 ered with coarse punctures and Synonymy.—None (195, p. minute cracks. The head is weakly 158). rounded to the eyes and contains Recorded distribution.—Brit- a median furrow that usually ex- ish Columbia and Pacific Coast tends to the clypeus. Antennae are States {195, p. 158 ; 2H, pp. 3, 62- robust and can attain the basal 63; 236, pp. 3, 26-27; 336, p. 35). fifth of the elytra. The scape of (Map 137.) the male reaches the middle of Host species.—Grasshoppers the eye, whereas in the female it {123, p. 4.93 ; 220, p. 9 ; 236, pp. 3, almost reaches the hind margin. 26-27; 336, p. 35). Antennal joints on the apical half Incidence of prédation.—Not are more slender, loosely united, given. and more or less compressed than Life history and habits.—SO: on the basal half. The thorax is Eggs, larvae, and pupae unknown. quadrate with the pronotum a Adults not given. LOD : Unknown. little longer than wide. The sides OWS and G/Y: Not given. SA: of the latter are weakly divergent anteriorly and strongly con- structed on the apical fifth. The inner spur of the protibia is dis- tinctly longer than the outer, and the spurs of the mesotibia are ex- panded and concave apically. The outer spur on the posterior tibia is broader than the inner and somewhat spoon shaped. {82, p. 15; 85, pp. 388-389; 86, pp. 295- 297; 14.2, pp. 96, 103-lOA; U9, pp. 3-20; 159, p. 151; 3i2, pp. 15,25; 358, p. 33) MAP 137.—Recorded distribution of Discussion.—Since adult feed- Epicauta punetieollis (Mannerheim). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 203

Egg pods. N/H : Unknown. MOA : They key out to E. pennsylvanica Larvae encounter food. AF : Sym- in Jaques (159). Except for the phoricarpos. HAB: Not given. A bluish sheen and luster they are very common species. (336, p. very difficult to separate. There- 35) fore the adult description will not Description.—Eggs unknown. be treated here. (1^2, pp. 95-97 ; Larvae (young) are approxi- 159, p. 151 ; 215, p. 288) mately 2.7 mm. long and brown. Discussion.—Treherne and The head and prothorax are yel- Buckell (336) indicated that this lowish brown. Body surface is species is possibly a predator of covered with a network of fine grasshopper eggs. lines. The head is about as long Epicauta segmenta (Say) as the prothorax and mesothorax combined, and the lateral margins (Lytta) 1823: 303-304 are scarcely rounded and sub- Synonymy.—Lytta segmentata parallel. Antennae are moderately Say, L. segmenta Say, L. seg- robust. The second segments are mentata LeConte, Macrobasis seg- less than 21/2 times as long as the mentata (Say), Macrobasis seg- third. At the apex of the second mentata Horn, Microbasis segment are one short and two cinclothorax Duges, Apterospasta long setae. Mandibles are moder- valida LeConte, Epicauta seg- ately robust and bear seven api- mentata (Say) (195, p. 158). cally rounded teeth, five notice- Common name.—It has been able in outline when viewed from called the white segmented blister above. The thorax is about one- beetle. fourth as long as the body. The Recorded distribution.—South- prothorax is shorter than the western United States except Cali- mesothorax and metathorax com- fornia and Nevada (118, p. 20; bined. On all segments of the tho- 195, p. 158; 2H, pp. i5, 53; 3U2, rax is a line of dehiscence, which pp. 17, 31-32 ; 358, p. 35). (Map is absent from the abdomen. 138.) There are 10 setae in a marginal Host species.—Melanoplus bi- row on tergites 1 to 8 of the ab- vittatus (Say), M. differentialis domen and spinelike evaginations (Thomas), and M. sanguinipes at the base of the marginal setae (Fabricius) (117, p. 20; 212, pp. on tergites 1 to 6. Sternum is U5-53; 355, p. 35). weakly sclerotized on segments 1 Incidence of prédation.—Not to 7 but well sclerotized on seg- given. ments 8 and 9. There are six Life history and habits.—SO: lanceolate setae on the anterior Eggs, larvae, and pupae unknown. femora. {212, pp. 3, 62-63) Adults not given. LOD : Unknown. Pupae unknown. OWS and G/Y: Not given. SA: Adults are 7 to 11 mm. long. Egg pods. N/H : Unknown. MOA : 204 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

metathorax combined. A line of M^> dehiscence extends along the full length of the pronotum and meso- notum but not on the metanotum. A marginal row of 10 setae is lo- cated on abdominal tergites 1 to 8, and weakly developed spinelike evaginations are at the base of the marginal setae on tergites 1 to 6. The sternum segments 1 to 7 are weakly sclerotized, whereas seg- ments 8 and 9 are well sclerotized. MAP 138.—Recorded distribution of Several lanceolate setae are lo- Epicauta segmenta (Say). cated on the anterior femora. {2H, pp. U5, 58) Pupae un- Larvae encounter larvae. AF: known. Although potatoes and several Adults are 15 to 25 mm. long, other garden crops are attacked, basically black, and covered with Russian-thistle seems preferred. black pubescence. The entire up- HAB: Not located. (118, p. 20; per surface is densely and finely 160, p. 181 ; 222, p. 5) punctate and covered with minute Description.—Eggs unknovs^n. cracks. Head, underside, mandi- Larvae (young) are 3.1 mm. long bles, genae and occiput, pronotal and basically yellow. The head apex and basal margin, scutellum, and prothorax are dark yellow and elytral apices, coxae, sterna, and abdominal segments 6 to 9 are abdominal sternites are covered brown. A network of fine lines or edged with white pubescence. covers the body surface. The head Head is one-fourth wider than is slightly longer than the protho- long and the sides are rounded to rax and mesothorax combined, the eyes. Antennae with the scape and the lateral margins are reaches nearly to the hind margin slightly emarginated. Antennae of the eye in the female and be- are moderately slender. The sec- yond in the male. Pronotum is ond segments are twice as long slightly longer than it is wide. as the third. At the apex of the The sides are parallel except at second segment are one short and the apex where they are strongly two long setae. Mandibles are ro- constricted. T\vo long spurs, bust and bear 11 apically rounded which are concave on the mesal teeth, nine to 10 of which can be facies are located on each tibia. seen in outline. The thorax is First segment of the tarsi is dis- about one-fourth the length of the tinctly longer than the next two body. The prothorax is slightly combined. {118, p. 20; H2, pp. shorter than the mesothorax and 89, 93-9A; 193, p. 169; 285, pp. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 205 303-30^; 3U2, pp. 17, 31-32; 358, Egg pods. N/H : Unknown. MOA : p. 35) Larvae encounter food. AF : Blos- Discussion.—Since there is gen- soms of Aster concinnus. HAB : erally an ample supply of Russian- Unknown. (55, pp. 563, 56^; thistle to satisfy the adults, they 160, p. 181 ; 222, pp. 6, 15) generally cause little economic Description.—Eggs unknown. damage. The predaceous larval Larvae (third instar) are 7 to 8 habit combined with the food hab- mm. long by 2.5 mm. wide. Color its of the adult make it a more de- and appearance of E. sericans and sirable species than most of the E. Immaculata are very similar. meloids. Coarctate larvae are 6.9 mm. long Epicauta sericans LeConte by 3.8 mm. wide. This stage is reddish brown but is not as dark 1866: 158 as in E. cinérea. The lateral ridges Synonymy.—Epicauta im- are present and distinctly angu- merita Walker (195, p. 158). lar. (222, pp. 6, 15; 3U, p. U3) Recorded distribution.—West- Pupae unknown. ern and central United States east Adults are 8 to 13 mm. long of the Rocky Mountains (85, pp. and black. They are covered with 385-386; 389) (Map 139.) a uniform, dense, pale ashy-gray Host species.—Melanoplus spp. pubescence. Upper surface of the (77, p. 12; 123, p. U93; 236, p. 26; body is finely punctate and is cov- 330, p. 181; 331, p. 77). ered with minute cracks; the Incidence of prédation.—Not elytra are more rough than the given. other parts. Sides of the head are Life history and habits.—SO: rounded to the eyes, and a me- Eggs, larvae, and pupae unknown. dian furrow on the top can be Adults not given. LOD : Unknown. seen when viewed from above. OWS and G/Y: Not given. SA: Pale pubescence covers the first two segments and the basal part of the third segment of the an- tennae. Antennae scarcely attain the base of the elytra but grad- ually thicken to the ninth segment. Scape of the female reaches one- fourth across the eye, whereas in the male it reaches one-third the ï^ distance across the eye. The pro- |.-'^,>x, notum is about as long as broad and nearly as wide as the head across the occiput. Sides of the MAP 139.—Recorded distribution of elytra are nearly parallel. The Epicauta sericans LeConte. outer spur of the protibia is 206 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE longer than the inner. Spurs of Eggs, larvae, and pupae unknown. the mesotibia are subequal; the Adults not given. LOD : Unknown. outer one is more robust. Inner OWS and G/Y: Not given. SA: spur of the metatibia is slightly Egg pods. N/H : Unknown. MOA : longer than the outer; the latter Larvae encounter food. AF and is a little wider apically. (85, pp. HAB : Not given. {300, p. 60) 385-386; H2, pp. 95, 98', 191, p. Description.—Eggs unknown. 158; 3Jf2, pp. 16, 29-30; 358, p. Larvae (full grown) are about 9 U) mm. long and grayish white ex- cept the dark mandibles. There Epicauta subglabra (Fall) are narrow rings around the base (Macrobasis) 1922: 173 of the antennae and claws. Much Synonymy.—Macrobasis sub- of the head and the entire body glabra (Fall) (196, p. 36). bear numerous short pointed Recorded distribution.—Al- setae. The body is plump and berta, Saskatchewan, and Mani- slightly curved, resembling a cur- toba south into Arizona, New culionid larva except three pairs Mexico, and northwest Texas (77, of short, fleshy, four-segmented p. 12; 107, p. 173; 358, p. U7 ; legs located far apart and termi- 386) (Map 140.) nating in tiny pointed claws. The Host species.—Grasshoppers head lacks the epicranial suture and Melanoplus spp. {77, p. 12; or clypeus but does possess a lab- 123, p. Í93; 196, p. 36; 236, p. 26; rum and a pair of inconspicuous 276, p. 15; 300, p. 60; 330, p. 181 ; eye spots. Spiracles are located on 331, p. 98). the lateral aspects of the meso- Incidence of prédation.—Not thorax and abdominal segments 1 given. to 8. {2A5, p. 81) Pupae un- Life history and habits.—SO: known. Adults are 7 to 10 mm. long, slender, and entirely black. The ¡IV : M head and thorax are sparsely but i > finely punctate. The elytra are finely rugose without being dis- tinctly punctate. Antennae are similar to those of E. unicolor (Kirby). The first segment of the male is slender and reaches be- yond the vertex of the head. The second antennal segment is slen- der and slightly curved. It is about two-thirds the length of the first MAP 140.—Recorded distribution oí and equal to the following three Epicauta subglabra (Fall). segments combined. Protibiae of ARTHROPOD AND NEMATODE PARASITES IN AMERICA 207 the male has two spurs. (,107, p. 178; 358, p. h7) Epicauta vittata (Fabricius) 1775: 260 Synonymy.—None (195, p. 158). Common name.—Striped blis- ter beetle. Recorded distribution.—South- eastern Canada and eastern United States. (Sweetman {1958) places E. vittata in the MAP 141.—Recorded distribution of West also). {86, pp. 295-297; Epicauta vittata (Fabricius). 323, p. 2Jt6) (Map 141.) Host species.—Melanoplus dif- from the egg chamber and seek fer entialis (Thomas), M. san- the grasshopper egg pod, which guinipes (Fabricius), and Mela- they immediately penetrate and noplus spp. (123, p. Í63; 264-, P- begin to feed. Within several days 298 ; 265, p. 260 ; 272, p. 264 ; 323, two of the eggs are devoured, p. 346; 330, pp. 181-182; 331, p. after which the larva rests for a 77). short time and then molts. The Incidence of prédation.—Not second instar is carabidoid, dif- given. fering greatly from the triun- Life history and habits.—SO: gulin. The next several molts oc- Eggs, larvae, and pupae unknown. cur about a week apart, after Adults not given. LOD : Eggs un- which the larva leaves the egg known. Larvae: Second and third pod and burrows into the soil to instars 1 week each. Pupae 5 to 6 form a smooth cavity. Within this days. Adults unknown. OWS: cavity it molts into the sixth in- Coarctate. G/Y: Two per year. star or coarctate stage. After SA : Eggs. N/H : Unknown. MOA : molting into the seventh instar it Larvae encounter food. AF : Adult burrows into the ground and beetles prefer the foliage of soy- forms a chamber where it pu- beans. HAB: Not given. pates. (15; 55, pp. 564-566 ; 160, The female lays her eggs loosely p. 180; 222, pp. 18-19; 264, p. in chambers within the soil. Each 298 ; 265, p. 260 ; 266, pp. 91-93 ; female lays several masses of 323, p. 246) about 130 eggs each. The cham- Description.—Eggs unknown. bers are usually in the same areas Larvae (young) are 2.9 mm. long that the host grasshopper uses and yellow. The head, prothorax, for its egg laying. After hatching, and abdominal segments 1 to 3 the young triungulins emerge and 7 to 8 are brown. The body 208 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE surface is covered by a fine net- notum are finely but densely punc- work of lines except the median tuated, and the elytra are densely > part of the head, which is smooth. granulated. The head is wider i The median dorsal parts of the than the thorax. Antennae are of mesothorax and metathorax and moderate length, filiform, and not abdomen are finely tuberculate. thickened toward the tip. Mandi- ; The head is as long as the pro- bles are short, not reaching be- thorax and mesothorax combined. yond the labrum, and obtuse at '> Antennae are moderately slender. the tip. Wings are usually present, ; The second segment is twice as and the elytra are of normal long as the third. On the apex of length. (86, pp. 295-297; H.2, - the second segment are three long pp. 96, 100; 159, p. 15; 160, p. < setae. Mandibles are moderately 180) robust and bear 15 apically rounded teeth, 11 of which can be Linsleya sphaericollis (Say) '^ seen in outline. The thorax is con- {Lytta) 1823: 299 siderably less than one-third as long as the body. The prothcrax Synonymy.—Lytta sphaericol- - is as long as the mesothorax and lis Say, L. ckalybea (LeConte) metathorax combined. There are (193, p. 159; 295, pp. 22, 202- > 10 setae in a marginal row on ter- 203). ; gites 1 to 8 of the abdomen, and Recorded distribution.—East- well-developed spinelike evagina- ern Washington, Oregon, Mon- tions are at the base of the mar- tana and Kansas (295, p. 22, 202- ginal setae on tergites 2 to 5. 203). (Map 142.) Sternum and segments 1 to 7 are Host species.—Grasshoppers weakly sclerotized, whereas seg- (77,p.l2;123,p.i93;276,p.l5; •, ments 8 and 9 are well sclerotized. 330, p. 181; 331, p. 78). There are nine and sometimes 10 lanceolate setae on the anterior femora. {2JÍ, p. 278; 2H, pp. A9-50', 3H) Pupae unknown. Adults are 12 to 18 mm. long, elongate, slender, and subcylin- drical. The upper body surface of the head, thorax, and elytra is dull clay yellow, and there are two longitudinal black stripes on each elytron. The underparts of the body and legs are black. The entire upper surface of the body is covered by a dense network of MAP 142.—Recorded distribution of fine cracks, the head and pro- Linsleya sphaericollis (Say). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 209

Incidence of prédation.—Not given. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Egg pods. N/H : Unknown. MOA : Larvae encounter food. AF and HAB : Not given. (297, p. 60) Description.—Eggs, larvae, and pupae unknown. Adults are black- ish green with a slight tinge of brass. Head is punctured. Anten- MAP 143.—Recorded distribution of Lytta cyanipennis LeConte. nae are robust and black. Joints are short, conical, and acute at the edge of the tip. The labrum Eggs not given. Larvae and pupae and palpi are blackish. Thorax is unknown. Adults March to Au- subglobular and sparsely punc- gust. LOD: Eggs 8 days. Larvae tured. Elytra are green, slightly 36 days total. Pupae and adults tinged with an olive or brassy unknown. OWS and G/Y: Not color, and somewhat rugose. given. SA : Eggs. N/H : Unknown. There are three weak longitudinal MOA: Larvae encounter food. lines on the disk and another near AF : Leaves and flowers of lupines the extreme margin. Beneath is and vetches. HAB : Not given. a blackish green. (192, p. 7;19S, During courtship the male ex- p. 168; 290, p. 299) tends the antennae forward and strokes the antennae of the fe- Lytta cyanipennis LeConte male, violently shakes his body, (Cantharis) 1851: 160 and vibrates his extended abdo- Synonymy.—Cantharis cyani- men so rapidly that it appears pennis LeConte, Lytta solids Le- blurred. Selander (296) once Conte {195, p. 158). observed that this dance lasted Recorded distribution.—Brit- for 5 hours with only brief inter- ish Columbia and United States vals of rest. Couplation takes 11 west of the Rocky Mountains to 12 hours, although insemina- (159, p. 150; 195, p. 158; 295, pp. tion is believed to be short. 3, 9, 11-15, 19, 23, U, U7, 207, The oviposition burrow varies 209-213; 336, p. 35). (Map 143.) from 1 to 21/^ inches deep and is Host species.—Grasshoppers at a 45° angle from the soil sur- (336, p. 35). face. The females apparently pre- Incidence of prédation.—Not fer to excavate the burrow at given. night. After ovipositing, they Life history and habits.—SO: cover the burrow entrance, which 210 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE is difficult to detect. First-instar larvae are pale and very active. When touched, they characteris- tically roll into a ball. They can survive for 15 days without food. {159, p. 150; 296, pp. 3, 9, 11-15, 19, 32, U, A7, 205, 209-213) Description.—Eggs, larvae, and pupae unknovi^n. Adults are 13 to 15 mm. long and metallic green. The head is shining and sparsely punctured. Antennae of the male are moderately thickened at the MAP 144.—Recorded distribution of joints, whereas those of the fe- Lytta infidelis Fall. male are gradually thickened to the tip. The thorax is sparsely Incidence of prédation.—Not punctured. Elytra are parallel, given. bluish or violet, and finely scab- Life history and habits.—SO rous. The body has a bluish or and LOD: Unknown. OWS and . greenish metallic luster. Spurs of G/Y : Not given. SA : Eggs. N/H : ' the hindtibia of the female are Unknown. MOA : Not given. AF : dissimilar; the inner is slender Found on lupines and vetches. - and acute and the outer is slightly HAB : Not given. (336, p. 35) stouter, very oblique truncate, and Description.—Eggs, larvae, and laminate at the tip. The hind tro- pupae unknown. Adults are 8 to chanters of the male bear short 15 mm. long, deep blue to black, spines at the middle and those of and slightly longer than wide. Fall ' the female are neither armed with (106, p. 303) did not give a de- • a spine nor subangulate. {1A2, tailed description but instead used pp. lOA-107; 159, p. 150; 187, p. these characteristics to distin- ' 160; 296, pp. 3, 9, 11-15, 19, 22, guish it from Linsleya sphaeri- U, h7, 205, 209-213) collis. Discussion.—Treherne and ^ Lytta infidelis Fall Buckell (336) indicated this spe- ^ 1901: 303 cies is a "possible" grasshopper Synonymy.—None (195, p. egg predator. 159). Recorded distribution.—West- Lytta nuttalli Say ern and south-central Canada and 1824: 5 s northwestern United States (106, Synonymy.—Cantharis nuttalli p. 303; 336, p. 35). (Map 144.) Horn, C. fulgifer LeConte, Lytta Host species.—Grasshoppers fulgifer LeConte, L. nuttallii Say, (336, p. 35). L. nutalli Say, L. nuttalli var. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 211 fulgifera Ulke, C. nuttali var. beets. (296, pp. 3, 9, 11, 32, U, fulgifera Putnam (195, p. 158; U7, 205-209) 296, pp. 3, 9, 11, 32, U, h7, 205- Description.—Eggs unknown. 209). Larvae (first instar) are slightly Recorded distribution.—Al- over 3 mm. long and dark brown. berta, Saskatchewan, Manitoba, The head and prothorax are western United States except slightly darker. The head is northwest Washington, and slightly wider than long, and the north-central United States (296, lateral margins are almost evenly pp. 3, 9, 11, 32, U, U7, 205-209; rounded at the base. Second seg- 358, pp. 1-62). (Map 145.) ment of the antenna is about one- Host species.—Grasshoppers half as long as the third. There (296, pp. 3, 9, 11, 32, U, U7, 205- are about eight distinct toothlike 209). serrations on the inner margin of Incidence of prédation.—Not the mandibles. Maxillary palpi are given. weakly sclerotized with the first Life history and habits.—SO : two segments subequal. On the Eggs, larvae, and pupae unknown. thorax is a line of dehiscence that Adults not given. LOD : Unknown. is complete on all segments. Most OWS and G/Y: Not given. SA: setae of the thoracic tergites are about as long as most of the ab- Eggs. N/H: Unknown. MOA: dominal setae. (2H; pp. 79, Larvae encounter food. AF : Oats, 87-88) Pupae unknown. barley, Leguminosae australagus, Adults are 16 to 18 mm. long Caragana arborescens, Cnemido- and bright green varied with a phacos pectinatus, Diholcos bisul- golden or brassy to purplish re- catus, Vicia americana, Thermop- flection. The head and pronotum sis montanus, and some beans and are brassy green and usually edged with purple. The under- surface is metallic dark green or blue green to purple along the edges of the sclerites. Femora and tibiae are metallic green, blue, or purple and the tarsi are black. Wings are nearly colorless. Antennae are of moderate length, heavy at the apex, and not thick- ened toward the tip. The elytra are sparsely covered with short pubescence. Mandibles do not reach beyond the labrum and are MAP 145.—Recorded distribution of obtuse at the tip. Tarsal claws are Lytta nuttalli Say. cleft to the base. (H2, pp. lA, 212 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE 106-107; 159, p. 150; 192, p. 5; 193, p. 168; 285, p. 301 ; 288, p. 5; 296, pp. S, 9, 11, 32, U, i7, 205- 209 ; 358, pp. 61-62) Discussion.—Selander (296) reported that only occasionally did this species injure oats and barley in western Canada. However, no incidence-of-prédation records were located.

Pomphopoea aenea (Say)

1823: 301-302 MAP 146.—Recorded distribution of Synonymy.—Pomphopoea nig- Pomphopoea aenea (SB.y). ^ ricornis LeConte, P. tarsalis Bland, P. filiformis LeConte, Description.—Eggs, larvae, and Lytta aenea Say, P. pyrivora pupae unknown. Adults are 13 to ' Fitch (195, p. 157). 16 mm. long and greenish blue to , Recorded distribution.—Most brassy. The elytra are glabrous of Canada and the United States and brassy to purplish, and the including Alaska (193, p. 168; body is hairy. The head is pune- ^ 286, pp. 301-302; 336, p. 35). tured and hairy, and the eyes are (Map 146.) oval. Antenane are longer than Host species.—Grasshoppers the thorax and black. Palpi, knees, (336, p.35). and trochanters are black black- Incidence of prédation.—Not ish. Thorax is also punctured and ' given. hairy. There are two obsolete * Life history and habits.—SO elevated lines on the elytra. (19S, and LOD: Unknown. OWS and p. 168; 285, pp. 301-302) ' G/Y: Not given. SA: Eggs. N/H: Discussion.—Treherne and Unknown. MOA : Not given. AF : Buckell (336) indicated it is a Symphoricarpos. HAB: Un- possible grasshopper egg pred- ^ known. (336, p. 35) ator.

Díptera

Anthomjâidae tion as to the validity of past This is the only known family identification of the predatory of Diptera with species that are members of this family, but since both parasitic and predatory on there is no new evidence to in- grasshoppers. There is some ques- dicate any change, the two spe- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 213 cies in question will be treated tubercles. (264, VP- 285-289) here. Pupae are pale brown and rounded normally at each end. Hylemya angustifrons The prothoracic spiracles, lips, (Meigen) anal spiracles, and lower tuber- (Anthomyia) 1826: 146 cules appear only as minute Synonymy.—Anthomyia an- points. (264, pp. 285-289) gustifrons Meigen, Anthomyza Adults are approximately 17 scatrophagina Zetterstedt, Hy- mm. long and ash gray with a lemya brevitarsata Malloch (320, metallic reflection. Wings are p. sm. faintly smoky with brown-black Recorded distribution.—South- veins. The haltères are yellowish ern Canada and the United States and appear crumpled. The face is including Alaska. Also reported in whitish and the eyes are brown- the Euro-Asian countries and ish and surrounded by the ground Greenland. {26Jf, pp. 285-289; color. Forehead is a dusky brown 320, p. 8^7). (Map 147.) becoming a bright yellowish red Host species.—Only Melanoplus toward the base of the antennae. spretus (Walsh) recorded. Mouth parts and antennae are Incidence of prédation.—Ten black; the stylus is simple and percent of the grasshopper eggs longer than the entire antennae. in Missouri, Kansas, and Ne- Three dusky longitudinal lines on braska during 1876 (26i, pp. 285- the thorax become obsolete be- 289). hind. The legs are black and ash Life history and habits.—SO: gray beneath. Very light, dusky Eggs, larvae, and pupae unknown. spots are on the medial-dorsal Adults not given. LOD : Unknown. part of the abdomen. The abdomen OWS and G/Y: Not given. SA: Eggs. N/H : Unknown. MOA and Et-'^ói AF: Not given. HAB: Wide range. (264, pp. 285-289) Description.—Eggs are oval, smooth, white, and about 1 mm. long (264, pp. 285-289). Larvae (instar stage unknown) are about 6 mm. long and normal in general shape. The skin is unarmed and the prothoracic spiracles are elon- gate. The mandibular hooks are conspicuous and diverging at the base. The anal spiracles are very Ki^ I } . small, yellowish brown, and sur- MAP 147.—Recorded distribution of rounded by eight small fleshy Hylemya angustifrons (Meigen). 214 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE is broad at the base and tapers Recorded distribution.—South- toward the end of each joint. The ern Canada and the United States eyes of the male are much larger including Alaska (320, p. 850). and the bristles narrower and (Map 148.) stronger than in the female. The Host species.—Camnula pellu- face is whiter and the abdomen cida (Scudder) and Melanoplus less tapering. The medial-dorsal spretus (Walsh) (123, p. 487; dark mark of the abdomen is con- 264, p. 288 ; 323, pp. 31, 264, 457). tinuous and the hindtibia are Incidence of prédation.—None armed with a comblike series of located. fine hairs on the underside. (217, Life history and habits.—SO: p. 146; 26A, pp. 285-289) Eggs, larvae, and pupae unknown. Discussion.—Riley (264) re- Adults not given. LOD : Unknown. ferred to H. angustifrons as the OWS and G/Y: Not given. SA: most common and widespread of Eggs and adults. N/H : Not given. all the egg enemies of the grass- MOA: Eggs and adults of host. hoppers in 1876 and credited it AF : Unknown. HAB : Not given. with destroying 10 percent of the (55, pp. 417-418; 257, p. 65; 264, eggs, on an average, in Missouri, p. 288; 323, pp. 31, 264, 457) Kansas, and Nebraska. He also Description.—Eggs unknown. reported it from Iowa, Minnesota, Larvae not located. Pupae not lo- Colorado, and Texas. This species cated. Adults are about 4 mm. long apparently attacks only the egg and basically gray. The thorax is stage. (264, pp. 285-289) dark with three black to brown stripes. The abdomen is also lined Hylemya platura (Meigen) with a dark stripe. Head of the (Anthomyia) 1826: 171 female is white and black pos- Synonymy.—Anthomyia pla- tura Meigen, A. cana Macquart, Aricia ruficeps Meigen, A. fusci- ceps Zetterstedt, Anthomyia tinia Walker, A. perrima Walker, A. viana Walker, Eriphia marginMa Walker, Dialyta cupreifrons Walker, Hylemya deceptiva Fitch, Hylemyia similis Fitch, Choro- phila cilicrura Rondani, Hylemya cilioraca (error) Hagen, An- thomyia zeas Riley (error zeae), Homalomyia rupecula Bigot (320, p. 850).

Common name.—Seedcorn MAP 148.—Recorded distribution of maggot. Hylemya platura (Meigen). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 215 teriorly; the face has yellowish fliers and can transport prey at bristles. The thorax is black to least as large as themselves in ash gray with some pale stripes the air. They have a voracious and some black spots. The abdo- appetite and are not particular as men is ash gray with black spots. to the type of insect collected so A stripe on the frons of the fe- long as it flies. On occasion jump- male in front is rust colored. ing insects are also included in {217, pp. 171^172) the diet. Discussion.—There is an in- Since very few species exhibit teresting problem with this spe- a preference for any type of in- cies. Riley (26Í) reported this sect, some writers question species as attacking the eggs of whether the family should be Camnula pellucida, whereas Boze- considered as predominantly ben- man records {389) indicated eficial or injurious. The flies at- about 80 specimens were collected tack destructive insects, such as from C. pellucida nymphs and grasshoppers, but also attack adults during the summer of 1925 beneficial insects, such as honey at Chester, Calif. Although both bees and other pollinators. For records were substantiated by example, Proctacanthus milbertii Diptera taxonomists, it is possible Macquart preys on grasshoppers that one or both may be in error. and other files and also attacks First, it is highly probable that honey bees, which gave it its nick- the Bozeman records may refer name **the bee killer.'' To make to the anthomyiid fly Acridomyia matters worse, many of these canadensis Snyder, which was not asilids feed on some of their own named until 1940. The patterns species. are similar. Second, H. pintura is Mating may occur any time the seedcorn maggot, a common after emergence and may or may pest in the larval stage of veg- not be preceded by a courtship etables. It is therefore improbable dance. The act may be initiated that the same larvae are also pred- in the air or on upright vegeta- atory. Identification of the first tion. This is generally accom- record may then have been in panied by both members loudly error, or possibly the maggots buzzing their wings. Should the were picked up in the area of the pair be disturbed, they usually egg pods and the assumption was depart in tandem with the same made of their feeding habits. loud buzzing noise. Eggs may be laid near water, on grass, soil, bark, or wood, the Asilidae location varying with the species. The robber flies can be said to Eggs may be laid singly but often be the veritable assassins of the are deposited in masses of con- sky. Adults are swift, strong siderable size and covered with a 216 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE soft chalky white material. After Of the 856 species in 82 genera a short incubation period the lar- in North America, 26 species in vae hatch and enter the soil. They seven genera are recorded here are vermiform in all instars and as preying on grasshoppers. Of feed mainly on the various stages these, six definitely prefer grass- of any soft-bodied insects or in- hoppers. sect eggs they find. In certain areas they are believed to have Diogmites angustipennis Loew destroyed an appreciable amount 1866: 23 of the white grub population. The pupae move freely and mi- Synonymy.—None (320, p. grate to the soil surface imme- 371). diately prior to adult emergence. Recorded distribution.—Colo- The period within the pupal case rado, Kansas, and Illinois {183, is dependent on the species; Ef- p. 13; 320, p. 371). (Map 149.) feria interrupta (Macquart) is Host species.—Aulocara elliotti reported to have a pupal period of (Thomas) {183, p. 13). from 20 to 26 days. Adults live Incidence of prédation.—None approximately 3 months during located. the summer, during which they Life history and habits.—SO: satisfy a ravenous appetite. Lar- Eggs, larvae, and pupae unknown. vae do not feed on such a wide Adults not given. LOD : Unknown. variety of insects, but they too OWS: Not given. G/Y: One. SA: have a voracious appetite. Life Mostly adults. N/H: Unknown. cycles of the more common spe- MOA: Prey in flight. AF: Body cies range from 1 to 3 years. contents of insects (hemolymph). There are only two short pe- HAB : Similar to the prey taken riods during the asilid's develop- when adults, in the soil as larvae, ment when it is not predaceous, the egg and pupal stages. During the remainder of its lifetime it is a relentless hunter. Its numbers never seem great during the adult stage, possibly because of its en- forcement of territorial rights. Any stray male asilid or some- times female asilids that wander into an area dominated by an- other are either chased or cap- tured. On several occasions male asilids have been noted draining the body contents of the female MAP 149.—Recorded distribution of during the act of mating. Diogmites angustipennis Loew. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 217 and in or near the egg beds of the Recorded distribution.—East- host. (183, p. 13; 323, p. 260) ern North America to central Description.—Eggs, larvae, and Saskatchewan, central Montana, pupae unknown. Adult flies are 21 and south through Arizona (320, mm. long and gray ocher. On the p. 393). (Map 150.) thorax are three obscure dark- Host species.—Egg pods, olive stripes. The head is red pol- nymphs, and adults of Melanoplus linose and the setae of the occiput spretus (Walsh) and other flying and vertical bristles are dark to insects (123, pp. 450, Â80; 264, black. The first two antennal pp. 303-304; 331, p. 97). joints bear black pile. The mystax Incidence of prédation.—None is a straw color, the palpi are red located. or nearly so, and the pile is dark Life history and habits.—SO: and rough or bristly. The dorsal Eggs July to August. Larvae late gray thorax is sprinkled with a fall until spring. Pupae spring. pale ocher, the pile is very short, Adults May to August. LOD : Un- and the setae are completely black. known. OWS: Larvae. G/Y: One Abdomen is dark to dark olive generation may last from 1 to 3 with the angle of the posterior years. SA: All. N/Y: Unknown. lateral margin a pale pollen color. MOA: Larvae are predaceous on Feet and apices of the tarsi and egg pods or soft-bodied grubs; the setae are all black. Legs are adults attack jumping or flying narrow and bear microscopic prey. AF: Hemolymph of prey. hairs. Wings are narrow and hy- HAB : For adults, similar to that aline with a pale yellow tinge ; the of the grasshopper hosts ; for lar- posterior margin is gray or dark vae, in or near egg beds of the gray. (30, pp. 82, 100; 31, p. host. 230; 158, p. 33; 205, p. 23) Females lay their eggs in clus- Discussion.—Although only grasshopper species are recorded as hosts, other insects are be- lieved to be included. Efferia aestuans (Linnaeus) (Asilus) 1763: 413 Synonymy.—Asilus aestuans Linnaeus, A. macrolabes Wiede- mann, A. nig er Wiedemann, Erax bastardi Macquart, E. femaratus Macquart, E. incisuallis Macquart, E. tibialis Macquart, E. aestuans Hine, E. niger (Wiedemann) (A, MAP 150.—Recorded distribution of pp. 275-276; 320, p. 393). Efferia aestuans (Linnaeus). 218 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE ters of from 40 to 90 in the seed Adult males are 19 mm. and heads of Brunella, Achillea, Rud- females 21 mm. long. Mystax is beckia, and Verbascum. Adults black with white hairs below. Face alight on upright stationary ob- and front are covered with a yel- jects such as buildings, fence- lowish-gray dust. Antennae are posts, and trees and have the black and covered on the first two unusual habit of chasing and land- joints with black and yellowish- ing on large moving animals such white short hair. The hair on the as cows and man. (26^, pp. 303- front is chiefly black. An abun- 30A \ 323, p. 260) dant black beard is mixed with Description.—Eggs unknown. silky white. On the dorsum of the Larvae are 27 mm. long and trans- thorax are two well-marked black lucent to yellowish white; they stripes of nearly equal width, taper toward both ends. There are which are separated by a pollinose 12 noticeable segments; the an- linear stripe that ends abruptly terior three and the posterior just beyond the suture. The tho- three are tapered gradually and rax is covered with a grayish- the middle six are of equal width. brown to gray pollen and black The head is dark brown, pointed hair of moderate length in the in shape, retractile, and divided front. Hair on the dorsum of the at the tip into two mandibular scutellum is white with thick black points with two uniform append- bristles. The abdomen is black and ages. The anal segment has two slightly shining. Venter and a depressed longitudinal lines above rather broad lateral margin on and is ridged on the anterior edge. each segment are covered with a (26i, pp. 303-30JÍ) thick gray pollinose. In the female Pupae are stout and honey yel- the fifth, sixth, and seventh seg- low. Legs and wings are en- ments are silvery white, whereas sheathed although separated from the large black hypogium supports the abdomen. The head has two black hair. Legs are black except large brown spines in front, a the yellowish-red tibia. Bristles laterial set of three small spines, are black and the hair is chiefly and large dark eyes. There are two white. The tip of the tibia and the small rounded dorsal projections first two joints of the metatarsi and a set of two small lateral support a golden-yellow pile. spines just behind the head on the Wings are hyaline and veins and thorax. Each segment of the ab- subcostal cell are brown. {30, domen is ridged in the middle pp. 86,105-108 ; HO, pp. 105,107, with a ring of brown blunt thorns 12U-125\ 202, p. U13\ 210, pp. that slope posteriorly. The anal 117-118 \ 367, pp. 97-98 \ 375, pp. segment has a few stouter spines. 6Jf, 71-72) {261^, pp. 303-30If) Discussion.—The larvae and ARTHROPOD AND NEMATODE PARASITES IN AMERICA 219 adults are predaceous on grass- hoppers and their eggs.

Efíeria alhibarbis (Macquart) {Erax) 1838: 118 Synonymy.—Erax alhibarbis Macquart, E. barbattis Hine, E. barbatus Fabricius, E. ciner- ascens Bellardi, E. furax Willis- ton, E. pumilis Walker, E. tricolor Bellardi U, p. 275; 320, p. 393). Recorded distribution.—South- MAP 151.—Recorded distribution of ern Canada and the United States Efferia alhibarbis (Macquart). {320, p. 393). (Map 151.) Host species.—Melanopltts dif- can be found near the ocelli. The ferentialis (Thomas) (123, p. thorax contains a dense gray pol- Í80; 155). linose and the dorsum is often Incidence of prédation.—None brownish. A weak median stripe located. can be seen on the thorax, and the Life history and habits.—SO: front of the dorsum is covered Eggs laid in fall. Larvae late fall with very short black hair, which to early spring. Pupae spring. becomes sparse posteriorly. Pile Adults: One of the first species on the pleura is white and very of asilids to appear in the spring. thin. Abdominal segments 1 to 5 In California at elevations up to and 8 of the male are gray, 6,000 feet it is seen between April whereas segments 6 and 7 are and October. LOD: Unknown. white. Segment 2 has small brown OWS : Larvae. G/Y : One. SA : All. basal spots and sometimes the N/H: Unknown. MOA: Larvae female has a blackish spot on each attack egg pods and soft-bodied side. Each remaining segment larvae in the soil. Adults attack contains a pair of large quadrate flying insects. AF: Hemolymph. spots rather narrowly separated. HAB : Similar to that of the prey. Legs are black with white pile This is a very common species and white with black macro- (155) chaetae. Front and middle tibia Description.—Eggs, larvae, and except the tip and the hindtibia pupae unknown. Adults are 16 to except the distal third or half are 19 mm. long, and the basic colora- reddish or yellow. The front and tion of the head and scutellum is middle tarsi are yellowish red, covered with white pile and hair. and the hindtarsi and hindfemora Occasionally a few black bristles usually blackish. Wings are hya- 220 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE line with brown veins. (HO, pp. 105, 107; 210, p. 118; 367, pp. 108-109)

Efferia arida (Williston) {Erax) 1893: 254 Synonymy.—Erax aridus Wil- liston U, p. 276; 320, p. 398). Recorded distribution.—Cali- fornia, Nevada, and Utah {320, p. 393). (Map 152.) Host species.—Grasshoppers MAP 152.—Recorded distribution of {123, p. Jf80; 367, pp. 1U3-1UU; Efferia arida (Williston). 377, p. 25U). Incidence of prédation.—None located. some black above the palpi. An- Life history and habits.—SO: tennae are black and support Eggs unknown. Larvae fall until blackish hair below with white spring. Pupae early spring. Adults above. The mesonotum is black March and April in California, with gray pollinose and a brown Nevada, and Utah. LOD: Un- central stripe in the intermediate known. OWS : Larvae. G/Y : One. area. Abdomen is black. Segments SA: Nymphs (grasshoppers). 1 to 5 of the male are thinly N/H: Unknown. MOA: Attacks brownish pollinose, whereas the jumping insects. AF: Hemo- sixth and seventh are a white lymph. HAB: Similar to that of pollinose. Abdominal segments 2 the prey. {367, pp. H3-1U; to 6 of the female are grayish pol- 377, p. 25A) linose. Hair around the forceps of Description.—Eggs, larvae, and the male is white with the ventral pupae unknown. fringe short and brown and with Adults are 22 to 26 mm. long sparse white hairs on the outer and basically black. Mystax of the margin. Legs are black with large male is white with black hairs white bristles. There are some intermixed on the sides and be- black bristles on the midfemora, low. The head is covered with hindfemora, and tibiae with some gray pollinose. Hair on the frons black hairs above and below on the is mixed black and white, whereas femora. Claws are black and the occipital bristles are yellowish brownish basally. Pulvilli and intermixed with some black. The empodium are brown. {HO, pp. beard and hair of the proboscis 105, 107, 117; 367, pp. U3-1U; and of the palpi are white with 377, p. 25A) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 221

Efferia bicaudata (Hîne) Egg pods and generally flying or {Erax) 1919: 138 adult stages. N/H: Unknown. Synonymy.—Erax hicaudatus MOA: Generally flying insects. Hine {320, p. 394). AF: Insect hemolymph. HAB: Recorded distribution.—Al- Similar to that of the prey. berta, Montana, Wyoming, Colo- La vigne (181) gave a good de- rado, New Mexico, and Texas scription of the general behavior (140, pp. 106, 138-139; 181, pp. of E. bicaudata. He reported that 341-3U; 320, p. 394) (Map flies usually rest on the vegetation 153.) of Carex, Stipa, Bromus, Opuntia, Host species.—Ageneotettix de- and Cleome about 5 inches to 2 orum (Scudder), Amphitornus feet above the ground. They sel- coloradus (Thomas), Cordillacris dom remain in one spot for more crenulata (Bruner), Melanoplus than 2 to 3 minutes except during infantilis (Scudder), M. occiden- cold or cloudy conditions and talis (Thomas), and species in make short flights ranging from seven orders of insects (181, pp. several to 20 feet in search of 341-344; 389). food. These forage flights usually Incidence of prédation.—Not occur 12 inches above the soil, given. depending on the height of the Life history and habits.—SO: vegetation. Prey are nearly al- Eggs summer to fall. Larvae fall ways taken in the air. Both prey until early spring. Pupae un- and predator fall to the earth knovs^n. Adults betv^een the last where the prey continues to part of July and late September. struggle for 30 to 60 seconds be- LOD: Unknown. OWS: Larvae. fore becoming paralyzed. The G/Y: Not more than one. SA: prey is then held with all six legs while the proboscis is inserted either into the frons or at the 111-, ,v< base of the wings. Feeding is generally accomplished on a grass stalk or other vegetation by hold- ing the victim up so that the fluids drain into one spot. The proboscis is inserted in various locations along the body to obtain all the internal juices. Time spent feeding on grasshoppers averages 59 to 89 minutes. Oviposition occurs by insertion MAP 153.—Recorded distribution of of the ovipositor into the sand and Efferia bicaudata (Hine). depositing two to three eggs per 222 TECHNICAL BULLETIN NO. 14^0, U. S. DEPT. OF AGRICULTURE location. After about li/^ minutes 138-139 ; 158, p. JÍ8 ; 367, pp. 165- the female moves to a new area 168) (Fig. 18.) 1 to 2 feet away and oviposits Discussion.—Although preda- again. Eggs are generally ce- ceous on insects, E. hicaudata is mented together in layers. {157, also the prey of three other asilids pp. 70-7^; 181, pp.Sil-SU; 389) —Diogmites angustipennis Loew, Description.—Eggs are about Proctacanthus milbertii Mac- 1.23 mm. long by 0.42 mm. wide, quart, and Efferia pallidulus creamy white when laid, and (Hine), which are themselves semioval {181, pp. 3Í1-3H). Lar- predators of grasshoppers. E. hi- vae and pupae unknown. caudata is thought to reduce Adult males are 24 mm. long grasshoppers when their popula- and females 25 mm. Both mystax tions are low and other flying and beard are white and the palpi insects limited. are black and bear black hair. The thorax is covered with white Efferia cana (Hine) hairs and bristles, whereas the {Erax) 1916: 22 scutellum has black hair on the Synonymy.—Erax canus Hine disk and numerous white bristles, {320, p. 39JÍ). which are irregular on the mar- Recorded distribution.—Brit- gin. The abdomen is dark and ish Columbia, California, Utah, partly gray poUinose with much Colorado, and Arizona {320, p. white hair and some short black 39A\ 367, pp. 196-197). (Map hair dorsally. Abdominal seg- 154.) ments 1 to 5 are brownish and Host species.—Small bees, flies, segments 6 and 7 are silvery. and grasshoppers {123, p. ^80; Hypopygium is black, reddish in 336, p. 33). part, medium sized, and covered Incidence of prédation.—None mostly with white hair. The ovi- located. positor of the female is about as Life history and habits.—SO long as abdominal segments 6 and and LOD: Unknown. OWS and 7. As viewed from the side it is G/Y : Not given. SA : Nymphs of narrowed abruptly before the grasshoppers, adult bees, and flies. apex with a backward projecting N/H: Unknown. MOA: Jumping and flying prey. AF : Insect hemo- point, which from the dorsal view lymph. HAB: Not given. {336, is seen to be furcated. The legs p. 33) are largely black with numerous Description.—Eggs, larvae, and white hairs and black bristles. pupae unknown. Adults are about The tibia is reddish on the basal 24 mm. long and grayish white third, especially the anterior and (produced by white pollen and the middle pairs. Wings are hyaline. covering hair). Black hair and {30, pp. 86, 107 \ HO, pp. 106, bristles can be found on the dor- ARTHROPOD AND NEMATODE PARASITES IN AMERICA 223

PN-2736 FIGURE 18.—Adult of Efferia bicaudata (Hine). sum of the thorax and legs. Both front. On the black palpi are yel- the mystax and beard are pale low hair and bristles. The meso- yellow with black bristles on the notum is black with grayish- yellow pollinose and black hairs. The dividing central stripe and intermediate spots are brown, and the scutellum is grayish-yellow pollinose covered with dense white hairs. The abdomen is black with a uniform grayish white. Long white hair on the dorsum is parted at the middle and directed for- ward on segments 2, 3, and 4. Hypopygium is black, of medium size, and covered with white hair. The female lacks long hair on MAP 154.—Recorded distribution of Efferia cana (Hine). segments 2, 3, and 4. Ovipositor 224 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE is shiny black and slightly longer known. OWS: Larvae. G/Y: Un- than abdominal segments 6 and 7 known. SA: Eggs. N/H: Un- "- combined. Legs are black with the known. MOA: Larvae attack " basal third of each tibia reddish. grasshopper egg pods. AF : Insect Wings are hyaline with brownish hemolymph. HAB: Unknown. veins. (,139, p. 22; HO, vv- 106, This species is generally associ- s 1A5-H6; 367, pp. 196-197) ated with tree trunks. {264) Description.—Eggs, larvae, and Efíeria femorata (Macquart) pupae unknown. Adult flies are {Erax) 1838: 115 26 to 30 mm. long. The sexes are , Synonymy.—Erax femoratus similar in appearance. Black and Macquart (Í, p. 276 ; 320, p. 39^.). white intermixed hairs make up - Recorded distribution.—North the mystax and the beard is pale. Carolina, South Carolina, Louisi- Palpi are black. Very short black ana, Georgia, Florida, and east- hair is on the anterior part of the ern Alabama {320, p. 39^). (Map mesothorax and several strong 155.) black bristles are above the roots Host species.—Melanoplus of the wings and just anterior of ^ spretus (Walsh) {123, pp. 4-50, the scutellum. On the scutellum ^ i.80; 26A). are fine white hairs and several Incidence of prédation.—Very black bristles on the disk. The low, since this species is appar- first four segments of the male ently rare {123, pp. 450, 480). abdomen are gray on the sides Life history and habits.—SO: and narrow on the hind margin. Eggs unknown. Larvae late fall Segments 5, 6, and 7 are silvery. to early spring. Pupae unknown. Hypopygium is black and contains Adults April to August in the black hair. Abdomen of the female States listed above. LOD: Un- is gray on the sides and narrow ■ near the posterior borders. The ovipositor is nearly as long as the last four abdominal segments. Femora are black and tibia and tarsi are dark red. Wings are • hyaline with a light-brown tinge ^ apically. Veins and subcostal cell are brown. {30, p. 86; 140, pp. 105, 123-124; 210, p. 115, 232; 867, p. 102) « Efferia harveyi (Hine) {Erax) 1919: 115 Synonymy.—Erax harveyi MAP 155.—Recorded distribution of Efferia femorata (Macquart). Hine {320, p. 394). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 225

1^ Recorded distribution.—Brit- ish Columbia and Pacific Coast States (320, p. 39i). (Map 156.) Host species.—Melanoplus dif- fer entialis (Thomas) and various small bees and nies (123, p. i80; 336, p. 33). Incidence of prédation.—Not given. Life history and habits.—SO: Eggs summer to fall. Larvae fall. Yxl I \ 'Í i:^ Pupae spring. Adults August and September. LOD : Unknown. MAP 156.—Recorded distribution of OWS : Probably larvae. G/Y: Un- Efferia harveyi (Hine). known. SA: Nymphal grass- hoppers, adult bees, and flies. are wedge shaped at the tip when N/H: Unknown. MOA: Jumping viewed from the side and furcate and flying insects. AF: Insect apically when viewed from the hemolymph. HAB: Unknown. Is top. (UO, pp. 105,115-116 ; 367, thought to be similar to other p. U7) asilid species of the same size. (336, p. 33) Efferia helenae (Bromley) Description.—Eggs, larvae, and (Erax) 1951: 30 pupae unknown. Adult males are Synonymy.—Erax helenae 18 mm. long and females 21 mm. Bromley (320, p. 394.). Anterior part of the dorsal meso- Recorded distribution.—Colo- thorax of the female is covered rado at an elevation between 4,300 with black hair, whereas the re- and 4,500 feet (183, pp. 28, 30; mainder of the thorax and the 320, p. 39i). (Map 157.) abdomen are covered with pale Host species.—Amphitornus hair; the general color is dark, coloradus (Thomas), Ageneotet- thin, gray pollinose. Mystax, tix deorum (Scudder), Cordilla- beard, and ocular and occipito- cris crenulata (Bruner), C occi- orbital bristles are pale yellow. pitalis (Thomas), Melanoplus Palpi are black and yellow and angustipennis (Dodge), M. occi- are covered with hair. Scutellum dentalis (Thomas), Opeia obscura is covered with numerous pale- (Thomas), Philibostroma qvxtdri- yellow marginal bristles. Legs are maculatum (Thomas), Psoloessa faded reddish and covered with delicatula (Scudder), and Trachy- a pale pile. Wings are hyaline. rhachys kiowa (Thomas) (183, The male is smaller than the fe- pp. 28, 30). male but is similar in color. Sixth Incidence of prédation.—None and seventh abdominal segments located. 226 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE bear black spots that are bare of the surrounding yellowish pollen. Segments 6, 7, and 8 are a silvery pruinose. Hypopygium of the male and the ovipositor of the female are a dark reddish brown. Legs ' are black as are the bristles. • Tibiae on the posterior sides ex- cept the tips are pale reddish brown. Claws are black and the ■ pulvilli yellow. Wings are hyaline and the haltères yellowish brown. =- MAP 157.—Recorded distribution of {33, p. 30; 367, p. 170) Efferia helenae (Bromley). Discussion.—Although nearly a dozen species are recorded as Life history and habits.—SO: prey, this predator is only re- Eggs late summer to fall. Larvae ported from Colorado. fall. Pupae spring. Adults August and September. LOD: Unknown. Efferia interrupta (Macquart) OWS: Probably as larvae. G/Y: {Asilus) 1834: 310 Not given. SA: Third-instar nymphs to adults. N/H: Un- Synonymy.—Asilv^ interrupta known. MOA : Jumping and flying Macquart, Erax lateralis Mac- prey. AF: Insect hemolymph. quart, E. ambiguus Macquart, A. HAB : Similar to that of the prey. maculatus Macquart, Eristicus (lU, pp. 28, 30) villosus Bellard (320, p. 39A). Description.—Eggs, larvae, and Recorded distribution.—United pupae unknown. Adults are 22 to States. Also Mexico. (320, p. 31 mm. long and pale-yellowish 3H). (Map 158.) pollinose to pruinose. Tip of the Host species.—Grasshoppers female ovipositor is bifurcate and (158, p. A8; 323, pp. 259-260', ' the mesonotum and scutellum are 331, p. 97). strongly bristled. Hair of the Incidence of prédation.—None head is white except a few black located. bristles on the vertex and on the Life history and habits.—SO: distal part of the palpi. White Eggs summer to fall. Larvae fall. hair intermixed with some black Pupae spring. Adults from first of covers the mesothorax, and most May in Texas, June in Georgia ^ of the bristles of the mesonotum and Florida, and July in Colorado and scutellum are whitish or yel- and Utah until fall. LOD: Un- lowish. Two median lines on the known. OWS, G/Y, and SA: Not dorsum are free from the yellow given. N/H: Unknown. MOA: pollen. Abdominal tergites 1 to 5 Not given. AF: Hemolymph. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 227

tibiae. Wings are reddish hyaline. (30, pp. 86, 107; HO, pp. 107, 153; 158, p. 48; 209, p. 310) Efferia pallidula (Hine) (Erax) 1911: 309 Synonymy.—Erax pallidula Hine (320, p. 395). Recorded distribution.—^Wy- oming, Colorado, and New Mex- ico (320, p. 395). (Map 159.) Host species.—Ageneotettix de- MAP 158.—Recorded distribution of orum (Scudder), Amphitornus Efferiainterrupta (Macquart). coloradus (Thomas), Aulocara el- liotti (Thomas), Cordillacris cren- HAB: Not given. {55, p. 373; ulata (Bruner), C. occipitalis 323, pp. 259-260) (Bruner), Hesperotettix viridis Description.—Eggs, larvae, and (Thomas), Melanoplu^ angv^ti- pupae unknown. Adult flies are pennis (Dodge), and M. infantilis 22 to 27 mm. long and brownish. Scudder (183, pp. 28-30). The mystax is pale and inter- Incidence of prédation.—None mixed with a few black bristles located. above. Beard is pale. Palpi sup- Life history and habits.—SO port a few black bristles. Thorax and LOD: Unknown. OWS and is largely gray pollinose on which G/Y: Not given. SA: Usually is a wide dark middorsal stripe. adult grasshoppers. N/H: Un- On the margin of the scutellum known. MOA: Generally flying are many black bristles. Abdomen insects. AF: Hemolymph. HAB: is nearly black and covered with yellowish gray pollinose spots on the posterior region of the cor- ners of each segment; spots do not meet on the midsegmental part of abdominal segments 1 to 4. Segment 5 is all pollinose on the hind margin and segments 6 and 7 of the male are completely pol- linose and in the female bear small black triangles with the base posteriorly. The male geni- talia are furcate. Legs are mostly reddish brown except the nearly MAP 159.—Recorded distribution of black underside of the femora and Efferia pallidula (Hine). 228 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Similar to that of the host. (18S, pp. 28-30) Description.—Eggs, larvae, and pupae unknown. Adults are 18 to 28 mm. long and pale. Mystax is a very pale yellowish and the beard is white. Palpi are black and support white hair. Occipital, orbital, and ocellar bristles are usually black. Dorsal part of the n - -^^ ~^- thorax is pale yellow gray and supports short black hair ante- riorly and black and white bristles MAP 160.—Recorded distribution of posteriorly. On the scutellum are Efferia rapax (Osten Sacken). pale hairs. Legs are black except the base of the tibia is pale. Male (Thomas) (123, p. i80; 336, p. abdomen is silvery white. The first 33). four segments contain long white Incidence of prédation.—None hair that is parted at the middle located. and directed outward on segments Life history and habits.—SO 2, 3, and 4. Segments 5, 6, and 7 and LOD: Unknown. OWS and are without long hair. Hypopy- G/Y : Not given. SA : Nymphs and gium of the male is small, black, adults of grasshoppers and adults and narrow toward the apex of bees and flies. N/H : Unknown. where it appears rounded when MOA : Jumping and flying insects. viewed from the lateral aspect. AF: Insect hemolymph. HAB: Wings are hyaline. {30, p. 86; Similar to that of the prey. (336, 138, pp. 309-310; HO, pp. 107, p. 33) lU) Description.—Eggs, larvae, and pupae unknown. Efferia rapax (Osten Sacken) Adults are 19 to 25 mm. long (Erax) 1887: 201 and dark brown basically. Face Synonymy.—Erax rapax and frons are covered with yellow Osten Sacken (4, p. 277; 320, p. pollinose and the occiput with 395). white pollinose. Palpi are black Recorded distribution.—From and support dull-yellow hairs. British Columbia throughout Antennae are also black except southwestern United States to the basal joints are covered with Mexico (320, p. 395). (Map short yellowish hairs. Beard is 160.) pale yellow. Thorax is brownish Host species.—"Grasshoppers, yellow and bears a well-marked, flies and small bees," including dark-brown dorsocentral stripe. Melanoplus differentialis Hair is black. Mesonotum is ARTHROPOD AND NEMATODE PARASITES IN AMERICA 229 black and the scutellum yellowish Host species.—Grasshoppers, gray. small bees, and nies {123, p. A80 ; The abdomen of the male is 336, p. 33). black at the base, but it is covered Incidence of prédation.—Not with yellowish-gray pollen and given. yellowish hairs on the sides. Seg- Life history and habits.—SO: ments 4, 5, and 6 and part of the Eggs late spring. Larvae summer third are silvery and contain sil- to early spring. Pupae unknown. very hairs parted in the middle so Adults April and May in Texas that they point outward. The and New Mexico. LOD : Unknown. seventh segment is blackish and OWS and G/Y: Not given. SA: contains a white stripe on the Nymphs and adults of grass- sides. Forceps of the male are hoppers and adults of bees and large, black, and covered with flies. N/H: Unknown. MOA: black and yellowish hairs. Jumping and flying insects. AF: Abdomen of the female is a Hemolymph. HAB : Similar to uniform golden, yellowish brown that of the prey. {336, p. 33) with shifting darker shadows on Description.—Eggs, larvae, and the side. On each side is a distinct, pupae unknown. Adults are about rather broad, dull-gray stripe that 22 mm. long and the sexes are extends from the first to the similar in coloration. Most of the seventh abdominal segment. Ven- head bristles are yellowish, ter is gray and the ovipositor is whereas the hair on the thorax equal in length to the three pre- (dorsum) and scutellum is black. ceding segments. Legs are black The flrst three abdominal seg- and covered with soft yellow hair. ments are mostly black in the The longer bristles on the hind male and thinly pollinose laterally. side of the tibia of the female are Hair is black above and pale also yellow. Wings are hyaline and very slightly tinged with yellowish brown, especially on the distal half. {138, pp. 307-308; HO, pp. 106, 1U2; 158, p. Í8; 367, pp. 216-217) Efferia snowi (Hine) {Erax) 1919: 116 Synonymy.—Erax snowi Hine {320, p. 395). Recorded distribution.—Brit- ish Columbia, Colorado, Kansas, New Mexico, and Texas {320, p. MAP 161.—Recorded distribution of 395). (Map 161.) Efferia snowi (Hine). 230 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE yellow on the sides. Abdomen of the female is entirely yellowish- gray pollinose and supports pale hair. Male hypopygium is black as is the ovipositor of the female. Legs are mostly black and support abundant long black hair that is intermixed with black and pale bristles. The basal half of each front and middle tibia and the basal fourth of the hindtibia are reddish. Wings are faintly yellow- ish and hyaline. (30,pp.87,108; MAP 162.—Recorded distribution of lAO, pp. 105, 116; 158, pp. il-49; Efferia staminea (Williston). 367, p. U9) to 21 mm. long and basically Efferia staminea (Williston) black. Hair and bristles on the (Erax) 1885: 68 head are yellowish and the beard Synonymy.—Erax stramineus is white. Face and frons are yel- Williston, E. staminea Williston, lowish white and the occiput is E. tanneri Bromley U, p. 277; grayish pollinose. Antennae are 320, p. 395). black with yellow hairs. Mesono- Recorded distribution.—Mon- tum is black and yellowish-brown tana, Wyoming, and Colorado pollinose ; a divided central stripe {320, p. 395; 367, pp. 217-218). and intermediate spots are brown. (Map 162.) Mesonotal bristles of the female Host species.—Ageneotettix de- are yellowish white. Scutellum is orum (Scudder), Aulocara elliotti yellowish-gray pollinose. The (Thomas), and Cordillacris occipi- hairs of the male are black and talis (Thomas) (183, pp. 28-30). those of the female are white with Incidence of prédation.—None a few black intermixed. Abdomen located. of the male is black and the fe- Life history and habits.—SO: male a gray pollinose. The venter Eggs summer. Larvae fall. Pupae of the male is brownish pollinose spring. Adults June, July, and basally becoming gray apically; August. LOD: Unknown. OWS that of the female is yellowish- and G/Y : Not given. SA : Adults. gray pollinose. Hairs of both sexes N/H: Unknown. MOA: Flying are white and bristly. Ovipositor insects. AF: Hemolymph. HAB: of the female is black. Wings are Grassland, near foothills, and hyaline and the veins light brown. mountains. (367, pp. 217-218) (140, pp. 107, IW-IU; 158, p. Description.—Eggs, larvae, and A8; 367, pp. 217-218; 375, pp. pupae unknown. Adult flies are 18 53-76) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 231 Machimus occidentalis (Hine) ment (arista not included) is (Asilus) 1909: 147 slightly longer than the first two Synonymy.—Asilus occiden- segments combined. Aristae are talis Hine (320, p. 396). barely as long as the third anten- Recorded distribution.—Brit- nal segment. Median dark mark- ish Columbia, Washington, Ore- ings on the thorax are weak. The gon, California, Nevada, and abdomen is dark; the posterior Wyoming (320, p. 396). (Map margins of each segment are gray. 163.) On the posterior margin of the Host species.—Grasshoppers venter of the eighth abdominal (183, pp. 28-30; 331, p. 99). segment in the male is a dense Incidence of prédation.—None cluster of long bristly hairs. Geni- located. talia of the male are bent upward Life history and habits.—SO, near the middle. (137, pp. 147- LOD, OWS, and G/Y: Unknown. U8) SA: Not given. N/H: Unknown. Ospriocerus latipennis (Loew) MOA: Not given. AF: Grasshop- per hemolymph. HAB : Not lo- (Stenopogon) 1866: 28 cated. (9A, p. 77U) Synonymy.—Stenopogon con- Description.—Eggs, larvae, and sanguineu^ Loew, S. latipennis pupae unknown. Adults are 14 to Loew (4, p. 256; 320, p. 381). 18 mm. long and generally gray Recorded distribution.—From with the thorax often a yellowish eastern Wyoming and South Da- gray. Mystax is largely black and kota south through New Mexico intermixed with yellowish hairs and Texas (320, p. 381). (Map below. The face is covered with 164.) yellowish dust. Beard is white. Host species.—Melanoplus Antennae are black ; the third seg- spretus (Walsh) (26A, p. 317). Incidence of prédation.—None located. Life history and habits.—SO, LOD, OWS, and G/Y : Unknown. SA: Not given. N/H and MOA: Unknown. AF: Grasshopper he- molymph. HAB : Unknown. (26^, p. 317) Description.—Eggs, larvae, and pupae unknown. Adults are nearly 20 mm. long and generally ocher. The head, antennae, prothorax, and setae of the dorsal thorax and MAP 163.—Recorded distribution of metanotum are black; the thorax Machimus occidentalis (Hine). is usually ocher. The three stripes 232 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE Life history and habits,—SO, LOD, OWS, and G/Y: Unknown. SA: Not given. N/H and MOA: Unknown. AF: Grasshopper he- molymph. HAB: Unknown. Description.—Eggs, larvae, and pupae unknown. Adult males are approximately 36 mm. long and dark to black except the abdomen is gray and the feet are pale brown. Antennae are black. Bris- tles are white and the mystax and MAP 164.—Recorded distribution of vibrissae gray. That section of Ospriocerus latipennis (Loew). the head between the antennae, eyes, and mouth is near ocher, the of the dorsal thorax are black frons is a lighter color, and on it with a compressed yellow pollen are dark-black setae. White pile covering. The abdomen is all on the occiput is intermixed with ocher with the end of the segment some black bristles on either side. margins sometimes black. Hind- The thoracic stripe is dark and coxae are black, but the feet are surounded by yellow; the scutel- all ocher. Setae and pile are yel- lum and pleura are white. Abdo- low. The forward section of the men is gray to dark gray and the wings is dark and the costal veins venter lighter gray. Wings are are near ocher. (l-í, pp. 205- clear to a slight yellowish tinge. 206; 30, p. 97; 32, p. 295; 158, p. Haltères are dark. (30, pp. 85, 3i; 205, p.28) 106; 365, p.183)

Proctacanthus longus (Wiedemann) (Asilus) 1821: 183 Synonymy.—Asilus longus Wiedemann, Proctacanthus long- ulvs Wiedemann H, p. 27A; 320, p. 399). Recorded distribution.—Texas, Georgia, and Florida (320, p. 399). (Map 165.) Host species.—"Grasshoppers" (323, p. 260). Incidence of prédation.—Not MAP 165.—Recorded distribution of given. Proctacanthus longus (Wiedemann). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 233

Proctacanthus milbertii Macquart 1838: 124 Synonymy.—Asilus agrión Jaennicke, A. missouriensis Riley. U, p. 274; 320, p. 399). Recorded distribution.—South- ern Canada and the United States (320, p. 399). (Map 166.) Host species.—Ageneotettix de- orum (Scudder), Amphitomus coloradus (Thomas), Aulocara MAP 166.—Recorded distribution of elliotti (Thomas), Hesperotettix Proctacanthus milbertii Macquart. viridis (Thomas), Melanoplus foe- dus Bruner, M. occidentalis red ground color is covered by (Thomas), M. sanguinipes (Fab- dense gray pollen. Sparse white ricius), bristles make up the mystax. An- (Thomas), Trimerotropis campe- tennae are generally black; the stris McNeill, and butterflies, joints of the third and sometimes honey bees, bumble bees, wild the tip of the second joints are bees, wasps, and the beetle Ci- red. Beard is white and moder- cindela purpurea Olivier (123, p. ately abundant. Two median black Jf80; 183, pp. 28-30; 26Í, p. 317; stripes on the thorax are sur- 326, p. 3; 331, p. 99 ; 336, p. 33). rounded by a reddish ground Incidence of prédation.—Not color. The fly is completely cov- given. ered with a nearly white pollen. Life history and habits.—SO : Abdomen is black but densely cov- Eggs, larvae, and pupae unknown. ered with nearly white pollen, Adults not given. LOD : Unknown. which sometimes appears brown- OWS and G/Y: Not given. SA: ish. Ovipositor of the female is Nymphs and adults. N/H: Un- shining and the hypopygium of known. MOA : Attacks and catches the male red. At times the ground flying insects and transports them color of the posterior margins of in the air where the body contents the segments may be red. Legs are sucked out. AF: Insect he- are usually dark red and the fe- molymph. HAB: Similar to that mora broadly black in front. Tarsi of the host (generally dry cli- are often blackish. (30, pp. 85, mate). (IBS, pp. 28-30; 26Í., p. 106; 66, pp. 3Jf-57; 158, p. 51; 317; 336, p. 33) 210, p. 12i; 375, pp. 73-7Jf) Description.—Eggs, larvae, and Discussion.—It is considered pupae unknown. Adults are 30 to an important predator of grass- 35 mm. long. Males and females hoppers in British Columbia (336, are similar in appearance. Facial p. 33). 234 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Proctacanthus rodecki James covered with densely grayish pol- 1933: 2-3 len. In the male the ground color Synonymy.—None {320, p. of the thorax and the basal ab- 399). dominal segments are completely Recorded distribution.—Wy- covered. Palpi and proboscis are oming, Colorado, and Kansas black ; palpi are covered with long (320, p. 399). (Map 167.) hairlike bristles, which are pale Host species.—Trachyrhachys to almost white. The first two kiowa (Thomas) and "grasshop- antennal segments are red and the pers" (183, pp. 28-30). third segment and the style except Incidence of prédation.—None the apex are black. The thorax is located. covered with a short, erect, bris- Life history and habits.—SO: tlelike pile, which is mostly black Eggs summer to fall. Larvae fall. but is intermixed with a few pale Pupae spring. Adults July. LOD: hairs at the posterior angles. Bris- Unknown. OWS and G/Y: Not tles of the thorax and scutellum given. SA: Adults. N/H: Un- are white as is the pile of bristles known. MOA: Not given. AF: of the pleura, coxae, and first five Hemolymph. HAB: Similar to abdominal segments. Bristles of that of the hosts. (183, pp. 28- the femora, tibiae, and tarsi are 30) black. The pile of the tibiae and Description.—Eggs, larvae, and tarsi is white intermixed with pupae unknown. Adult flies are some black anteriorly, at the mid- 36 to 42 mm. long and reddish dle part of the hindtibiae, and all brown to black. The ground color tarsi. Anterior femora are red is apparent on the face, legs, and and the posterior femora reddish apical segments of the abdomen black. Genitalia of the male are in the female, but otherwise it is red and of the female black. Ter- minal segments of the abdomen and the genitalia are covered with gray pilosity in the male and black pile in the female except just anterior to the ovipositor. Female genitalia are encircled with distinct black spines. (156, pp. 1-3) Promachus vertebratus (Say) (Asilus) 1823: 47 Synonymy.—Asilus vertebra- tus Say, Promachius vertebratus

MAP 167.—Recorded distribution of Schiner, Trapanea vertebratus Proctacanthus rodecki James. Macquart (4, p. 280; 320, p. WO). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 235

Recorded distribution.—From Wyoming, Colorado, and New Mexico into the Atlantic Coast States (320, p. iOO). (Map 168.) Host species.—Melanoplus sanguinipes (Fabricius) and spe- cies of Coleóptera, Hymenoptera, and Hemiptera (323, p. 260 ; 330, p. 182; 331, p. 99; 375, pp. 60, 62). Incidence of prédation.—None located. Life history and habits.—SO: MAP 168.—Recorded distribution of Eggs summer. Larvae and pupae Promachus vertebratus (Say). unknown. Adults not given. LOD : Eggs 1 week. Larvae unknown. len. Antennae are black except Pupae 32 days. Adults unknown. for a yellowish base. Front and OWS : Larvae. G/Y : One genera- face are densely covered with a ' tion is believed to last 3 years. SA : light grayish-yellow pollen. Sides Larvae and adults. N/H: Un- of the face are bare except for known. MOA: Adults attack fly- some hair the same color as the ing insects and larvae attack pollen. The bristles at the middle , white grubs. AF: Insect hemo- of the oral margin are strong and lymph. HAB : Not given. sparse, mostly lighter than the The female lays several hun- face but intermixed with a few j, dred eggs together in small crev- black ones. Thorax is grayish and ices, cracks, and earthen cavities bears dorsally two broad darker * that she constructs. In the sum- stripes, which are narrowly sep- I mer they hatch in a week and the arated. The abdomen is elongated larvae migrate through the soil and covered with dense, light * to locate prey. The larvae pierce grayish-yellow pollen on the pos- s the body of the prey and suck the terior half of each segment. Sides, ^ body fluids from the wound, after venter, and front part of each seg- ' which they seek other victims. ment are opaque black. The line *■ (55, p. 373; 323, p. 260) between the black and yellow is Description.—Eggs are white sharply marked. Posterior and and eWiTptical (323, p. 260). Lar- hypopygium are black. Legs are » vae are creamy white, smooth, and black with black and whitish pile. ^ cylindrical, tapering toward the All tibiae are light red except the extremities (323, p. 260). Pupae tip. The wings are tinged with * unknown. yellow. (193, p. 62; 287, p. h7; ' Adult males are 25 mm. long 375, pp. 60, 62) and covered with a yellowish pol- Discussion.—A single larva of 236 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE this species can destroy a large appear in the field at the end of number of white grubs. June or early July. LOD: Un- known. OWS and G/Y : Not given. Stenopogon coyote Bromley SA : Nymphs and adults. Nymphs 1931: 429 seldom survive an attack. N/H: Synonymy.—None (320, p. Unknown. MOA : Although many 383). asilid species attack their prey Recorded distribution.— only in flight, S. coyote becomes Alberta, Montana, Wyoming, Col- interested in any form of move- orado, Arizona, and New Mexico ment. It usually attacks quickly, {29, pp. Í28-Í29; 180, pp. 103- grasping its prey with its strong 107; 320, p. 383). (Map 169.) legs while it inserts its proboscis Host species.—Amphitornus between the head and thorax. The coloradus (Thomas), Ageneotet- struggle is very short. It lifts its tix deorum (Scudder), Cordil- prey up and sucks out the body lacris crenulata (Bruner), C. oc- contents. Small grasshoppers very cipitalis (Thomas), Melanoplus seldom escape an attack, medium- occidentalis (Thomas), Aulocara sized ones may escape sometimes, elliotti (Thomas), and species of and large grasshoppers are sel- Sarcophagidae (Diptera) and Epi- dom attacked. AF: Hemolymph. cauta (Coleóptera) (180, pp. 103- HAB : In Wyoming, short to me- 107; 183, pp. 28-30). dium grass rangeland at eleva- Incidence of prédation.—None tions of 4,200 to 8,000 feet. located. Copulation is generally on grass Life history and habits.—SO: stems about 3 to 5 inches above the Eggs summer to fall. Larvae un- ground. The mating pair cling to known. Pupae spring. Adults: In separate stems and face away Wyoming and Colorado they first from each other. This position forms a "V." Later the female constructs a burrow in the soil with her ovipositor between grasa clumps. These holes are 9 to 10 mm. long. Eggs are laid on top of each other. After oviposition the ovipositor is used to pack sand and soil into the hole above the eggs and to sweep the area clear. More than one clutch of eggs is generally laid within 2 to 14 min- utes, and each one contains six to 12 eggs. Examination of gravid MAP 169.—Recorded distribution of females by Lavigne (180) re- Stenopogon coyote Bromley. vealed that each female produces ARTHROPOD AND NEMATODE PARASITES IN AMERICA 237

16 to 18 eggs at a time and only Stenopogon helvolus (Loew) two ovipositions are needed to {Scleropogon) 1874: 355 empty the ovaries. (180, pp. Synonymy.—Scleropogon hel- 103-107) volus Loew {320, p. 38U). Description.—Eggs are oval Recorded distribution.—Wy- and cream to white, averaging oming, Nebraska, Kansas, and 2.18 mm. by 0.69 mm. {180, p. Texas {320, p. 38i). (Map 170.) 107). Larvae and pupae unknown. Host species.—Ageneotettix de- Adults are 15 to 19 mm. long and orum (Scudder) {183, p. 13). black basically. Pollen on the ab- Incidence of prédation.—None domen is more grayish than that located. on the thorax, which is tinged Life history and habits.—Un- with yellow, causing a slight con- known. trast between the two regions. Description.—Eggs, larvae, and Proboscis and palpi are dark red- pupae unknown. Adult nies are dish brown. The first and second 21 to 22 mm. long and chamois antennal segments are reddish yellow to nearly creamy white, yellow and the third segment and which in dead specimens turns to the bristlelike aristae are black. fawn. The thorax is covered with Covering of the head is straw yel- a dull-yellow pollen that extends low ; the darker region is the ver- over most of the body except the tex and the lighter one the beard. hind abdominal segment of the The thorax is covered by a gray- female, which is glossy. Body is ish-yellow bloom and the abdomen covered with short pale or yellow- by grayish pollen and fine straw- ish bristles. The third segment of yellow hairs. Genitalia of the the antenna is scarcely longer male are reddish as is the oviposi- than both the first two segments tor of the female, which also sup- combined. The antennae are com- . ports dark reddish-brown bristles. Legs are reddish and the femora black above. Hindtibiae are darker than the other legs. Bris- tles and hairs of the legs are straw colored. Wings are hyaline. {29, pp. U28-U29 ; 30, p. 81 ; 32, p. 296 ; 159, p. 3U) Discussion.—S. coyote attacks small grasshopper nymphs and adults and seems to prefer grass- hoppers over other insects. It ap- pears to reduce the grasshopper MAP 170.—Recorded distribution of population in its habitat. Stenopogon helvolus (Loew). 238 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

pletely black except the extreme base, which is sometimes yellow. The style is also black and con- siderably longer than half the dis- tance of the third segment. Pro- boscis is black and the palpi are the same color as the rest of the body. A thin stripe on the thorax of dark pollen is noticeably darker than the brownish surroundings. The hypopygium of the male is of the same color as the rest of the body. On the upper side of the MAP 171.—Recorded distribution of legs are long, rather wide, black- Stenopogon inquinatus Loew. ish longitudinal stripes, which on the midleg are darker at the edge. AF: Hemolymph. HAB: Un- Tarsi are black. Wings are clear known. It is assumed that the life with a weak gray to yellowish or history is similar to that of other pale sandy turbidity. Veins of the Stenopogon species. Once the prey leading edge of the wings are has been caught, the fly drops im- brown ; the remainder of the wing mediately to the ground to suck surface is brown to black. {lA, out the body contents. (32, p. pp. 208-209; 29, p. Í28; 30, pp. 293; 163, p. 331 ; 316, p. 500) 81, 97; 32, p. 296; 158, p. 3^; 207, Description.—Eggs, larvae, and p. 358) pupae unknown. Adults are 28 mm. long and dark gray. The my- Stenopogon inquinatus Loew stax, beard, setae, and thorax are 1866: 27 straw yellow. Antennae, humérus, Synonymy.—Stenopogon mo- and abdomen are reddish brown destus Loew, S. morosus Loew (4, and the palpi are dark to dark p.256;320,p.38i.). red. There is a weak stripe on Recorded distribution.—West- the thorax. Pile of the thorax is ern United States except south- short, black, and intermingled central region {320, p. 384.). with straw-colored setae. Hu- (Map 171.) mérus is red to pale yellow Host species.—"Grasshoppers" pollinose. Abdomen is dark to (123, p. 480; 336, p. 33). bricklike; last three abdominal Incidence of prédation.—None segments of the female are red. located. Hair on the abdomen is short and Life history and habits,—SO, straw color. Feet are a burnt LOD, OWS, and G/Y : Unknown. clay red ; femora are largely black SA: Nymphs and adults; prefers with many setae. Tibiae and tarsi adults. N/H and MOA : Unknown. are nearly completely black. ARTHROPOD AND NExMATODE PARASITES IN AMERICA 239

Wings are gray and smooth with Life history and habits.—SO: darkish veins, (i^, VV- 198-201 ; Eggs, larvae, and pupae unknown. 29, p. ^28 ; S2, p. 293 ; 205, p. 27) Adults not given. LOD : Unknown. Discussion.—Treherne and OWS and G/Y: Not given. SA: Buckell {386, p. 33) reported this Nymphs and adults. N/H: Un- species as predaceous on nymphal known. MOA: Captures moving and adult grasshoppers in Can- insects. AF: Insect hemolymph. ada. HAB : Between 5,000- and 9,000- foot elevation in Wyoming. (183, ' Stenopogon neglectus Bromley pp. 28-30) i 1931: 430 Description.—Eggs, larvae, and Synonymy.—None (320, p. pupae unknown. Adults are 19 to 384). 28 mm. long, basically black, and Recorded distribution.—Wash- covered with dark-gray pollinose ington, Oregon, Idaho, Montana, dust. The antennal style is bristle- Wyoming, Utah, and Colorado like and the face and occiput are (320, p. 384). (Map 172.) covered with silvery pruinosity. Host species.—Aeropedellus Beard is nearly white to straw yel- clavatus (Thomas), Ageneotettix low, and the antennae, proboscis, deorum (Scudder), Aulocara el- and palpi are black. The thorax liotti (Thomas), Cordillacris oc- is black. The humeri and beneath cipitalis (Thomas), Hesperotettix and also areas along the sutures viridis (Thomas), Melanoplus con- of pleura are reddish. Thoracic fustis Scudder, M. occidentalis hairs are straw colored except (Thomas), and Spharagemon for fine black hairs on the meso- equale (Say) (183, pp. 28-30). notum. Abdomen is black and cov- Incidence of prédation.—None ered with gray pollinose dust and located. fine straw-colored hairs. The ven- ter and margins of the segments are pale reddish brown and more narrowed in the female. The male genitalia range from reddish brown below and blackish above to totally black. The female has reddish spines on the ovipositor. The front coxae are black and the others reddish with blackened areas anteriorly. Midfemora and hindfemora are black above and reddish below. Tibiae are light reddish brown. Legs are covered MAP 172.—Recorded distribution of with straw-colored bristles and Stenopogon neglectus Bromley. hairs. Wings are short and nearly 240 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE hyaline. The first posterior cell is tarsi are black ; the anterior apices open. (29, pp. ^29-^30; 32, p. of the femur and the basal tarsus 297; 158, p. 3A) of the tibia are all reddish. My- stax is a muddy color, as is the Stenopogon obscuriventris Loew pile on the venter and the thorax. 1872: 69 (29, p. 428; 32, p. 294; 206, p. 69) Synonymy.—Stenopogon ob- scuriventria Loew (4, P- 256; 320, Stenopogon picticornis (Loew) p. 384). (Scleropogon) 1866: 26 Recorded distribution.—Cali- Synonymy.—Scleropogon picti- fornia {320, p. 384). (Map 173.) cornis Loew (320, p. 384). Host species.—"Grasshoppers" Recorded distribution.—Cali- (331, p. 100; 389). fornia and Wyoming (183, p. 13; Incidence of prédation.—None 320, p. 384). (Map 174.) located. Host species.—Ageneotettix de- Life history and habits.—SO: orum (Scudder), Aulocara elliotti Eggs and larvae unknown. Pupae (Thomas), Hadrotettix trifascia- spring. Adults about May 1. LOD : (Say), Melanoplus occidentalis Unknown. OWS and G/Y: Not (Thomas), M. sanguinipes (Fab- given. SA: Nymphs. N/H: Un- ricius), Spharagemon equale known. MOA : Attack jumping in- (Say), and Trachyrhachys kiowa sects. AF: Hemolymph. HAB: (Thomas) (183, p. IS; 331, p. Grasslands. (386) 100). Description.—Eggs, larvae, and Incidence of prédation.—None pupae unknown. Adults are 14 to located. 20 mm. long, generally black Life history and habits.—SO: brown, and covered with muddy to Eggs, larvae, and pupae unknown. ashy-gray pollen. The venter and Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Fifth-instar nymphs and adults. ^^S^\ N/H: Unknown. MOA: Captures jumping and ñying insects. AF: Hemolymph. HAB: Not given. (183, p. 13) Description.—Eggs, larvae, and pupae unknown. Adults are 21 mm. long and basically reddish. Setae are rigid and light yellow or white. Palpi are black. An- tennae are red and the second segment and base of the third

MAP 173.—Recorded distribution of segment are black. Pile on the Stenopogon obscuriventris Loew. front is short. Pile on the dorsal ARTHROPOD AND NEMATODE PARASITES IN AMERICA 241

most prefer the warmer regions. Circular flights are usually made in the warm sunshine and over feeding and oviposition areas. These flies can often be found above or on flowers and near waterholes in the more arid areas. When at rest, the wings are held away from the body. Adults are pollen and nectar feeders. Larvae of this group are either parasitic or predaceous on other MAP 174.—Recorded distribution of insects. Since the general habits Stenopogon picticomis (Loew). of the two groups are so variable, only those that are predaceous on part of the thorax is dark or grasshopper egg pods are treated black. The three stripes on the here. dorsal thorax are black, scutellum Eggs are deposited in cracks is black, and the pleura contain and crevices near ovipositing black spots. On the first abdominal grasshoppers. After a brief in- segment is a black stripe and on cubation period, the eggs hatch segments 6 and 7 are three black and the larvae seek food. Ap- stripes. Segment 8 is black or parently an encounter with grass- nearly so. Pile of the abdomen is hopper eggs is accidental since the short. Tarsi are red; femora and larvae seem to wander at random. tibiae are black. Veins of the Once an egg pod is located, the wings are dark. {H, pp. 209- covering is penetrated and the 211; 29, p. JÍ28; 82, p. 296; 163, larva begins feeding on the con- p. 3U; 205, p. 26) tents of the first encountered egg. After several pods have been de- Bombyliidae stroyed, the larva often abandons Most bee flies are stout bodied, them and seeks new ones. The furry, and capable of hovering in opening in the affected pod al- midair or rapidly darting from lows entrance of other insects and place to place. They are generally fungi. In this way more pods are black basically but covered with destroyed than by direct consump- a lighter colored pile. Wings are tion alone. The larval feeding clouded and often contain stig- period often ranges from 1 to 3 mata. A few exceptional species months, and winter is passed as are slender and do not appear a larva. The following spring the hairy. larvae leave the egg pods and There are about 500 species of pupate some distance away in the bee flies in the United States, and soil. After 1 to 4 weeks the pupae 242 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE move near the soil surface for adult emergence. There is generally one genera- tion a year. However, under ex- cessively dry conditions the larval stage may last 3 to 4 years. The number of grasshopper egg pods destroyed per individual varies with the species but often ex- ceeds three. Six species are known to attack grasshopper eggs; one also feeds MAP 175.—Recorded distribution of on larvae of the tiger beetle Cic- Anastoechus barbatus Osten Sacken. indela scutellaris Say. pupae unknown. Adults are 5 to 7 Anastoechus barbatus mm. long exclusive of the length Osten Sacken of the pile. The body is basically 1877: 252 grayish black and covered with Synonymy.—Bombylius nitidu- a dense grayish-yellow pile inter- lus Fabricius, Anastoechus nitidu- mingled with some black pile at lus (Fabricius) U, p. 237; 320, the end of the abdomen. The beard p. All). is white with intermingling black Recorded distribution.—From hairs above. The base of the an- Alberta east into Massachusetts, tennae is slightly yellowish, above south into Texas, west into Cali- which on the front is a tuft of long fornia, and north through south- black hair descending on each side east Washington. Also Europe along the eyes to about the middle and Asia. {320, p. ill) (Map of the inner orbit. Antennae are 175.) black. On the chest is a white pile, Host species.—Grasshoppers and on the abdomen in addition (3A, p. S73; 123; 183, pp. US, to the yellowish pile are black U81 ; 231, pp. 252-253; 236, p. 25; hairs arranged in rows. Femora 332, p. 33; 389) except the tips are black and Incidence of prédation.—None densely covered with white scales ; located. the tip of the femora and the Life history and habits.—SO: tibiae and tarsi are reddish yel- Eggs, larvae, and pupae unknown. low. Wings are grayish hyaline, Adults not given. LOD : Unknown. and the knobs of the haltères are OWS and G/Y: Not given. SA: yellow. Wing veins are black and Egg pods. N/H : Unknown. MOA : those at the root and near the Larvae encounter food. AF: Un- costa are often pale brown. (3A, known. HAB: Not given. (389) p. 373; 68, p. 110; 231, pp. 252- Description.—Eggs, larvae, and 253) ARTHROPOD AND NEMATODE PARASITES IN AMERICA 243

Discussion.—This species also known. MOA: Larvae encounter occurs in Europe and Asia, where food. AF: Unknown. HAB: Not it is predaceous on grasshopper given. Eggs are deposited in holes egg pods in the larval stage. and fissures or in the powdery superficial soil layer in the egg Aphoebantus hirsutas Coquillett beds of grasshoppers. (220, p. 9 ; 1886: 85 323, pp. 259, 482; 379, pp. 413- Synonymy.—None (^, p. 2U3\ 416; 389) 320, p. Í29). Description.—Eggs, larvae, and Recorded distribution.—Ore- pupae unknown. Adult females gon, California, and Arizona {320, are 7 to 8 mm. long, generally p. 429). (Map 176.) blackish basically, and covered Host species.—Camnula pellu- with a yellowish pile. The front cida (Scudder) (123, p. Í81 ; 236, has a mixture of very fine yellow pp. 3, 25; 332, pp. 259, 482; 325, and black pile and the occiput p. 7; 326, p. U; 389). whitish hair. The styliform part Incidence of prédation.—Up to of the third antennal joint is about 62.4 percent, the highest rate re- twice as long as the thick basal ported coming from the egg beds part. The thorax is yellowish and of C. pellucida in peat or sandy the pleura are white pilose. Legs loam soils (379, pp. U3-416). have a white tomentum and the Life history and habits.—SO: bristles are reddish. Wings are hy- Eggs not given. Larvae: In Cali- aline with a yellowish costal cell. fornia second instar in mid-May. In the male the eyes are very nar- Pupae: June in California. rowly separated on the front. The Adults: July in California. LOD: abdomen is about as wide as the Unknown. OWS and G/Y: Not thorax and greatly depressed. Hy- given. SA: Egg pods. N/H: Un- popygium is very small. Both the front and middle tibia support a row of white pile on the outside. (62, p. 85; 68, p. 106)

Aphoebantus mus mus (Osten Sacken) (Triodites) 1877: 246 Synonymy.—Triodites muss Osten Sacken (4, p. 243; 320, p. 430). Recorded distribution.—Cali- fornia, Utah, and Arizona (320, p.43). (Map 177.) MAP 176.—Recorded distribution of Host species.—Camnula pellu- Aphoebantus hirsutus Coquillett. cida (Scudder), Melanoplus spre- 244 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

11*^'-;V- larva begins to feed immediately and in a short time has destroyed most or all the eggs in it. It may ^ feed on several egg pods before completing development. (9^, p. ' 773 ; 323, p. 259) Description.—Eggs, unknown. Larvae are similar to those of Sys- toechus, although the head parts < are slightly broader and less flat. Mandibles are sharper than in ' Systoechus and have a smoother MAP 177.—Recorded distribution of outer edge. Labrum is lighter in Aphoebantus THUS mus (Osten Sacken). color, and the maxillae are more blunt. Thoracic spiracles are also tus (Walsh), and Melanoplus spp. less conspicuous. (265, pp. 268- (123, pp. US, A81; 236, p. 25; 269) 265, pp. 268-269; 323, p. 259; PMpae; The head is broader and -\ 325, p. 13; 326, p. ^). more bulbous than that of Sys- Incidence of prédation.—None toechus. There is a spine on the ' located. front and anterior border of each Life history and habits.—SO: wing, and the legs are all shorter. Eggs summer to fall. Larvae fall Prothoracic spiracles are less con- to spring. Pupae spring. Adults spicuous and the hairs on the ab- early summer. LOD: Eggs not dominal joints shorter. The basal given. Larvae and pupae un- abdominal segment lacks spines known. Adults not given. OWS: but possesses long stout hairs, and Mature or nearly mature larvae. the dorsal tubercles of abdominal G/Y : Usually only one ; occasion- segment 9 are replaced by a sin- ally one generation requires 2 gle spine. (265, pp. 268-269) years. SA: Egg pods. N/H: Un- Adult nies are 8 to 9 mm. long known. MOA: Larvae encounter and basically black to grayish food. AF: Flowers in the areas black. The body is uniformly cov- associated with grasshoppers. ered with a whitish-gray pile. The , HAB : Not given. face also is covered with whitish Eggs are deposited either on pile, whereas the frontal triangle the ground or in crevices near of the male is black and contains , where grasshoppers oviposit. short erect black pile and ap- These hatch after the grasshop- pressed white hairs along the or- ' pers have completed egg laying. bitals of the face. The thorax is The larvae migrate to the soil grayish black and densely covered surface to seek grasshopper egg with downy whitish-gray pile, pods. On finding an egg pod, the which is longer on the sides. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 245

Haltères are whitish with the HAB : Not given. (265, pp. 265- knob brownish. Wings, including 268) the costal cell, are hyaline except Description.—Eggs, unknown. at the root, which is black. Legs Larvae (young) are about 13 mm. are black and densely covered long and opaque whitish with with white scales. The spines on transparent yellowish mottlings the femora and tibiae are a whit- and some venous markings at the ish yellow. The female is very sim- sutures. The body is curved, gla- ilar to the male except the front, brous, swollen anteriorly, and tap- which is brownish pruinose. Also, ered posteriorly. The head is the hind margin of the abdominal small, flattened, and dark brown. segments is beset with some short, Sutures are deep. When the larvae appressed, whitish hairs, which are full and plump, the thoracic form crossbands. (68, p. 106, joints swell considerably, and 231, pp. 246-2A7; 265, pp. 268- near pupation they become more 269) opaque and more contracted. Discussion.—This species ap- (265, pp. 265-268) pears to be an effective predator Pupae are 8.5 mm. long and although its recorded distribution honey yellow when young, becom- is limited. ing darker with maturity. The head is narrow with two sets of Systoechus oreas Osten Sacken three stout dark spines on the top. 1877: 254 The two lower ones of each set are Synonymy.—None, U, p. 237; connected at the base. The thorax 320, p. 410). is unarmed. Prothoracic spiracles Recorded distribution.—South- are very large and raised on a western Canada and western curved tubercle. Mesothoracic United States (320, p. 410). spiracles are on swellings at the (Map 178.) base of the wings. The front tibiae Host species.—Camnula pellu- cida (Scudder), Melanoplus spre- tus (Walsh), and Melanoplus spp. (123, p. 481 ; 236, p. 25; 265, pp. 265-268; 325, p. 14; 326, p. 7; 336, p. 33; 389). Incidence of prédation.—None located. Life history and habits.—SO: Eggs, larvae, and pupae unknown. Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Egg pods. N/H : Unknown. MOA : MAP 178.—Recorded distribution of Larvae encountered food. AF and Systoechus oreas Osten Sacken. 246 TECHNICAL BULLETIN NO. 14Ó0, U. S. DEPT. OF AGRICULTURE

are stout, reaching the tips of The mystax is more mixed with the wings. Middle tarsi extend to fulvous pile, the proboscis is the first abdominal segment and longer, the legs are darker, and * beyond, whereas the hindtarsi the wings are more grayish. Also reach to the third abdominal seg- the ground color is less dark when ment or beyond. The abdomen is denuded. The reddishness is lack- ^ curved. On the dorsal part of each ing on the scutellum, and on the segment is a series of parallel, average the body is somewhat longitudinal, narrow chitinous longer. {231, p. 25i ; 265, pp. ' plates, each with a spine at the 265-268) extremity. There are eight pairs of abdominal spiracles and two Systoechus solitus (Walker) * pairs of thoracic spines. The first (Bombyliiis) 1849: 288 and last pairs of the abdominal spiracles are rather difficult to Synonymy.—Bombylius solitus detect. {265, pp. 265-268) Walker (320, p. 410). Adult length varies from 8 mm. Recorded distribution.—New in the female to 10 mm. in the Jersey and southeastern United < male. The male has a blackish- States (320, p. 4-10). (Map 179.) gray body basically, which is en- Host species.—Grasshoppers tirely concealed under a thick cov- (77, p. 12; 123, p. ^81 ; 326, p. 7). ■ ering of pale-yellow pile. This Incidence of prédation.—Not gives the body an elongated oval given. shape slightly broader above the Life history and habits.—SO: middle of the abdomen. Antennae Eggs, larvae, and pupae unknown. are black. The third joints are Adults not given. LOD : Unknown. considerably extended on the OWS and G/Y: Not given. SA: proximal half. Legs are also black. Egg pods. N/H : Unknown. MOA : Femora are densely covered with the usual appressed whitish hairs, which conceal the ground color. Tibiae are reddish but covered with the same whitish pubes- cence ; the outer part of the tibiae is black. On the inner side this color extends farther upward than on the outer part. Tarsi are a deep black. Wings are decidedly grayish to brownish yellow. In the female the femora and tibia except the tip are yellowish red. This species differs only slightly MAP 179.—Recorded distribution of from Systoechus vulgaris Loew. Systoechus solitus (Walker). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 247

Larvae encounter food. AF: Un- known. HAB : Not given. Description.—Eggs, larvae, and pupae unknovs^n. Adults are small, approximately 6 to 7 mm. long. They are black basically and cov- ered by a thick, shiny, light-yellovs^ pile, w^hich is longest on the ab- domen. Black hairs are on the head and clypeus. Eyes are very dark red. Antennae are black. The second joints are half the length of the first. Third antennal joint MAP 180.—Recorded distribution of is spindle shaped. Mouth parts Systoechus vulgaris Loew. are much longer than the chest. Legs are tavs^ny. Femora and tibia Life history and habits.—SO : are covered with short light-yel- Eggs not given. Larvae fall to low hairs and black spines. Tarsi spring. Pupae spring. Adults May are black and covered with short in Canada. LOD : Eggs and larvae black hairs and bristles. Wings not given. Pupae about 2 weeks. are hyaline, and the veins are fer- Adults not given. OWS: Larvae. rugineous to piceous. Haltères are G/Y: One. SA: Eggs. N/H: Un- pale tawny. (231, p. 253 ; 3^.6, p. known. MOA: Larvae encounter 288) food. AF: Nectar. HAB: Not given. Systoechus vulgaris Loew Adult flies are usually seen hov- 1863: 302 ering over areas containing grass- Synonymy.—None (4, p. 283; hoppers that are about to oviposit. 320, p. AlO). They may be so thick that the air Recorded distribution.—South- is filled with a continuous buzz. central Canada and the United About the time of mating, they States {320, p. UO). (Map 180.) circle about 8 to 12 feet above the Host species.—Camnula pellu- ground, seemingly chasing each cida (Scudder), "Acrididae," and other, and this may continue "grasshoppers" (20, pp. 169-178; throughout the warm daylight 3U, pp. 372-373; 61, p. 231; 77, hours. The flies alight from time p. 12 ; 78, p. 98 ; 123, pp. A76, ^82 ; to time to deposit their eggs di- 183, p. 7; 236, pp. 3, 25; 239, pp. rectly next to a grasshopper egg A92-Í93; 276, pp. H-18; 325, pp. pod or very near it. When the eggs 5, 7 ; 326, p. 7 ; 336, pp. 33-34-). hatch, the first-instar larvae im- Incidence of prédation.—Often mediately search out an egg pod. destroys a very high percentage They seek the protection and food of grasshopper eggs (77, p. 12). of a pod as winter approaches. 248 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Pupation occurs the next spring. pending on age. There are eight (20, pp. 169-178 ; 3Jf, pp. 372-373 ; stout, thornlike black to brown 61, p. 231; 78, p. 98; 171, p. 32; cephalic processes on the head. On 183, p. 7; 239, pp. 4^92-^93; 389) each side of the thorax above the Description.—Eggs: Berg {20, base of the wing sheath are three p. 169) described the eggs taken hairs. Prothoracic spiracles are directly from gravid females. prominent and directed backward. They are 0.87 mm. long by 0.19 Wing sheaths extend to about the mm. wide, white, bluntly ellipti- middle of the third abdominal seg- cal, and with no external mark- ment. Legs extend down the ven- ings. Larvae (first instar) are 1.5 tral midpart of the body, and the mm. to 1.7 mm. long, white, cylin- hindlegs extend as far as the pos- drical elongate, and with a trans- terior margin of the fifth abdomi- parent body wall. Body is nor- nal segment. There is a transverse mally curved anteriorly and row of reclining brown spines on thorax raised. Head and first the dorsal surface of each ab- thoracic segment are directed dominal segment. Anterior to the downward. Body is widest at the anal spines is a group of small middle and tapers slightly both tubercles and on the ventral sur- ways. There are two widely sep- face of the males is a pair of swell- arated pseudopods near the ante- ings, each with a tubercle at the rior margins of abdominal seg- apex. (20, pp. 175-177) ments 2 to 6, one on segment 7, Adults are 6 to 7.5 mm. long and two double pseudopods near exclusive of the pile. The black the posterior margin of segment basal color is covered with a 8. A pair of raised spiracles is on golden-yellow to rusty-yellow pile. the dorsolateral surface of the A black pile covers the head be- posterior margin of segment 8. hind the mentum and the occiput. The abdomen ends by tapering Legs are rusty yellow and the abruptly upward to the apex, on tarsi darker to blackish. Wings which is a pair of long curved are hyaline with a cloudy base. anal hairs. Articulating with the Veins and haltères are rusty yel- infrahypostomal bridge is the hy- low. (20, p. 177; 20JÍ, p. 302) postomal sclerite. Only the mouth (Fig. 19.) hooks, labrum, and maxillae are Discussion.—This species is visible externally. The second in- often credited with a general re- star is 4 mm. long by 0.8 mm. duction of the potential grasshop- wide. The third instar is about 10 per populations. mm. long when extended. All three stages are completely described by Berg (20, pp. 169-175), Chloropîdae Pupae are 8 to 9 mm. long and The adults of all known species yellowish to darker brown de- of Chloropidae are small and ARTHROPOD AND NEMATODE PARASITES IN AMERICA 249

sanguinipes (Fabricius) {277, pp. 151-156; 386). Incidence of prédation.—None located. Life history and habits.—SO: Eggs fall. Larvae and pupae un- known. Adults May to June in 1% Idaho. LOD and OWS : Unknown. G/Y: One or possibly two. SA: Eggs. N/H : One per several eggs. MOA: Larvae encounter food by undetermined manner. AF: Un- known. HAB : Sagebrush and PN-2737 cheatgrass complex in Idaho, FIGURE 19.—Adult of Systoechus where the vegetation is neither vulgaris Loew. compact nor sparse and farming and cattle grazing are in the vi- often chunky. Antennae are short, cinity. although the arista may be com- Adults can be seen on vegeta- paratively large. Most of the spe- tion, running about on the soil cies are black, some with yellow. surface, or in cracks and crevices. Wings are usually broad and They also burrow into the soil. rounded at the tip. Adults are not strong flyers. The larvae of most species live Neither premating courtship, and feed on grasses and other mating, nor manner of oviposition plants. Some are pests of wheat has been observed. However, once and other cereals; others feed on inside the grasshopper egg pod, the egg cases of spiders and on the larvae generally destroy or root aphids. One species in Aus- tralia is parasitic beneath the skin of certain frogs. Only one species is known to be predaceous on grasshopper eggs in the United States. Goniopista oophaga Sabrosky 1967: 153 Synonymy.—None {279, pp. 151-156). Recorded distribution.—Brit- ish Columbia and Idaho {279, pp. 151-156; 386). (Map 181.) MAP 181.—Recorded distribution of Host species.—Melanoplus Goniposita oophaga Sabrosky. 250 TECHNICAL BULLETIN NO. 14Ó0, U. S. DEPT. OF AGRICULTURE

consume the contents of several sensory areas. Haltères are yel- eggs during their development. low, and the veins of the wings Often each infested egg pod will are black. (279, pp. 151-156) contain several larvae. After com- Discussion.—Although this pletion of development the larvae species is reported from only one usually migrate outside the eggs Province and one State, it prob- or to the edge of the egg pod ably occurs elsewhere and has not where they pupate. With the ad- been noted because it is extremely vent of warm weather, adults small. Its ability to destroy sev- emerge and migrate to the soil eral eggs rather than just one surface. (389) may indicate an advantage over Description.—Eggs unknown. the Scelio wasps. Larvae are small, white, fusi- form, and slender, tapering gradually toward the anterior Therevidae end. Segmentation is prominent. Therevid flies are very similar Mouth hooks are small. (386) in appearance to the robber flies Pupae are approximately 2.3 mm. or asilids. Close observation of long by 0.85 mm. wide and a yel- the adult, however, indicates no low to mahogany color that seems furrow between the eyes and a to be dependent on size and age. slightly different wing venation. Larger and older pupae are Although the adults were pre- darker. (386) sumed to be predaceous at one Adults are 2 to 2.5 mm. long time, their mouth parts are ap- and shiny black. Palpi are black parently not equipped for pred- and of ordinary size in the female atory feeding. Legs are feebly but broadened and elongated in attached and the mannerism of the male. The ventral third of flight is generally different from each antennal segment is orange the asilids. yellow. Lower half of the face is All the species reported here whitish and the upper third of the belong to one genus, Psilocephala cheeks is silvery. Thorax is highly Zetterstedt. This is a rather large shiny. Mesonotum is convex and genus and cosmopolitan in distri- contains numerous fine prolifer- bution. Approximately 35 species ous punctures. Scutellum is flat- are reported from North America. tened, wrinkled, and broadly The larvae are predaceous, at- rounded distally. Knees, the bases tacking other insect larvae and and apices of all tibiae, and the eggs within the soil. In this re- midtarsi and hindtarsi (except spect they are similar to the distally) are yellow to orange yel- asilids. Although only three spe- low. Hair and bristles are black. cies are described here, it is possi- Legs are well developed, and the ble that many more are predaceous hindtibiae contain conspicuous on grasshopper eggs. There is no ARTHROPOD AND NEMATODE PARASITES IN AMERICA 251 evidence to indicate any prefer- Description.—Eggs, larvae, and ence for one type of egg over an- pupae unknown. Adults are 7 to 8 other or whether eggs are pre- mm. long and basically black. ferred to larvae. Knobs of the haltères, hypopy- gium of the male, and base of the Psilocephala aldrichii Coquillett tarsi are yellow. Posterior mar- gins of the second and third ab- 1893: 227 dominal segments of the male are Synonymy.—None (i, p. 24-6; white. The frons of both sexes is 320, p. 350). a shining white pollinose as is the Recorded distribution.—West- face. The oral pile and the occiput ern Canada, western United hairs are also white, whereas the States, and New Jersey (320, p. hairs of the cheeks and the bris- 350). (Map 182.) tles of the antennae are mostly Host species.—Camnula pellu- black. First antennal segment is cida (Scudder) (389). three-fourths the length of the Incidence of prédation.—Not third; the third is lanceolate and given. approximately six times as long Life history and habits.—SO, as the style. The thorax is some- Eggs, larvae, and pupae unknown. what shiny and bears two white Adults not given. LOD : Unknown. pollinose stripes, which may be OWS and G/Y: Not given. SA: indistinct. The pile of the thorax Eggs. N/H: Unknown. MOA: and scutellum is yellowish to Larvae encounter food. AF: Un- white, and four black bristles are known. HAB : Wide range of eco- on the scutellum. The abdomen of logical conditions from coastal the male is a light-white pollinose, type to desert areas. (61, p. 173; shining, and with white pile ex- 389) cept for the hypopygium, which contains some black pile. The fe- male supports mostly black pile on the abdomen. Wings of both sexes are hyaline; the veins and the stigma are light yellow. (67, pp. 22U, 227-228) Psilocephala haemorrhoidalis (Macquart) (Thereva) 1840: 26 Synonymy.—Thereva hoemor- rhoidalis Macquart (i, p. 266; 320, p. 351). MAP 182.—Recorded distribution of Recorded distribution.—East- Psilocephala aldrichii Coquillett. ern and south-central United 252 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

States except Maine (320, p. 351). light hair on the abdomen and (Map 183.) sides as well as below. The seventh Host species.—Lepidoptera, abdominal segment and the geni- Coleóptera, and grasshopper {211, talia are orange. Feet are tan and p. 26; 326, p. HO). the thighs black with white hair. Incidence of prédation.—Not Tibiae also have small hairs. given. Wings are yellowish hyaline and Life history and habits.—SO: the veins are slightly yellowish. Eggs, larvae, and pupae unknown. (211, p. 26) Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Psilocephala lateralis Adams Eggs. N/H: Not given. MOA: 1904: 444 Larvae encounter egg pods. AF: Not given. HAB: Larvae in egg Synonymy.—None (320, p. 351 ). beds. (389) Recorded distribution.—Brit- Description.—Eggs, larvae, and ish Columbia, California, Ari- pupae, unknown. Adults are bas- zona, and Colorado (320, p. 351 ; ically black with the face and part 336, p. 33). (Map 184.) of the front covered with white Host species.—Camnula pellu- hair. Antennae are black. The cida (Scudder) (123, p. A50; 330, first joints are somewhat long and p. 178; 336, p. 33). have bristles, whereas the third joints are reduced or missing. Incidence of prédation.—Not Hair on the thorax is slate colored. given. A dorsal stripe and two black Life history and habits.—SO: spots are on the side, and hair Eggs, larvae, and pupae unknown. on the sides is white. There is Adults not given. LOD : Unknown. OWS and G/Y: Not given. SA: Egg pods. N/H : Unknown. MOA : Larvae encounter food. AF: Not given. HAB: Desert areas. (61, p. 173; 336, p. 33) Description.—Eggs, larvae, and pupae unknown. Adult males are about 7 mm. long and basically black. Face is white pollinose and there are white areas on the lower corners of the head. Front is shining and black. Antennae are black. The third joints have a faint reddish cast at the base. MAP 183.—Recorded distribution of Psilocephala haemorrhoidalis (Mac- Black bristles are on the first and quart) . second antennal joints. Aristae ARTHROPOD AND NEMATODE PARASITES IN AMERICA 253

rax is black and the dorsal part of the mesonotum and the scutel- lum are grayish-brown pollinose. Pile is yellowish white and bristles are black. Abdomen is also black and shining. The first segment has abundant white pollinose dust as do the apices of the second and third segments. All segments con- tain a sparse white pilosity. Tip of the hypopygium has a yellowish cast. Legs are black with white MAP 184.—Recorded distribution of pile and black bristles. There is Psilocephala lateralis Adams. white pilosity on the coxae, and / the middle tibia, hindtibia, and are small and acute. Mouth parts tarsi are reddish yellow. Wings are black, whereas the palpi bear have a yellowish-brown tinge ; the white pile. At the junction of the darkest part is on the anterior face and cheeks is a deep black half. Haltères are lemon yellow. spot with short black pile. Tho- (3, p. iU)

Hymenoptera

Formicidae largest members of the group and are winged prior to mating. Males One rarely considers the possi- are also winged but are consider- bility that ants may have an im- ably smaller than the queens. portant eiïect on grasshoppers. After mating the male dies and They can destroy tremendous the queen sheds her wings and numbers of first-instar grass- begins a new colony or takes over hopper nymphs as they emerge one already established. She feeds from the egg pod. This has been and cares for her first brood, ' observed by Lavigne and Pfadt which consists of workers. Once ^ (183) and others. However, little these workers appear, they take additional information has been over the work of the colony, such ** recorded. as nest construction, gathering ^ Ants are probably the most food, and caring for the young, successful of all the insect groups. and thus relieve the queen of all r They occur nearly everywhere in duties but egg laying. ^ terrestrial habitats. They are social in habit, living in colonies Ants utilize a great variety of I** with three castes—queens, males, food sources, ranging from plant and workers. The queens are the sap, fungi, seeds, and honeydew 254 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

secreted by various insects to flesh of other animals living or dead. Some are predaceous on a wide variety of insects and v^ere the first insect group utilized in an attempt to control other insects. One of the most effective species is in the genus Solenopsis, which has been credited with destroying up to 75 percent of the larvae, pupae, and adults of the south- western corn borer (Diatraea grandiosella (Dyar)). MAP 185.—Recorded distribution of Although ants prey most effec- Formica obtusopilosa Emery. tively on grasshoppers as they exit from the egg pod, their ef- Incidence of prédation.—Not fectiveness decreases as the grass- given. hoppers increase in size until only Life history and habits.—SO: sick, dying, or dead grasshoppers Eggs, larvae, and pupae unknown. are taken. Only four species of Adults not given. LOD : Eggs, lar- ants are noted here, but presum- vae, and pupae unknown. Adults ably a large number of species not given. OWS and G/Y: Not destroy young grasshoppers as given. SA: First-instar nymphs. they emerge. N/H: Variable. MOA: First- instar nymphs as they exit from Formica obtusopilosa Emery their egg pods. AF: Insects and 1893: 643, 648 plant material. HAB: Gravel al- luvia, clay, and under rocks in Synonymy.—Formica sangu- canyon bottom meadows; piñon- ínea obtusopilosa Emery, F. cedar woodland, forests, short munda Wheeler, F. pergandei grass prairie, sagebrush desert, Emery (225, p. 869). pastures, roadside, and exposed Recorded distribution.—Al- bedrock usually at higher eleva- berta, Montana, North Dakota, tions. In Colorado it is generally South Dakota, Minnesota, Ne- found between 6,000 and 7,000 braska, Colorado, Utah, and New feet. Colonies are comparatively Mexico. {125, pp. 566-572). small and consist of only a few (Map 185.) hundred individuals. Nests are Host species.—Ageneotettix de- obscure and generally constructed orum (Scudder), Aulocara elliotti in open meadows. (125, pp. 566- (Thomas), and Cordillacris oc- 572 ; 183, p. 31 ; 352, pp. 165-206 ; cipitalis (Thomas) (183, p. 31 ; S5Jf, pp. H-O-IU) 331, p. 150). Description.—Eggs, larvae, and ARTHROPOD AND NEMATODE PARASITES IN AMERICA 255

pupae unknown. Adults: Head Phoetaliotes nebrascensis and thorax are clear f errugineous (Thomas), and species of Coleó- red and the gaster is brown to ptera, Hemiptera, Homoptera, piceous black. Size was not given. Lepidoptera, Hymenoptera, Díp- Long, stout, silvery, erect hairs tera, and other Orthoptera (183, cover the gaster more abundantly pp. H, 31 ; 354-, pp. 140-141). than the rest of the body. There Incidence of prédation.—Not are erect hairs on the dorsum of given. the pronotum, mesonotum, and Life history and habits.—SO: epinotum. The gaster is opaque Eggs by April. Larvae spring to with a pubescence that partially summer. Pupae summer. Adults obscures the fine sculpturing. fall to spring. LOD: Eggs 23 to (59, pp. 30-33; 72, pp. 1-5; 73, 53 days. Larvae 7 to 33 days. pp. h73-U79, Í81, A90, h92-U9U; Pupae and adults not given. OWS . 112, p. 39 ; 362, pp. 392, Í33-A3Í) Adults. G/Y: Not given. SA Discussion.—During a short First-instar grasshoppers. N/H period this species can have a very Variable. MOA: Adults attack pronounced eiïect in reducing weak, dying, or young insects. potential grasshopper popula- They also attack first-instar tions. nymphs as they exit from their egg pods. AF: Various insects. Formica rufa obscuripes Forel HAB : Nesting sites consist of •" 1886: 39 conspicuous thatched mounds in Synonymy.—Formica rufa ob- open areas devoid of cover. The scuriventris var. rubigenosa habitat encompasses a wide va- Emery, F. rufa aggerans Wheeler, riety of climatic conditions and F. obscuripes Forel, F. rufa Mc- extends from hot, dry areas to -^ Cook, F. rufa obscuriventris mel- anotica Emery, F. rufa obscuHpes melonotica Wheeler, F. rufa ob- scuripes melanotica Weber, F. rufa melanotica Creighton (,225, ' p. 867). Recorded distribution.—North- western North America (91, pp. - 6Jf3, 648; 225, p. 867; 362, pp. ^ 389, U-Í-Í18). (Map 186.) Host species.—Ageneotetti de- orum (Scudder), Aulocara cUiotti (Thomas), Cordillacris occipitalis (Thomas), Psoloessa delicatula ^ Scudder, Psoloessa spp., Melano- MAP 186.—Recorded distribution of plus spp., M. bivittaius (Say), Formica rufa obsuripes Forel. 256 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

those with moderate to heavy the nest. Weber reported that rainfall. seven insect orders are attacked. Nesting mounds are 8 to 18 Orthoptera is the largest group inches high and contain never comprising about 26 percent of more than three openings usually- all the insects caught. Most of the located near the summit. Cham- grasshoppers taken to the nest bers extend down more than 41/2 were in such damaged condition feet. Materials for the nest con- that specific identification was al- sist of twigs, herb stems, and most impossible. Specimens of grasses mixed with soil. Mounds Melanoplus were most common. may be found in deserts, woods, This species is not without its or near streams and all are in own enemies. Man often cultivates direct sunlight and not sur- the area that is or can be utilized rounded by heavy vegetation. for nest making. Large animals, When vegetation encroaches, the such as cows and horses, trample colony moves. The mound is con- its nests. This species is eaten by stantly in danger of strong winds birds, frogs, and rodents and is and encroaching vegetation. attacked by such parasites as Thatch lost by winds is replaced, mites. (183, pp. 1^, 31; 352, pp, and the workers attempt to har- 165-206) vest the encroaching vegetation. Description.—Eggs are 0.60 The vegetation benefits this mm. long by 0.31 mm. wide, ellip- species in two ways : First, it acts soidal, and milky white {352, pp. as a pasture ground for the 165-206). Larvae (first instar) aphids, which these ants *'milk" are about 0.60 mm. long or the for honeydew, a product that same size as the eggs and develop supplies much nourishment; and to a maximum length of about 6 second, the vegetation attracts mm. {352, pp. 190-191). Pupae membracids, grasshoppers, and (male and female) are about 9 aphids, which are the chief source mm. long and the worker pupae of food for this species. may vary from 3.5 to 7 mm. {352, Insects for food may be ob- p. 191). tained by several methods. Those Adult workers are 3.8 to 8 mm. that are weak or dying may be long. The head and thorax are attacked and dragged into the reddish and the abdomen is black. nest. Weber {352) observed a This ant is similar to the Euro- worker deliberately fall several pean species Formica rufa Lin- inches onto a fluttering moth in naeus. The small workers are an attempt to capture it. He also much darker and have brown observed a struggling specimen of spots on the head and thorax, Coccinella transversoguttata var. whereas the abdomen is unpol- quinquenotata Kirby being ished and black with a gray dragged toward the entrance to pubescence that shows up a little ARTHROPOD AND NEMATODE PARASITES IN AMERICA 257 stronger than in F. rufa. The pile is also slightly more feeble. The larger workers are lighter red and entirely without spots on the head and thorax, whereas the feet are brown to black. Erect hairs on the thorax are rather unequal in length, whereas the cephalic hairs are less abundant and about the same size as those on the tho- rax. Median lobe of the clypeus descends to the clypeal fossae, which are shallow and scarcely MAP 187.—Recorded distribution of pitlike. Scale of the petiole is Myrmica sabuleti americana Weber. angularly produced upward in the middle, as seen from behind, Incidence of prédation.—Not whereas the sides taper inward given. evenly from the crest to the Life history and habits.—SO: peduncle. Antennal scapes lack Eggs, larvae, and pupae unknown. erect hairs except some on the Adults not given. LOD : Unknown. extreme tip. {60, pp. 376-378; OWS and G/Y: Not given. SA: 73, pp. Jt92-JÍ9A; 91, pp. 6A3, 6A8; First-instar nymphs. N/H: Var- 360, p. 267; 861, pp. A33-Í3Í) iable. MOA : Adults attack young Discussion.—Under ideal con- prey, sick, weak, and dying ditions and for a limited time this nymphs, and adults, and sick and species can reduce the potential dying insects of young to old. AF : grasshopper population. Insects of various species. HAB: This species is found at an alti- tude of 3,500 to 7,000 feet in Colo- Myrmica sabuleti americana Weber rado and is generally associated with the Sonoran and Transition 1939: 144 zones. Its habitat includes gravel, Synonymy.—Myrmica sabuleti sand, rocks, boulders, and wood. trullicornis Burmeister, M. amer- It is found around ponderosa icana Weber {225, p. 790). pines, foothills, deciduous canyon Recorded distribution.—From forests, grass prairies, and grass- eastern Canada and northeastern lands. {125, pp. 309-312; 183, United States west to Rocky pp. H, 31) Mountains and including moun- Description.—Eggs, larvae, and tains in Utah {125, pp. 309-312; pupae unknown. Adult workers 225, p. 790). (Map 187.) are 4.6 to 6.2 mm. long. The head Host species.—Trachyrhachys is brown and the thorax and ap- kiowa {Thomas) {183,pp.lA,31). pendages are a rusty reddish 258 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE brown. The gaster is dark brown, lighter brown. Wings are hyaline slightly lighter apically, smooth, when present with a distinct and shining, whereas the rest of brownish cast. Veins are brown. the body is punctate as the base The pilosity is moderately abun- of the sculpturing. Dorsally the dant, acute, and fine. There is an petiole has a deep, irregular appressed pubescence on the an- rugose appearance. The thorax is tennae and legs. Surface of the coarsely and sharply rugose. Sur- head is shining with compara- face of the head is regularly and tively numerous ridges or moderately sculptured and the wrinkles punctate at the base of frontal area distinct and finely the sculpturing. Thorax is also striate punctuate. Clypeus has 10 shining and abundantly wrinkled, to 14 wrinkles. The thorax in pro- except on the large smooth scutel- file is slightly convex and has a lum of the mesonotum. Petiole is distinct, rounded mesoepinotal thinly but distinctly wrinkled and suture. The epinotal spines pro- finely punctate basally. The post- ject upward and backward at petiole and the gaster are smooth about a 55° angle. The petiole in and shining. (73, pp, 88, 90, 93- profile has a concave anterior face 95, 100; 125, pp. 309-312; 353, that meets the dorsal convex sur- p. lU) face at a round angle and is as Discussion.—This species can long from the apex of the ventral be an effective predator for only tooth to the postpetiole as it is a limited time. high. The gaster is ovate and the legs are of moderate length. Solenopsis molesta validiuscula The female is 5.2 to 7 mm. long. Emery The head is rusty brown becoming 1895: 278 brownish middorsally. The tho- Synonymy.—None (225, p. rax is rusty red with two elongate 8U), brown blotches on the mesonotum, Recorded distribution.—From whereas the remainder of the Pacific Coast States east into body and the appendages are Idaho, Wyoming, Colorado, and brownish rusty red and the gaster New Mexico (225, p. 8H). (Map has a brown transverse medial 188.) band. Wings when present are Host species.—Aulocara elliotti hyaline with a brownish case and (Thomas) (183, pp. 15, 31). the veins are brown. Incidence of prédation.—Not Males are 5.2 to 6.6 mm. long. given. The head is blackish brown and Life history and habits.—SO: the remainder of the body and Eggs, larvae, and pupae unknown. appendages are dark brown. The Adults not given. LOD : Unknown. pedicel and base of the gaster, the OWS and G/Y: Not given. SA: antennal club and the tarsi are First-instar grasshopper nymphs. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 259

Description.—Eggs, larvae, and pupae unknown. Adults are 1.8 to 2 mm. long and grayish-brown- ish yellow. Eyes of the female cover less than half the side of the head. Epinotum is rather rounded at the junction of the base. The female thorax is not slender. The mesoepinotal suture of the thorax is not impressed. Cephalic punctures are sparse and small, often visible only under MAP 188.—Recorded distribution of high magnification of a micro- Solenopsis molesta validiuscula Emery. scope and not much larger in diameter than the hairs arising , N/H : Variable. MOA : Adults at- from them. Eyes of the worker tack very young, weak, sick, dy- have not more than 15 facets and ing, and dead insects. AF : Insects. usually 10 or less. Funicular HAB : It is generally found at an joints 2 and 3 of the worker at elevation of 3,500 to 8,378 feet in most are very slightly longer than ■* Colorado in a wide variety of eco- broad, usually broader than long. ^ logical conditions. It is extremely {73, pp. 230, 2SA-237; 89, pp. 22- common and prefers the Sonoran 23; 92, p. 278; 125, pp. 375-377; and Transition zones. It can be 361, p. Jf30) ^ found around gravel, sand, under Discussion.—Like the other rocks and boulders, in duff, and species of ants, S. m. validiuscula * under dung. {125, pp. 375-377; is only effective as a grasshopper ^ 183, pp. 15,31) predator for a short time.

Araneida

Possibly the least studied of While attempting to escape, it vi- » the grasshopper predators are the brates the web and thus arouses ^ spiders (fig. 20). This is rather the spider. The spider locates the surprising since large numbers of victim and immobilizes it either * grasshoppers may often be seen by attacking and injecting a para- » entangled in spider webs or woven lyzing substance into the body or into cocoons. Spiders are not host by quickly enshrouding the prey 10' specific in their diet and most in- with the tough webbing. If the ■ sects are utilized, although some spider is hungry, the oral appara- may be rejected. The victim be- tus is inserted into the hemocoel comes entangled in the web. of the victim and digestive juices 260 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

PN-2738 FIGURE 20.—Spider. are injected. After a short wait tatus victims. The grasshopper the predigested material is sucked population was rather high, and out of the body shell. Later the as the closeness of these insects carcass is usually discarded— caused them to jump, some would often in a pile directly under the become ensnared in the many web. If the spider is not hungry, webs. Immediately they were the victim is generally wrapped pounced upon and wrapped until and stored in the web for later they were no longer visible. The use. spider would return to the top of During the summer of 1965 the web while the wrapped grass- near Horseshoe Bend, Idaho, I ob- hoppers continued to struggle. served a field of "silver harvester" Even when several became caught spiders at work. Their webs were at the same time, the spider stretched among sunñower plants quickly and efficiently wrapped and under each web were 20 to 90 each one in turn before retreating empty shells of adults and nymphs to the top of the web. of Mekmo'plus bivittatus (Say). Web design is usually unique In each web were several M. bivit- for each species as is the location ARTHROPOD AND NEMATODE PARASITES IN AMERICA 261 of construction. Webs may be enopsis Oklahoma Gertsch, Agiope found at the entrance of small tHfasciata Forskal, Dendryphan- animal burrows, vertically be- tes insoleus Hentz, Latrodectus tween or within bushes, or hori- mactons (Fabricius), Pelens spp., zontal to the ground. Some species Schizocosa minnesotensis Gertsch, do not use a web and prefer to SteatodacoroUataEmertoji,Than- ambush their victims. atus altimontis Gertsch, Xystiem Only nine species of Araneida coloradensis Bryant. I believe the are recorded as being predatory list would be very long if spider on grasshoppers. These are Agel- food habits were well known.

REFERENCES

(1) ABBOTT, R. L. 1931. INSTINCT OR INTELLIGENCE IN THE GREAT GOLDEN DIGGER.' lOWa Acad. Sei. 38: 255-258. (2) ADAMS, C. C. ..,, 1915. AN ECOLOGICAL STUDY OF PRAIRIE AND FOREST INVERTEBRATES. 111. State Acad. Nat. Hist. 11: 33-280. (3) ADAMS, C. F. 1904. NOTES ON AND DESCRIPTIONS OF NORTH AMERICAN DÍPTERA. Kans. Univ. Sei. Bul. 2: 433-455. (4) ALDRICH, J. M. 1905. CATALOGUE OF NORTH AMERICAN DÍPTERA (OR TWO-WINGED FLIES). Smithsn. Inst. Mise. Colleet. XLVI, No. 1444, 680 pp.

1916. SARCOPHAGA AND ALLIES IN NORTH AMERICA. 301 pp. ThomaS Say Foundation, Lafayette, Ind. (6) 1927. THE DIPTEROUS PARASITES OF THE OF TROPICAL AMERICAN, SCHISTOCERCA PARANENSIS BURMEISTER. Jour. Eeon. Ent. 20: 588-593. (7) ANDREWES, C. 1969. THE LIVES OF WASPS AND BEES. 204 pp. New York. (8) ARNAUD, P. H., JR. 1963. TYPES OF THE TACHINIDAE (DIPTERA) IN THE AMERICAN MUSEUM OF NATURAL HISTORY. Amer. Mus. Nat. Hist. Bui. 125, pp. 103-107. (9) ARNETT, R. H., JR. 1968. THE BEETLES OF THE UNITED STATES (A MANUAL FOR IDENTIFICA- TION). 1112 pp. Ann Arbor, Mieh. (10) ASHMEAD, W. H. 1887. STUDIES OF THE NORTH AMERICAN PROCTOTRUPIDAE WITH DESCRIP- TIONS OF NEW SPECIES FROM FLORIDA. Pt. I. Amer. Ent. 3: 117-119.

1893. A MONOGRAPH OF THE NORTH AMERICAN PROCTOTRUPIDAE. U.S. Nati. Mus. Bul. 45, 472 pp. 262 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(12) 1894. THE HABITS OF THE ACULEATE HYMENOPTERA. III. Psyche 7: 59-66. (13) 1899. FOUR NEW ENTOMOPHiLOUS WASPS. Ent. News 10: 9-10. (14) BACK, E. A. 1909. THE ROBBER-FLIES OF AMERICA, NORTH OF MEXICO, BELONGING TO THE SUBFAMILIES LEPTOGASTRINAE AND DASYPOGONINAE. Amer. Ent. Soc. Trans. 35: 137-400. (15) BALDUF, W. V. 1935. THE BIONOMICS OF ENTOMOPHAGUS COLEÓPTERA. 220 pp. St. Louis, Mo. (16) BANKS, N. 1921. NEW NEARCTic FOSSORIAL HYMENOPTERA. Ent. Soc. Amer. Ann. 14: 16-26. (17) 1942. NOTES ON THE UNITED STATES SPECIES OF TACHYTES (HYMENOPTERA: LARRiDAE). Harvard Univ. Mus. Compar. Zool. Bui. 89, pp. 421-435. (18) BEQUERT, J. 1934. NOTES ON AMERICAN NEMESTRINIDAE, SECOND PAPER. N.Y. Ent. Soc. Jour. 42: 163-184. (19) 1950. A NEMESTRINID BRED FROM A GRASSHOPPER IN THE UNITED STATES. Brooklyn Ent. Soc. Bui. 45: 104. (20) BERG, V. L. 1940. THE EXTERNAL MORPHOLOGY OF THE IMMATURE STAGES OF THE BEE FLY, SYSTOECHUS VULGARIS LOEW (DÍPTERA, BOMBYLIIDAE) , A PRED- ATOR OF GRASSHOPPER EGG-PODS. Ganad. Ent. 72: 169-178. (21) BLATCHLEY, W. S. 1910. EPiCAUTA PENNSYLVANICA (DEG.). In The Coleóptera of Indiana, Bui. 1, 1363 pp. Ind. Univ. Dept. Geol. and Nat. Resources, Indianapolis. (22) BLICKENSTAFF, C. C, and SHARIFULLAH, M. 1962. INFESTATION OF OVERWINTERING NYMPHS OF CHORTOPHAGA VIRIDI- FASCIATA BY MERMITHIDS. Jour. Econ. Ent. 55: 268. (23) BORROR, D. J., and DELONG, D. M. 1954. AN INTRODUCTION TO THE STUDY OF INSECTS. 1030 pp. New York. (24) BoviNG, A. G., and CRAIGHEAD, F. C. 1931. AN ILLUSTRATED SYNOPSIS OF THE PRINCIPAL LARVAL FORMS OF THE ORDER COLEÓPTERA. Amer. Ent. (n.s.) 11: 1-351. (25) BRADLEY, J. C. 1919. A NEW TACHYTES FROM GEORGIA (HYMENOP. : LARRIDAE) . Ent. NeWS 30: 298. (26) BRAUER, P., and BERGENSTAMM, J. V. VON. 1891. DIE ZWEIFLÜGLER DAS KAISERLICHEN MUSEUMS ZU WIEN. V. VORAR- BEITEN ZU EINER MONOGRAPHIE DER MUSCARIA SCHIZOMETOPA (EX- CLUSIVE ANTHOMYiDAE) PARS II. K. Akad. der Wiss. Wien, Math.- Nat. Kl. Denkschr. 58: 305-446. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 263

(27) BRIAND, L. J., and RIVARD, I. 1964. OBSERVATIONS SUR MERMIS SUBNIGRESCENS COBB (MERMITHIDAE) , NEMATODE PARASITE DES CRIQUETS AU QUÉBEC. Phjrtoprotection 45: 73-76. (28) BRIDWELL, J. C. 1920. MISCELLANEOUS NOTES ON HYMENOPTERA, 2ND PAPER, WITH DESCRIP- TIONS OF NEW SPECIES. Hawaii Ent. Soc. Proc. 4, pp. 386-403. (29) BROMLEY, S. W. 1931. NEW ASILIDAE, WITH A REVISED KEY TO THE GENUS STENOPOGON LOEW: (DÍPTERA). Ent. Soc. Amer. Ann. 24: 427-435. (30) 1934. THE ROBBER FLIES OF TEXAS (DÍPTERA, ASILIDAE) . Ent. Soc. Amer. Ann. 27: 74-113. (31) 1936. THE GENUS DIOGMITES IN THE UNITED STATES OF AMERICA WITH DE- SCRIPTIONS OF NEW SPECIES (DÍPTERA: ASILIDAE). N.Y. Ent. Soc. Jour. 44: 225-237. (32) 1937. THE GENUS STENOPOGON LOEW IN THE UNITED STATES OF AMERICA (ASILIDAE: DíPTERA). N.Y. Ent. Soc. Jour. 45: 291-309. (33) 1951. ASILID NOTES (DíPTERA) WITH DESCRIPTIONS OF THIRTY-TWO NEW SPECIES. Amer. Mus. Nat. Hist. Amer. Mus. Novitates 1532, 36 pp. (34) BROOKS, A. R. 1952. IDENTIFICATION OF BOMBYLIID PARASITES AND HYPERPARASITES OF PHALAENIDAE OF THE PRAIRIE PROVINCES OF CANADA, WITH DESCRIP- TIONS OF SIX OTHER BOMBYLIID PUPAE (DÍPTERA). Cañad. Ent. 84: 357-373. (35) 1958. ACRIDOIDEA OF SOUTHERN ALBERTA, SASKATCHEWAN, AND MANITOBA (ORTHOPTERA). Canad. Ent. Sup. 9, 92 pp.

(36) BRUES, C. T. 1917. ADULT HYMENOPTEROUS PARASITES ATTACHED TO THE BODY OF THEIR HOST. Nati. Acad. Sei. Proc. 3: 136-140. (37) BUCKELL, E. R. 1945. THE GRASSHOPPER OUTBREAK OF 1944 IN BRITISH COLUMBIA. Canad. Ent. 77: 115-116. (38) and SPENCER, G. J. 1945. A PRELIMINARY LIST OF THE FLIES OF BRITISH COLUMBIA. Brit. Co- lumbia Ent. Soc. Proc. 42: 6. (39) BURGESS, A. F., and COLLINS, C. W. 1917. THE GENUS CALOSOMA: INCLUDING STUDIES OF SEASONAL HISTORIES, HABITS, AND ECONOMIC IMPORTANCE OF AMERICAN SPECIES NORTH OF MEXICO AND OF SEVERAL INTRODUCED SPECIES. U.S. Dept. Agr. Bul. 417, 124 pp. (40) BURKS, D. B. 1936. THE ILLINOIS SPECIES OF BRACHYMERIA (HYMENOPTERA: CHALCIDAE) . 111. State Acad. Sei. Trans. 33: 251-254. (41) 1940. THE HOST OF ANOTHER ILLINOIS SPECIES OF BRACHYMERIA (HYME- NOPTERA). ni. state Acad. Sei. Trans. 33: 208. 264 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(42) CARTWRIGHT, W. B., BLANCHARD, R. A., and WILSON, C. C. 1933. NOTES FOR 1932 ON CEREAL FORAGE INSECTS IN CALIFORNIA. Calif. Dept. Agr. Monog. Bui. 22, pp. 156-160. (43) CASEY, T. L. 1913. STUDIES IN THE CICINDELIDAE AND CARABIDAE OF AMERICA. Mem. Coleóptera 4, pp. 1-192. (44) 1914. A REVISION OF THE NEARCTic HARPALINAE. Mem. Coleóptera 5, pp. 45-305. (45) 1918. STUDIES AMONG SOME OF THE AMERICAN AMARINAE AND PTEROSTI- CHiNAE. Mem. Coleóptera 8, pp. 224-393. (46) 1920. RANDOM STUDIES AMONG THE AMERICAN CARABOIDEA. Mem. Coleóp- tera 9, pp. 133-299. (47) CHABORA, P. C. 1970. STUDIES OF PARASITE-HOST INTERACTION USING GEOGRAPHICAL STRAINS OF THE BLOW FLY PHAENCIA SERICATA AND ITS PARASITE NASONiA viTRiPENNis. Ent. Soc. Amer. Ann. 63: 495-501. (48) CHAUDOIR, B. M. 1850. PASIMACHUS, LEBiA, CARABUs. . . . Soc. Nat. de Moscou Bul. 22: 3-85. (49) CHITTENDEN, F. H. 1903. A BRIEF ACCOUNT OF THE PRINCIPAL INSECT ENEMIES OF THE SUGAR BEET. U.S. Dept. Agr. Div. Ent. Bui. 43, 71 pp. (50) CHRISTIE, J. R. 1928. NOTES ON LARVAL NEMAS FROM INSECTS. Jour. Parasitol. 15: 127- 130. (51) 1936. LIFE HISTORY OF AGAMERMIS DECAUDATA, A NEMATODE PARASITE OF GRASSHOPPERS AND OTHER INSECTS. Jour. Agr. Res. 52: 161-198. (52) 1937. MERMIS SUBNIGRESCENS, A NEMATODE PARASITE OF GRASSHOPPERS. Jour. Agr. Res. 55: 353-364. (53) 1960. SOME INTERESTS ENTOMOLOGISTS AND NEMATOLOGISTS HAVE IN COM- MON. Fla. Ent. 43: 43-48. (54) CLAASSEN, P. W. 1917. GRASSHOPPERS OF KANSAS. SYSTEMATIC TREATISE OF THE MELANOPLI WITH REFERENCE TO VARIATIONS FROM SCUDDER'S DESCRIPTIONS. Kans. Univ. Dept. Ent. Bui. 11, pp. 7-126. (55) CLAUSEN, C. P. 1940. ENTOMOPHAGOUS INSECTS. 688 pp. New York. (56) COBB, N. A. 1926. THE SPECIES OF MERMIS. Jour. Parasitol. 13: 66-72 (57) 1927. NEMAS SOMETIMES AID MAN IN HIS FIGHT TO CONTROL INSECT PESTS. U.S. Dept. Agr. Ybk. Separate 1022, pp. 479-480. (58) STEINER, G., and CHRISTIE, J. R. 1923. AGAMERMIS DECAUDATA COBB, STEINER AND CHRISTIE, A NEMA PARA- SITE OF GRASSHOPPERS AND OTHER INSECTS. Jour. Agr ReS 23: 921-926. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 265

(59) COLE, A. C, JR. 1932. THE THATCHING ANT, FORMICA OBSCURIPES FOREL. Psyche 39: 30- 33. (60) 1942. THE ANTS OF UTAH. Amer. Midland Nat. 28: 358-388. (61) COLE, F. R. 1969. THE FLIES OF WESTERN NORTH AMERICA. 693 pp. Berkeley and Los Angeles. (62) COQUILLETT, D. W. 1886. MONOGRAPH OF THE LOMATINA OF NORTH AMERICA. Canad. Ent. 18: 81-87. (63) 1886. REPORT ON THE LOCUSTS OF THE SAN JOAQUÍN VALLEY, CALIFORNIA. U.S. Dept. Agr. Ann. Rpt. 1885: 289-313. (64) 1892. THE DIPTEROUS PARASITE OF MELANOPLUS DEVASTATOR IN CALIFORNIA. Insect Life 5: 22-24. (65) 1892. A NEW ASILID GENUS RELATED TO ERAX. Canad. Ent. 25: 175-177.

1892. REPORT ON THE LOCUST INVASION OF CALIFORNIA. In Riley, C. V., ed.. Reports on the Damage by Destructive Locusts During the Season 1891. U.S. Div. Ent. Bui. 27, pp. 34-57. (67) 1893. SYNOPSIS OF THE DIPTEROUS GENUS PSILOCEPHALA. Canad. Ent. 25: 222-229. (68) 1894. NOTES AND DESCRIPTIONS OF NORTH AMERICAN BOMBYLIIDAE. Amer. Ent. Soc. Trans. 21: 89-112. (69) 1895. DESCRIPTIONS OF NEW GENERA AND SPECIES. In Johnson, C. W., Diptera of Florida. Phila. Acad. Nat. Sei. Proc. 1895: 303-340. (70) 1897. REVISION OF THE TACHINIDAE OF AMERICA NORTH OF MEXICO. A FAMILY OF PARASITIC TWO-WINGED INSECTS. U.S. Dept. Agr. Div. Ent. Tech. Ser. 7, pp. 1-156. (71) COSTA, A. 1862. OSSERVAZIONI SU TALUNE SPECIE D'INSETTI STRANIERI ALL'EUROPA. Articola 6°. Napoli Mus. Zool. Ann. 1: 65-71. (72) CREIGHTON, W. S. 1940. A REVISION OF THE NORTH AMERICAN VARIANTS OF THE ANT, FORMICA RUFA. Amer. Mus. Nat. Hist. Amer. Mus. Novitates 1055, 10 pp. (73) 1950. THE ANTS OF NORTH AMERICA. Harvard Mus. Compar. Zool. Bui. 104, 585 pp. (74) CRESSON, E. T. 1865. CATALOGUE OF HYMENOPTERA IN THE COLLECTION OF THE ENTOMOLOG- ICAL SOCIETY OF PHILADELPHIA, FROM COLORADO TERRITORY. Phila. Ent. Soc. Proc. 4: 242-313, 426-488. (75) 1872. HYMENOPTERA TEXANA. Amer. Ent. Soc. Trans. 4: 153-292. 266 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(76) 1872. SYNOPSIS OF THE NORTH AMERICAN SPECIES BELONGING TO THE GENERA LEUCOSPis, SMICRA AND CHALCis. Amer. Ent. Soc. Trans. 4: 29-60. (77) GRIDDLE, N. 1931. GRASSHOPPER CONTROL IN CANADA EAST OF THE ROCKY MOUNTAINS. Canada Dept. Agr. BuL 143, 18 pp. (78) 1933. NOTES ON THE HABITS OF INJURIOUS GRASSHOPPERS IN MANITOBA. Canad. Ent. 65: 97-102. (79) CURRAN, C. H. 1931. THE NEARCTIC SPECIES OF THE NEMESTRINID GENUS RHYNCHOCE- PHALUS FISCHER (DÍPTERA). Cañad. Ent. 63: 68-72. (80) DEBACH, P., and SMITH, H. S. 1941. ARE POPULATION OSCILLATIONS INHERENT IN THE HOST-PARASITE RE- LATION? Ecology 22: 363-369. (81) DEGEER, C. 1752. MéMOIRES POUR SERVIR *A' L'HISTOIRE DES INSECTES, V. 1. Stock- holm. (82) 1775. MéMOIRES POUR SERVIR *A' L'HISTOIRE DES INSECTES. V. 15. Stock- holm. (83) DEJEAN, P. F. M. A.

1825. SPECIES GéNéRAL DES COLéOPTèRES DE LA COLLECTION DE M. LE COMTE DEJEAN. V. 1, 451 pp. Paris. (84) 1829. SPECIES GÉNÉRAL DES COLÉOPTÈRES DE LA COLLECTION DE M. LE COMTE DEJEAN. V. 4, 334 pp. Paris. (85) DILLON, L. S. 1952. THE MELOIDAE (COLEÓPTERA) OF TEXAS. Amer. Midland. Nat. 48: 330-420. (86) and DILLON, E. S. 1961. A MANUAL OF COMMON BEETLES OF EASTERN NORTH AMERICA. 884 pp. New York. (87) DOUTT, R. L. 1964. BIOLOGICAL CHARACTERISTICS OF ENTOMOPHAGOUS ADULTS. In DeBach, P., Biological Control of Insect Pests and Weeds, pp. 145-167. New York. (88) DUFOUR, L. 1841. DES METAMORPHOSES DES CHALCIS, ET DESCRIPTION D'UNE ESPECE PEU CONNUE DE CE GENRE D'HYMENOPTERES. Soc. Ent. Le France Ann. 10, 19 pp. (89) ECKERT, J. E., and MALLIS, A. 1941 (1937). ANTS AND THEIR CONTROL IN CALIFORNIA. Calif. Agr. Expt. Sta. Gir. 342, pp. 1-39. (90) EDWARDS, R. L. 1955. THE HOST-FINDING AND OVIPOSITION BEHAVIOR OF MORMONIELLA VITRIPENNIS (WALKER) (HYM., PTEROMALIDAE) , A PARASITE OF MUSCOID FLIES. Behaviour 7: 88-112. (91) EMERY, G. 1893. BEITRAGE ZUR KENTNISS DER NORDAMERIKANISCHEN AMEISEN-FAUNA. Zool. Jahrb. Abt. f. System. 7: 633-682. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 267

(92) 1895. BEITRAGE ZUR KENTNISS DER NORDAMERIKANISCHEN AMEISEN-FAUNA. Zool. Jahrb. Abt. f. System. 8: 257-360. (93) ESSIG, E. O. 1926. INSECTS OF WESTERN NORTH AMERICA. 1035 pp. New York. (94) 1947. COLLEGE ENTOMOLOGY. 900 pp. New York. (95) EVANS, A. C. 1933. COMPARATIVE OBSERVATIONS ON THE MORPHOLOGY AND BIOLOGY OF SOME HYMENOPTEROUS PARASITES OF CARRION-INFESTING DÍPTERA. Ent. Res. Bui. 24: 385-405. (96) EVANS, H. E. 1958. STUDIES OF THE LARVAE OF DIGGER WASPS (HYMENOPTERA, SPHE« CIDAE) PART IV : ASTATINAE, LARRINAE AND PEMPHREDONINAE. Amer. Ent. Soc. Trans. 84: 109-139. (97) 1958. STUDIES ON THE NESTING BEHAVIOR OF DIGGER WASPS OF THE TRIBE SPHECINI. PT. I. GENUS PRIONONYX DAHLBOM. Ent. Soc. Amer. Ann. 51: 177-186. (98) 1964. FURTHER STUDIES ON THE LARVAE OF DIGGER WASPS (HYMENOPTERA: SPHECIDAE). Amer. Ent. Soc. Trans. 90: 235-299. (99) 1966. THE BEHAVIOR PATTERNS OF SOLITARY WASPS. Ann. Rev. Ent. 11: 123-154. (100) and KURCZEWSKI, F. E. 1966. OBSERVATIONS ON THE NESTING BEHAVIOR OF SOME SPECIES OF TACHYTES (HYMENOPTERA: SPHECIDAE, LARRINAE). Kans. Ent. Soc. Jour. 39: 323-332. (101) and LiN, C. S. 1955. STUDIES ON THE LARVAE OF DIGGER WASPS. (HYMENOPTERA: SPHE- CIDAE). PT. I. SPHECINAE. Amer. Ent. Soc. Trans. 81: 131-153. (102) FABRICIUS, J. C. 1775. SYSTEMA ENTOMOLOGIAE, SISTENS INSECTORUM CLASSES ORDINES, GENERA, SPECIES ADIECTIS SYNONYMUS, LOCIS, DESCRIPTIONIBUS, OBSERVATIONIBUS. 832 pp. Flensburg! et I/epsiae. (103) 1798. SUPPLEMENTUM ENTOMOLOGIAE SYSTEMATICA. 572 pp. Hafniae. (104) 1801. SYSTEMA ELEUTHERATORUM. 2 V. Kiliae. (105) 1805. SYSTEMA ANTLIATORUM SECUNDUM ORDINES, GENERA, SPECIES. 373 pp. Brunsvigae (Brunswick). (106) FALL, H. C. 1901. NOTES ON DICHELONYCHA AND CANTHARIS WITH DESCRIPTIONS OF NEW SPECIES IN OTHER GENERA. Amer. Ent. Soc. Trans. 27: 277- 310. (107) 1922. NEW COLEÓPTERA, X. Cañad. Ent. 54: 170-173. 268 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(108) FALLEN, C. F. 1817. BESKRIFNING öFVER DE I SVERIGEFUNNA FLUGE ARTER, SOM KUNNA FORAS TIL SLÄGTET MUSCA. FÖRSTA AFDELNINGEN. K. Vetensk. Akad. HandL (Ser. 3), 1816: 226-254. 109) 1820. RHIZOMYZIDES SVECIAE. 10 pp. Lundae (Lund). 110) FERNALD, H. T. 1945. A COLONY OF SOLITARY WASPS ( HYMENOPTERA: CHLORIONINI). Ent. Soc. Amer. Ann. 38: 458-460. 111) FORD, N. 1922. AN UNDESCRIBED PLANIDIUM OF PERILAMPUS FROM CONOCEPHALUS (HYM). Canad. Ent. 54: 199-204. 112) FOREL, A. 1886. ESPÈCES NOUVELLES DE FOURMIS AMÉRICAINES. Soc. Ent. de Belg. Ann. (C.R.) 30: 38-49. 113) FöRSTER, J. R. 1771. A CATALOGUE OF THE ANIMALS OF NORTH AMERICA. 143 pp. Lon- don. 114) Fox, W. J. 1892. MONOGRAPH OF THE NORTH AMERICAN SPECIES OF TACHYTES. Amer. Ent. Soc. Trans. 19: 234-252. 115) 1893. THE NORTH AMERICAN LARRIDAE. Phila. Acad. Nat. Sci. Proc. 45: 467-551. 116) FRISCH, J. A. 1937. THE LIFE-HISTORY AND HABITS OF THE DIGGER-WASP AMMOBIA ICH- NEUMONEA (LINN). Amer. Midland Nat. 18: 1043-1062. 117) GAHAN, A. B. 1914. DESCRIPTIONS OF NEW GENERA AND SPECIES, WITH NOTES ON PARA- SITIC HYMENOPTERA. U.S. Nati. Mus. Proc. 48: 155-168. 118) GiLBERTSON, G. I., and HORSFALL, W. R. 1940. BLISTER BEETLES AND THEIR CONTROL. S. Dak. Agr. Expt. Sta. Bui. 340. 23 pp. 119) GiRAULT, A. A., and SANDERS, G. E. 1909. THE CHALCIDOID PARASITES OF THE COMMON HOUSE OR TYPHOID FLY (MUSCA DOMESTICA LINNAEUS) AND ITS ALLIES. Psyche 16: 119- 132. 120) and SANDERS, G. E. 1910. THE CHALCIDOID PARASITES OF THE COMMON HOUSE OR TYPHOID FLY (MUSCA DOMESTICA LINN.) AND ITS ALLIES. Psyche 17: 9-28. 121) GLASER, R. W., and WILCOX, A. M. 1918. ON THE OCCURRENCE OF A MERMIS EPIDEMIC AMONGST GRASSHOPPERS. Psyche 25: 12-15. 122) GRAHAM, K. 1932. VARIATIONS WITHIN THE SPECIES OF THE NEMESTRINID FLY RHYNCO- CEPHALUS SACKENi WILLISTON. Ganad. Ent. 64: 167-168. 123) GREATHEAD, D. J. 1963. A REVIEW OF THE INSECT ENEMIES OF ACRIDOIDAE (ORTHOPTERA) . Roy. Ent. Soc, London, Trans. 114: 437-517. 124) GREENE, C. T. 1926. THE PUPARIA AND LARVAE OF SARCOPHAGID FLIES. U.S. Nati. MuS. Proc. 66 (art. 29), 26 pp. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 269

(125) GREGG, R. E. 1963. THE ANTS OF COLORADO WITH REFERENCE TO THEIR ECOLOGY, TAXON- OMY, AND GEOGRAPHIC DISTRIBUTION. 792 pp. Univ. Colo. PreSS, Boulder. (126) HAGAN, K. S. 1964. DEVELOPMENTAL STAGES OF PARASITES. In DeBach, P., Biological Control of Insect Pests and Weeds, pp. 168-246. New York. (127) HALDEMAN, S. S. 1843. DESCRIPTION OF NORTH AMERICAN SPECIES OF COLEóPTERA, PRE- SUMED TO BE UNDESCRiBED. Phila. Acad. Nat. Sei. Proc. 1: 298- 305. (128) HAMILTON, C. C. 1925. STUDIES ON THE MORPHOLOGY, TAXONOMY, AND ECOLOGY OF THE LARVAE OF HOLARCTIC TIGER-BEETLES (FAMILY CICINDELIDAE) . U.S. Nati. Mus. Proc. 65 (art. 17) (2530), 73 pp. (129) HARTMAN, C. 1905. OBSERVATIONS ON THE HABITS OF SOME SOLITARY WASPS OF TEXAS. Tex. Univ. Bui. Sei. Ser. 6, 72 pp. (130) HATCH, M. H. 1949. STUDIES ON THE FAUNA OF PACIFIC NORTHWEST GREENHOUSES

(ISOPTODA, COLEóPTERA, DERMAPTERA, ORTHOPTERA, GASTROPODA). N.Y. Ent. Soc. Jour. 57: 141-165. (131) 1953. THE BEETLES OF THE PACIFIC NORTHWEST. PT. I. INTRODUCTION AND . V. 16, 340 pp. Univ. Wash. Press, Seattle. (132) HAYES, W. P., and DECOURSEY, J. D. 1938. OBSERVATIONS OF GRASSHOPPER PARASITISM IN 1937. Jour. Econ. Ent. 31: 519-522. (133) HAYWARD, R. 1908. STUDIES IN AMARA. Amer. Ent. Soc. Trans. 34: 13-65. (134) HENTZ, N. M. 1830. DESCRIPTION OF ELEVEN NEW SPECIES OF NORTH AMERICAN INSECTS. Amer. Phil. Soc. Trans. Ser. 2, III, pp. 253-258. (135) HERBST, J. F. W. 1806. NATURSYSTEM. V. 10. Kafer, Berlin. (136) HERRICK, C. A. 1921. A SARCOPHAGID PARASITE OF THE COMMON FIELD CRICKET. Amer. Micros. Soc. Trans. 40: 116-117. (137) HiNE, J. S. 1909. ROBBERFLIES OF THE GENUS ASILUS. Ent. Soc. Amer. Ann. 2: 136- 172. (138) 1911. NEW SPECIES OF DÍPTERA OF THE GENUS ERAX. Ohio Nat. 11: 307- 311. (139) 1916. DESCRIPTIONS OF ROBBERFLIES OF THE GENUS ERAX. Ohio Jour. Sci. 17: 21-22. (140) 1919. ROBBERFLIES OF THE GENUS ERAX. Ent. Soc. Amer. Ann. 12: 103- 154. 270 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(141) HORN, G. H. 1870. CONTRIBUTIONS TO THE COLEOPTEROLOGY OF THE UNITED STATES. Amer. Ent. Soc. Trans. 3: 69-97. (142) 1873. REVISION OF THE SPECIES OF MELOIDAE OF THE UNITED STATES. Amer. Phü. Soc. Proc. 13, pp. 88-117. (143) 1875. SYNONYMICAL NOTES AND DESCRIPTIONS OF NEW SPECIES OF NORTH AMERICAN COLEÓPTERA. Amer. Ent. Soc. Trans. 5: 126-156. (144) 1878. DESCRIPTIONS OF THE LARVAE OF THE NORTH AMERICAN GENERA OF CICINDELIDAE, ALSO OF DICAELUS W^ITH A NOTE ON RHYNCHOPHORUS. Amer. Ent. Soc. Trans. 7: 28-40. (145) 1880. A REVIEW OF THE SPECIES OF ANISODACTYLUS. Amer. PhiL SoC. Proc. 19, pp. 162-178. (146) 1893. AMBLYCHiLA CYLiNDRiFORMis SAY. Ent. News 4: 281-283. (147) HORSFALL, W. R. 1941. BIOLOGY OF THE BLACK BLISTER BEETLE (COLEÓPTERA: MELOIDAE). Ent. Soc. Amer. Ann. 34: 114-126. (148) 1942. BIOLOGY OF THE SQUASH BLISTER BEETLE. Kans. Ent. Soc. Jour. 15: 93-99. (149) 1943. BIOLOGY AND CONTROL OF COMMON BLISTER BEETLES IN ARKANSAS. Ark. Agr. Expt. Sta. BuL 436, 55 pp. (150) HOWARD, C. W. 1918. A PRELIMINARY REPORT ON THE TROMBIDIIDAE OF MINNESOTA. MINN. State Ent. Rpt. 17: 111-144. (151) HOWARD, L. 0. 1897. A STUDY OF INSECT PARASITISM : A CONSIDERATION OF THE PARASITES OF THE WHITE-MARKED TUSSOCK MOTH, WITH AN ACCOUNT OF THEIR HABITS AND INTERRELATIONS, AND WITH DESCRIPTIONS OF NEW SPECIES. U.S. Dept. Agr. Div. Ent. Tech. Ser. 5, 57 pp. (152) HUGGANS, J. L., and BLICKENSTAFF, C. C. 1966. PARASITES AND PREDATORS OF GRASSHOPPERS IN MISSOURI. Mo. Agi. Expt. sta. Res. Bui. 903, 40 pp. (153) HUNGERFORD, H. B., and WILLIAMS, F. X. 1912. BIOLOGICAL NOTES ON SOME KANSAS HYMENOPTERA. Ent. NeWS 23: 241-260. (154) HUNTER, S. J. 1898. PARASITIC INFLUENCES ON MELANOPLUS. Kans. Univ. Quart. Ser. A: 205-210. (155) 1899. ALFALFA, GRASSHOPPERS, BEES; THEIR RELATIONSHIP. A REPORT OF THE FIELD WORK OF THE DEPARTMENT OF ENTOMOLOGY, SUMMER OF 1898. Kans. Univ. Dept. Ent. Rpt. 65, 152 pp. (156) JAMES, M. T. 1933. NEW ASILIDAE FROM COLORADO. Amer. Mus. Nat. Hist. Amer. Mus. Novitates 596, 3 pp. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 271

(157) 1938. A SYSTEMATIC AND ECOLOGICAL STUDY OF THE ROBBER FLIES (ASILIDAE) OF COLORADO. In Lester, O. C, ed., Abstracts of Theses and Reports for Higher Degrees, 1938. Colo. Univ. Studies 26, pp. 70-74. (158) 1941. THE ROBBER FLIES OF COLORADO (DÍPTERA, ASILIDAE). Kans. Ent. Soc. Jour. 14: 27-53. (159) JAQUES, H. E. 1951. HOW TO KNOW THE BETTLES. 372 pp. Dubuque, Iowa. (160) KANSAS STATE COLLEGE. 1962. INSECTS IN KANSAS. 307 pp. Kans. State Col., Topeka. (161) KASTON, B. J. 1953. HOW TO KNOW THE SPIDERS. 220 pp. Dubuque, Iowa. (162) KEILIN, D. 1924. ON A CASE OF INTESTINAL MYLIASIS IN MAN PRODUCED BY THE LARVAE OF A SARCOPHAGINE FLY. Parasitology 16: 318-320. (163) KELLOGG, V. L. 1914. AMERICAN INSECTS. 694 pp. New York. (164) KELLY, E. 0. G. 1914. A NEW SARCOPHAGID PARASITE OF GRASSHOPPERS. Jour. Agr. ReS. 2: 435-446. (165) KING, P. E., and RICHARDS, J. G. 1969. OOGÉNESIS IN NASONIA VITRIPENNIS (WALKER) (HYMENOPTERA : PTEROMALIDAE). Roy. Ent. Soc, London, Proc. 44: 143-157. (166) KiRBY, W. 1837. THE INSECTS. In Richardson, J., Fauna Boreali-Americana. Pt. 4, 325 pp. London. (167) KNIPLING, E. F. 1936. A COMPARATIVE STUDY OF THE FIRST-INSTAR LARVAE OF THE GENUS SARCOPHAGA (CALLIPHORIDAE, DÍPTERA), WITH NOTES ON THE BIOL- OGY. Jour. Parasitol. 22: 417-454. (168) KNOWLTON, G. F., and JANES, M. J. 1931. NOTES ON SOME BENEFICIAL UTAH DÍPTERA. Utah Acad. Sci. PrOC. 8: 147-148. (169) KROMBEIN, K. V. 1948. NEW PREY RECORDS IN OXYBELUS (HYMENOPTERA,. SPHECIDAE) Wash. Ent. Soc. Proc. 50: 67. (170) 1960. BIOLOGICAL NOTES ON SEVERAL SOUTHWESTERN GROUND-NESTING WASPS (HYMENOPTERA, SPHECIDAE). Brooklyn Ent. Soc. Bui. 55: 75-79. (171) KuiTERT, L. C, and CONNIN, R. V. 1952. BIOLOGY OF THE AMERICAN GRASSHOPPER IN SOUTHEASTERN UNITED STATES. Fla. Ent. 35: 22-33. (172) KURCZEWSKi, F. E. 1966. TACHYSPHEX TERMINATUS PREYING ON TETTIGONIIDAE—AN UNUSUAL RECORD (HYMENOPTERA: SPHECIDAE, LARRINAE) . KaUS. Ent. SoC. Jour. 39: 317-322. 272 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(173) 1966. COMPARATIVE BEHAVIOR OF MALE DIGGER WASPS OF THE GENUS TACHY- SPHEX (HYMENOPTERA: SPHECIDAE, LARRINAE) . Kans. Ent. Soc. Jour. 39: 436-453. (174) 1967. HEDYCHRIDIUM FLETCHERI (HYMENOPTERA: CHRYSIDIDAE, ELAM- PINAE), A PROBABLE PARASITE OF TACHYSPHEX SIMILIS (HYMENOP- TERA: SPHECIDAE, LARRINAE). Kans. Ent. Soc. Jour. 40: 278-284. (175) LA FERTé-SéNECTèRE, M. LE MARQUIS DE. 1841. DESCRIPTION DE DIX CARABIQUES NOUVEAUX DU TEXAS ET D'UNE ESPÈCE NOUVELLE DE BUPRESTIDE DE FRANCE. ReV. et Mag. de Zool. Pure et Appl. 1841: 37-50. (176) LANHAM, U. N., and EVANS, F. C. 1958. PHORETIC SCELIONIDS ON GRASSHOPPERS OF THE GENUS MELANOPLUS (HYMENOPTERA: SCELIONIDAE). Pan-Pacific Ent. 34: 213-214. (177) and EVANS, F. C. 1960. OBSERVATIONS ON THE SCELIONID COMPONENT OF A GRASSLAND INSECT FAUNA. Brooklyn Ent. Soc. Bui. 55: 84-87. (178) LA RIVERS, I. 1945. THE WASP CHLORION LAEVIVENTRIS AS A NATURAL CONTROL OF THE MORMON CRICKET. Amer. Midland Nat. 33: 743-763. (179) 1949. ENTOMIC NEMATODE LITERATURE FROM 1926 TO 1946, EXCLUSIVE OF MEDICAL AND VETERINARY TITLES. Wasmann Collector 7, pp. 177- 206. (180) LAVIGNE, R. J. 1963. NOTES ON THE BEHAVIOR OF STENOPOGON COYOTE BROMLEY WITH A DESCRIPTION OF THE EGGS. Pan-Pacific Ent. 39: 103-107. (181) 1964. NOTES ON THE DISTRIBUTION AND ETHOLOGY OF EFFERIA BICAUDATA (DíPTERA: ASILIDAE), WITH A DESCRIPTION OF THE EGGS. Ent. Soe. Amer. Ann. 57: 341-344. (182) and HOLLAND, F. R. 1969. COMPARATIVE BEHAVIOR OF ELEVEN SPECIES OF WYOMING ROBBER FLIES (DíPTERA: ASILIDAE). Wyo. Agr. Expt. Sta. Sei. Monog. 18, 61 pp. (183) — and PFADT, R. E. 1966. PARASITES AND PREDATORS OF WYOMING RANGELAND GRASSHOPPERS. Wyo. Agr. Expt. Sta. Sei. Monog. 3, 31 pp. (184) LECONTE, J. L. 1848. MONOGRAPH OF THE SPECIES OF PASIMACHUS INHABITING THE UNITED STATES. Lyceum Nat. Hist. Ann. 4: 141-163. (185) 1848. A DESCRIPTIVE CATALOGUE OF GEODELPHAGOUS COLEPTERA. Lyceum Nat. Hist. Ann. 4: 173-474. (186) 1850. GENERAL REMARKS ON COLEÓPTERA. Agassiv Lake Superior, Bos- ton 4, pp. 201-242. (187) 1851. DESCRIPTIONS OF NEW SPECIES OF COLEÓPTERA FROM CALIFORNIA. Lyceum Nat. Hist. Ann. 5: 125-216. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 273

(188) 1853. SYNOPSIS OF MELOIDS OF THE UNITED STATES. Phila. Acad. Nat. Sei. Proc. 6: 328-350. (189) 1859. THE COLEÓPTERA OF KANSAS AND EASTERN NEW MEXICO. Smithsn. Contrib. to Knowledge No. 126, 58 pp. (190) 1865. NOTE ON THE SPECIES OF HARPALUS. Phila. Acad. Nat. Sei. Proe. 17: 98-104. (191) 1866. NEW SPECIES OF NORTH AMERICAN COLEÓPTERA. PT. I. Smithsn. Mise. Colleet. 4, No. 167, pp. 1-177. (192) 1869. AMERICAN ENTOMOLOGY. A DESCRIPTION OF THE INSECTS OF NORTH AMERICA BY THOMAS SAY, WITH ILLUSTRATIONS DRAWN AND COLORED AFTER NATURE. V. 1, 412 pp. New York. (193) 1869. AMERICAN ENTOMOLOGY. A DESCRIPTION OF THE INSECTS OF NORTH AMERICA BY THOMAS SAY, WITH ILLUSTRATIONS DRAWN AND COLORED AFTER NATURE. V. 2, 814 pp. New York. (194) and HORN, G. H. 1883. CLASSIFICATION OF THE COLEÓPTERA OF NORTH AMERICA. Smithsn. Misc. Colleet. 26 (4), 567 pp. Washington, B.C.

(195) LENG, C. W. ._ 1920. CATALOGUE OF THE COLEÓPTERA OF AMERICA, NORTH OF MEXICO. 47U pp. Cambridge, Mass. (196) 1939. CATALOGUE OF THE COLEÓPTERA OF AMERICA, NORTH OF MEXICO. 4th sup., 146 pp. New York. (197) and BEUTENMüLLER, W. 1894 PRELIMINARY HAND-BOOK OF THE COLEÓPTERA OF NORTH EASTERN AMERICA. N.Y. Ent. Soe. Jour. 2, pp. 42-48, 87-96, 133-141, 175- 190. (198) LEPELETIER, A. DE S-F. . . 1845. HISTOIRE NATURELLE DES INSECTES. HYMENOPTERES. In Librairie Eneyelopedique de Roret. V. 3, 355 pp. Paris. (199) LIN, C. S. 1963. PREY-SELECTION AND OVIPOSITING BEHAVIOR OF SPHECID WASPS (HY- MENOPTERA: ACULEATA). Internatl. Cong. Zool. Proc. 16, p. 5. (200) 1967. NESTING BEHAVIOR OF TACHYTES (TACHYPLENA) PRAEDATOR FOX, WITH A REVIEW OF THE BIOLOGY OF THE GENUS (HYMENOPTERA: SPHECIDAE: LARRINAE). Amer. Midland Nat. 77: 241-245.

(201) LINNAEUS, C. 1758. SYSTEMA NATURAE PER REGNA TRIA NATURAE. Ed. 10, V. 1, 824 pp. Holmiae (Stockholm). (202) 1763. AMOENITATES ACADEMICAE, SEU DISSERTATIONES VARIAE PHYSICAE, MEDICAE, BOTANICAE, ANTEHAC SEORSIM EDITAE, NUNC COLLECTAE ET AUCTAE CUM TABULis. V. 6, 486 pp. Holmiae (Stockholm). 274 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(203) LLOYD, D. C. 1951. A SURVEY FOR GRASSHOPPER PARASITES IN TEMPERATE SOUTH AMER- ICA. Ganad. Ent. 83: 213-230. (204) LOEW, H. 1863. DíPTERA AMERICAE SEPTENTRIONALIS INDíGENA. Centuria Quarta. Berlin. Ent. Ztschr. 7: 275-326. (205) 1866. DíPTERA AMERICAE SEPTENTRIONALIS INDíGENA. Centuria Séptima. Berlin. Ent. Ztschr. 10: 1-54. (206) 1872. DíPTERA AMERICAE SEPTENTRIONALIS INDíGENA. Centuria Decima. Berlin. Ent. Ztschr. 16: 49-115. (207) 1874. NEU NORDAMERIKANISCHE DÍPTERA. Berlin. Ent. ztschr. 18: 353- 384. (208) LUGGER, 0. 1891. MIGRATORY LOCUSTS IN MINNESOTA IN 1891. Minn. Agr. Expt. Sta. Bui. 17, pp. 87-108. (209) MACQUART, J. 1834. HISTOIRE NATURELLE DES INSECTES—DÍPTERAS. Tome Premier, Diptera, v. 1. In Roret, N. E., ed.. Collection des Suites à Buffon, q.v., 578 pp. Paris. (210) 1838. DIPTÈRES EXOTIQUES NOUVEAUX OU PEU CONNUS. V. 1 pt 2 pp 5-207. Paris. * ' (211) 1840. DIPTÈRES EXOTIQUES NOUVEAUX OU PEU CONNUS. V. 2, pt. 1, pp. 5-135. Paris. (212)

1846. DIPTÈRES EXOTIQUES NOUVEAUX OU PEU CONNUS. (LER) SUP. SOC. Roy. des Sei. de l'Agr. et des Arts, Lille, Mém. (1845) 1844. pp. 133-364. (213) 1848. DIPTÈRES EXOTIQUES NOUREAUX OU PEU CONNUS. Suite de 2me sup. Soc. Roy. des Sei. de l'Agr. et des Arts, Lille, Mém. 1847 (2), pp. 161-237. (214) MACSWAIN, J. W. 1956. A CLASSIFICATION OF THE FIRST INSTAR LARVAE OF THE MELOIDAE (COLEÓPTERA). Calif. Univ. Pubs., Ent. 12, pp. 1-182. (215) MANNERHEIM, G. C. G. VON. 1843. BEITR. Z. KAFERF. DER ALEUTISCHEN INS. . . . Moscou Bul. 16. PP. 175-314. (216) MASSEY, W. F. 1893. THE CULTURE OF ORCHARD AND GARDEN FRUITS. N.C. Agr. Expt Sta. Bul. 92, 144 pp. (217) MEIGEN, J. W. 1826. SYSTEMATISCHE BESCHREIBUNG DER BEKANNTEN EUROPAISCHEN ZWEIFLUGELIGEN INSEKTEN. V. 5, 412 pp. Hamm (218) 1830. SYSTEMATISCHE BESCHREIBUNG DER BEKANNTEN EUROPAISCHEN ZWEIFLUGELIGEN INSEKTEN. V. 6, 401 pp. Hamm. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 275

(219) MERWE, J. S. VEN DER. 1843. INVESTIGATIONS ON THE BIOLOGY AND ECOLOGY OF MORMONIELLA VITRIPENNIS WALKER (PTEROMALIDAE! HYMENOPTERA) . So. Africa Ent. Soc. Jour. 6: 48-61. (220) MiDDLEKAUFF, W. W. 1958. BIOLOGY AND ECOLOGY OF SEVERAL SPECIES OF CALIFORNIA RANGELAND GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE) . Pan-Pacific Ent. 34: 1-11. (221) 1959. SOME BIOLOGICAL OBSERVATIONS ON SARCOPHAGA FALCIFORMIS, A PARASITE OF GRASSHOPPERS (DÍPTERA: SARCOPHAGIDAE) . Ent. Soc. Amer. Ann. 52: 724-728. (222) MiLLIKEN, F. B. 1921. RESULTS OF WORK ON BLISTER BEETLES IN KANSAS. U.S. Dept. Agr. Bul. 967, 26 pp. (223) MOORE, R. 1906. NOTES OF THE HABITS OF CICINDELA. Ent. News 17: 338-343. (224) MORGAN, H. A. 1901. SOME MISCELLANEOUS RESULTS OF THE WORK OF THE DIVISION OF ENTOMOLOGY. V. THE DIFFERENTIAL GRASSHOPPER IN THE MISSIS- SIPPI DELTA. . . . OTHER COMMON SPECIES. U.S. Bur. Ent. Bul. (n.s.) 30: 7-33. (225) MUESEBECK, C. F. W., KROMBEIN, K. V., and TOWNES, H. K. 1951. HYMENOPTERA OF AMERICA NORTH OF MEXICO. SYNOPTIC CATALOG. U.S. Dept. Agr. Monog. 2, 1420 pp. (226) NEWCOMER, E. J. 1930. NOTES ON THE HABITS OF A DIGGER WASP AND ITS INQUILINE FLIES. Ent. Soc. Amer. Ann. 23: 552-563. (227) NEWTON, R. C. 1954. NEW RECORDS OF DIPTEROUS PARASITES OF GRASSHOPPERS. Jour. Econ. Ent. 47: 935-936. (228) 1956. DIGGER WASPS, TACHYSPHEX SPP., AS PREDATORS OF A RANGE GRASS- HOPPER IN IDAHO. Jour. Econ. Ent. 49: 615-619. (229) 1960. THE NESTING HABITS OF THE WASP, SOLIERELLA SP., AND THE LOCA- TION OF ITS EGG ON THE GRASSHOPPER, AGENEOTETTIX DEORUM. Jour. Econ. Ent. 53: 958-959. (230) OLIVER, A. G. 1790. ENTOMOLOGIE II. 485 pp. Paris. (231) OSTEN SACKEN, C. R. 1877. WESTERN DíPTERA: DESCRIPTIONS OF NEW GENERA AND SPECIES OF DíPTERA FROM THE REGION WEST OF THE MISSISSIPPI AND ESPECIALLY FROM CALIFORNIA. U.S. Geol. and Geog. Survey of Ter. Bul. 3, pp. 189-354. (232) 1887. DíPTERA. V. 1. In Godman, F. D., and Salvin, 0., eds.. Biología Centrali-Americana (q.v.). Zoologia-Insecta-Diptera. V. 1, 378 pp. London. 276 TECHNICAL BULLETIN NO. 14Ó0, U. S. DEPT. OF AGRICULTURE

(233) PANDELLE, L. 1898. ÉTUDES SUR LES MUSCIDES DE FRANCE. Ill" PARTIE. ReV. d'Ent (Caen) 17, pp. 1-244. (234) PARISH, H. E., and GUSHING, E. C. 1938. LOCATIONS FOR BLOWFLY TRAPS : ABUNDANCE AND ACTIVITY OF BLOW- FLIES AND OTHER FLIES IN MENARD COUNTY, TEXAS. Jour Econ Ent. 31: 750-763. (235) PARKER, H. L. 1924. RECHERCHES SUR LES FORMES POST-EMBRYONNAIRES DES CHALCIDIENS. Soc. Ent. de France Ann. 93: 261-379. (236) PARKER, J. B. 1915. NOTES ON NESTING HABITS OF SOME SOLITARY WASPS. Wash. Ent See. Proc. 7: 70-76. (237) PARKER, J. R., and WAKELAND, G. 1957. GRASSHOPPER EGG PODS DESTROYED BY LARVAE OF BEE FLIES, BLISTER BEETLES, AND GROUND BEETLES. U.S. Dept. Agr. Tech. Bul 1165 29 pp. (238) PARKER, R. R. 1923. NEW SARCOPHAGIDAE FROM ASIA, WITH DATA RELATING TO THE DUX GROUP. Mag. Nat. Hist, and Ann., Ser. 9, 11: 123-129 (239) PATTON, W. H. 1879. DESCRIPTION OF A NEW FOSSORiAL WASP. Ganad. Ent 11 • 133-134 (240) PAUL, L. C., and PUTNAM, L. G. 1960. MORPHOMETRICS, PARASITES AND PREDATORS OF MIGRANT MELANOPLUS BILITRATUS (WLK.) (ORTHOPTERA: ACRIDIDAE) IN SASKATCHEWAN IN 1940. Ganad. Ent. 92: 488-493. (241) PEARCE, E. K. 1915. TYPICAL FLIES. A PHOTOGRAPHIC ATLAS OF DÍPTERA, INCLUDING APHANIPTERA. 47 pp. Cambridge, Mass. (242) PECKHAM, G. W., and PECKHAM, E. G. 1898. INSTINCTS AND HABITS OF SOLITARY WASPS. Wis. Geol. and Nat. Hist. Survey BuL 2, 245 pp. (243) and PECKHAM, E. G. 1900. ADDITIONAL OBSERVATIONS ON THE INSTINCTS AND HABITS OF THE SOLITARY WASPS. Wis. Nat. Hist. See. Bul. (n.s.) 1, pp. 85-93. (244) and PECKHAM, E. G. 1905. WASPS SOCIAL AND SOLITARY. 310 pp. Boston and New York (245) PETERSON, A. 1960. LARVAE OF INSECTS. PT. II. COLEÓPTERA, DÍPTERA, NEUROPTERA, SIPHONAPTERA, MECOPTERA, TRICHOPTERA. 416 pp. Columbus, Ohio (246) 1962. LARVAE OF INSECTS. PT. I. LEPIDOPTERA AND HYMENOPTERA. 315 pp. Columbus, Ohio. (247) PRESCOTT, H. W. 1955. NEORHYNCHOCEPHALUS SACKENII AND TRICHOPSIDEA CLAUSA, NEME- STRINID PARASITES OF GRASSHOPPERS. Ent. Soc. Amer. Ann 48- 392-402. (248) 1960. SUPPRESSION OF GRASSHOPPERS BY NEMESTRINID PARASITES (DIP- TERA). Ent. Soc. Amer. Ann. 53: 513-521. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 277

(249) 1961. RESPIRATORY PORE CONSTRUCTION IN THE HOST BY THE NEMESTRINID PARASITE NEORHYNCHOCEPHALUS SACKENII (DIlfTERA) , WITH NOTES ON RESPIRATORY TUBE CHARACTERS. Ent. Soc. Aiiner. Ann. 54: 557- 566. (250) PROVANCHER, M. L. . ^ J XT *. 1881. FAUNE CANADIENNE LES INSECTES—HYMéNOPTèRES. Cañad. Mat. 12, pp. 193-207, 225-241, 257-269, 291-304, 321-333, 353-362.

1882 FAUNE CANADIENNE LES INSECTES—HYMÉNOPTÈRES. Canad. Nat. 13, pp. 4-15, 33-51, 65-81, 97-110, 129-144, 161-175, 193-209, 225- 242, 257-269, 289-311, 321-336, 353-368. (252) RAU, P. a^ T • 1922. ECOLOGY AND BEHAVIOR NOTES ON MISSOURI INSECTS. bt. Í.0U1S Acad. Sei. Trans. 24, pp. 1-71. (253) 1923. A NOTE ON THE NESTING HABITS OF TACHYTES DISTINCTUS. Smith. Psyche 30: 220-221. (254) 1926. THE ECOLOGY OF A SHELTERED CLAY BANK; A STUDY OF INSECT SO- CIOLOGY. St. Louis Acad. Sei. Trans. 25, pp. 157-260. (255)- 1934. BEHAVIOR NOTES ON CERTAIN SOLITARY V7ASPS. Canad. Ent. 66: 259-261. (256) 1946. NOTFS ON THE BEHAVIOR OF A FEW SOLITARY WASPS. Brooklyn Ent. Soc. Bui. 41: 10-11. (257) and RAU, N. 1918. WASP STUDIES AFIELD. 372 pp. London. (258) REHN, J. A. G., and REHN, J. W. H. 1938. THE POST-OAK LOCUST (DENDROTETTIX QUERCUS) IN THE EASTERN UNITED STATES, WITH NOTES ON MACROPTERISM IN THE SPECIES (ORTHOPTERA: ACRIDIDAE). Amer. Ent. Soc. Trans. 64: 79-95. (259) RiLBY, C. V. 1872. FOURTH ANNUAL REPORT ON THE NOXIOUS, BENEFICIAL AND OTHER INSECTS OF THE STATE OF MISSOURI. Mo. State Bd. Agr. Ann. Rpt. 7: 1-145. (260) 1873. FIFTH ANNUAL REPORT ON THE NOXIOUS, BENEFICIAL AND OTHER INSECTS OF THE STATE OF MISSOURI. Mo. State Bd. Agr. Ann. Rpt. 8: 1-160. (261) 1874. DESCRIPTIONS AND NATURAL HISTORY OF TWO NEW SPECIES WHICH BRAVE THE DANGERS OF SARRACENIA VARIOLARIS. St. Louis Acad. Sei. Trans. 3: 235-240. (262) 1875. SEVENTH ANNUAL REPORT ON THE NOXIOUS, BENEFICIAL AND OTHER INSECTS OF THE STATE OF MISSOURI. Mo. State Bd. Agr. Ann. Rpt. 10: 1-196. (263) 1877. THE LOCUST PLAGUE IN THE UNITED STATES. 236 pp. Chicago. 278 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(264) 1878. INVERTEBRATE ENEMIES. In First Annual Report of the United States Entomological Commission for the Year 1877 Relating to the Rocky Mountain Locust, pp. 284-334. Washington, D.C. (265) 1880. FURTHER FACTS ABOUT THE NATURAL ENEMIES OF LOCUSTS. In Sec- ond Report of the United States Entomological Commission for the Years 1878 and 1879, Relating to the Rocky Mountain Locust, pp. 259-271. Washington, D.C. (266) 1893. SOME INJURIOUS INSECTS OF MARYLAND. Md. Agr. Expt. Sta. Bul. 23, pp. 71-93. (267) RiSTlCH, S. S. 1953. A STUDY OF THE PREY, ENEMIES, AND HABITS OF THE GREAT GOLDEN DIGGER WASP CHLORION ICHNEUMONEUM (L.). Canad. Ent. 85: 374-386. (268) ROBERTS, R. A. 1933. ACTIVITY OF BLOWFLIES AND ASSOCIATED INSECTS AT VARIOUS HEIGHTS ABOVE THE GROUND. Ecology 14: 306-314. (269) 1933. BIOLOGY OF BRACHYMERIA FONSCOLOMBEI (DUFOUR) , A HYMENOPTER- ous PARASITE OF BLOWFLY LARVAE. U.S. Dept. Agr. Tech. Bul. 365, 21 pp. (270) ROBINEAU-DESVOïDY, J. B. 1830. ESSAI SUR LES MYODAIRES. (Paris) Inst, de France, (Cl. Des) Sei. Math, et Phys., Acad. Roy. des Sei., Mém. Présentés par Divers Savans (Ser. 2) 2, pp. 1-813. (271) ROEMHILD, G., and ANDERSON, N. L., JR. 1957. GRASSHOPPER CONTROL HANDBOOK FOR 1957. Mont. State Col. Ext. Serv. Pub. E-239, 24 pp. (272) RoEPKE, W. 1917. EENIGE OPMERKINGEN OVER TWEEN JAVAANSCHE CANTHARIDEN : MY- LABRIS PUSTULATA THUNB. EN EPICAUTA RUFICEPS ILL. Tijdschr. V. Ent. 60: 252-267. (273) ROHWER, S. A. 1908. SOME LARRiD WASPS FROM COLORADO. Ent. News 19: 220-224. (274) 1909. NEW HYMENOPTERA FROM WESTERN UNITED STATES. Amer. Ent. Soc. Trans. 35: 99-136. (275) 1910. SOME NEW WASPS FROM NEW JERSEY. Wash. Ent. Soc. Proc. 12: 49-52. (276) ROMANOW, W. 1954. PREDATORS OF GRASSHOPPERS. Manitoba Ent. Soc. Proc. 9: 14-18. (277) ROOT, F. M. 1923. NOTES ON LARVAL CHARACTERS IN THE GENUS SARCOPHAGA. Jour. Parasitol. 9: 227-229. (278) Ross, H. H. 1959. A TEXTBOOK OF ENTOMOLOGY. 519 pp. New York. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 279

(279) SABROSKY, C. W. 1967. TWO MEN AND ECONOMICALLY SIGNIFICANT CHLOROPIDAE, WITH THE DESCRIPTION OF A NEW GENUS (DÍPTERA). KanS. Ent. Soc. Jour. 40,: 151-156. (280) SACHAROV, N. L. 1913. REPORT ON THE ACTIVITIES OF THE ENTOMOLOGICAL STATION IN 1912. 25 pp. Astrakhan, U.S.S.R. (281) ST. AMAND, W., and CLOYD, W. J. 1954. PARASITISM OF THE GRASSHOPPER, CHORTOPHAGA VIRIDIFASCIATA (DEGEER) (ORTHOPTERA: LOCUSTIDAE), BY DIPTEROUS LARVAE. Jour. Parasitol. 40: 83-87. (282) SAY T. 1817. AMERICAN ENTOMOLOGY. V. I (i), 18 pp. Philadelphia. (Fore- runner of his famous three volume work.) (283) 1818. A MONOGRAPH OF NORTH AMERICAN INSECTS OF THE GENUS CICIN- DELA. Amer. Phil. Soc. Trans, (n.s.) 1, pp. 401-426. (284) 1823. A DESCRIPTION OF SOME NEW SPECIES OF HYMENOPTEROUS INSECTS. West. Quart. Rpt., Med. Surg. and Nat. Sei. 2: 71-82. (285) 1823. DESCRIPTIONS OF COLEOPTEROUS INSECTS COLLECTED IN THE LATE EXPEDITION TO THE ROCKY MOUNTAINS, PREFORMED BY ORDER OF MR. CALHOUN, SECRETARY OF WAR, UNDER THE COMMAND OF MAJOR LONG. Phila. Acad. Nat. Sei. Jour. 3: 139-216. (286) ^, ., 1823. DESCRIPTIONS OF DIPTEROUS INSECTS OF THE UNITED STATES. Phlia. Acad. Nat. Sei. Jour. 3: 9-54, 73-104. (287) 1823. DESCRIPTIONS OF INSECTS OF THE FAMILIES OF CARABICI AND HYDRO- CANTHARI OF LATREILLE, INHABITING NORTH AMERICA. Amer. Phil. Soc. Trans, (n.s.) 2, pp. 1-109. (288) 1824. AMERICAN ENTOMOLOGY, OR DESCRIPTIONS OF INSECTS OF NORTH AMERICA. V. 1, 101 pp. Philadelphia. (289) 1824. APPENDIX TO THE NARRATIVE OF AN EXPEDITION TO THE SOURCE OF ST. PETER'S RIVER, LAKE WINNEPECK, LAKE OF THE WOODS. ... UN- DER THE COMMAND OF STEPHEN H. LONG, U.S.T.E. PT. I. NATURAL HISTORY. 1. ZOOLOGY. E. CLASS INSECTA. In Keating, W. H., Major Long's Second Expedition (q.v.), v. 2, pp. 268-378. Philadelphia. (290) 1824. DESCRIPTIONS OF COLEOPTEROUS INSECTS COLLECTED IN THE LATE EX- PEDITION TO THE ROCKY MOUNTAINS, PREFORMED BY ORDER OF MR. CALHOUN, SECRETARY OF WAR, UNDER THE COMMAND OF MAJOR LONG. Phila. Acad. Nat. Sei. Jour. 3: 216-462. (291) 1828. A DESCRIPTION OF SOME NEW SPECIES OF HYMENOPTERA OF THE UNITED STATES. Contributions of the Maclurian Lyceum to the Arts and Sciences. V. 1, 67-83. Philadelphia. 280 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(292) 1828. AMERICAN ENTOMOLOGY, OR DESCRIPTIONS OF INSECTS OF NORTH AMERICA. V. 3, 136 pp. Philadelphia. (293) 1834. DESCRIPTIONS OF NEW NORTH AMERICAN INSECTS, AND OBSERVATIONS ON SOME ALREADY DESCRIBED. Amer. Phil. Soc. Trans, (n.s.) 4, pp. 409-470. (294) 1837. DESCRIPTIONS OF NEW NORTH AMERICAN HYMENOPTERA AND OBSER- VATIONS ON SOME ALREADY DESCRIBED. Boston Jour. Nat. Hist. 1: 361-416. (295) SCHLINGER, E. I., and DOUTT, R. L. 1964. SYSTEMATICS IN RELATION TO BIOLOGICAL CONTROL. In DeBach, P., Biological Control of Insect Pests and Weeds, pp. 247-280. New York. (296) SELANDER, R. B. 1960. BIONOMICS, SYSTEMATICS AND PHYLOGENY OF LYTTA, A GENUS OF BLISTER BEETLES (COLEÓPTERA, MELOIDAE) . 111. Biol. Monog. 28, 295 pp. (297) SEVERIN, H. C. 1926. THE COMMON BLACK FIELD CRICKET, GRYLLUS ASSIMILIS (FAB.) AND ITS CONTROL. Jour. Econ. Ent. 19: 218-227. (298) 1944. THE GRASSHOPPER MITE EUTROMBIDIUM CRIGONUM (HERMENN) AN IMPORTANT ENEMY OF GRASSHOPPERS. S. Dak. Agr. Expt. Sta. Tech. Bui. 3, pp. 1-36. (299) SHOTWELL, R. L. 1930. A STUDY OF THE LESSER MIGRATORY GRASSHOPPER. U.S. Dept. Agr. Tech. Bui. 190, 35 pp. (300) SMITH, C. W. 1940. AN EXCHANGE OF GRASSHOPPER PARASITES BETWEEN ARGENTINA AND CANADA WITH NOTES ON PARASITISM OF NATIVE GRASSHOPPERS. Ent. Soc. Ontario Ann. Rpt. 70: 57-62. (301) SMITH, D. S. 1959. NOTE ON DESTRUCTION OF GRASSHOPPER EGGS BY THE FIELD CRICKET ACHETA ASSIMILIS LUCTUOSUS (SERVILLE) ( ORT H OPTERA : GRYLLIDAE) . Ganad. Ent. 91: 127. (302) SMITH, P. 1856. CATALOG OF THE HYMENOPTERA OF THE BRITISH MUSEUM. V. 4, 366 pp. London. (303) SMITH, H. E. 1915. THE GRASSHOPPER OUTBREAK IN NEW MEXICO DURING THE SUMMER OF 1913. U.S. Dept. Agr. BuL 293, 12 pp. (304) SMITH, H. S. 1912. TECHNICAL RESULTS FROM THE GYPSY MOTH PARASITE LABORATORY. IV. THE CHALCIDOID GENUS PERILAMPUS AND ITS RELATION TO THE PROBLEM OF PARASITE INTRODUCTION. U.S. Dept. Agr. Bur. Ent. Tech. Ser. 19, 67 pp. (305) 1917. THE HABIT OF LEAF-OVIPOSITION AMONG THE PARASITIC HYMENOP- TERA. Psyche 24: 63-68. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 281

(306) SMITH, R. W. 1944. OBSERVATIONS ON PARASITES OF SOME CANADIAN GRASSHOPPERS. Canad. Ent. 76: 28-33. (307) 1958. PARASITES OF NYMPHAL AND ADULT GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE) IN WESTERN CANADA. Canad. Jour. Zool. 36: 217-262. (308) 1965. A FIELD POPULATION OF MELANOPLUS SANGUINIPES (FAB.) AND ITS PARASITES. Canad. Jour. Zool. 43: 179-201. (309) and FINLAYSON, T. U. 1950. LARVAE OF DIPTEROUS PARASITES OF NYMPHAL AND ADULT GRASS- HOPPERS. Canad. Jour. Res. D 28: 81-117. (310) SMITH, T. C, and KELLY, E. G. 1937. THE SIXTH ANNUAL INSECT POPULATION SUMMARY OF KANSAS COVER- ING THE YEAR 1936. Kans. Ent. See. Jour. 10: 113-132. (311) SNYDER, F. M. 1940. THE GENUS ACRIDOMYIA STACKELBERG IN NORTH AMERICA (DÍP- TERA: MUSCIDAE). Amer. Mus. Nat. Hist. Amer. Mus. Novitates 1076, 2 pp. (312) SPENCER, G. J. 1931. THE OVIPOSITION HABITS OF RHYNCOCEPHALUS SACKENII WILL., (DÍP- TERA : NEMESTRINIDAE) . Brit. Columbia Ent. Soc. Proc. 28: 21-24. (313) 1932. FURTHER NOTES ON RHYNCOCEPHALUS SACKENI WILL. (DÍPTERA: NEMESTRINIDAE). Brit. Columbia Ent. Soc. Proc. 29: 25-27. (314) 1934. NOTES ON RHYNCOCEPHALUS SACKENI, WILL., A CORRECTION (NEMES- TRINIDAE: DÍPTERA). Canad. Ent. 66: 87. (315) 1945. A NOTE ON THE TANGLEWING FLIES OF BRITISH COLUMBIA (DÍPTERA: NEMESTRINIDAE). Brit. Columbia Ent. Soc. Proc. 42: 18. (316) 1958. THE NATURAL CONTROL COMPLEX AFFECTING GRASSHOPPERS IN THE DRY BELT OF BRITISH COLUMBIA. 10th Internatl. Cong. Ent. Proc. (Montreal 1956) 4, pp. 497-502. (317) 1958. ON THE NEMESTRINIDAE OF BRITISH COLUMBIA DRY RANGE LANDS. 10th Internatl. Cong. Ent. Proc. (Montreal 1956) 4, pp. 503-509. (318) and BUCKELL, E. R. 1957. ON THE ACRIDIOPHAGOUS SARCOPHAGIDAE OF BRITISH COLUMBIA WITH RECORDS OF ALL OTHERS TAKEN IN THE PROVINCE. Brit. Columbia Ent. Soc. Proc. 54: 29-36. (319) STANELY, W. W. 1933. OUTBREAK OF GRASSHOPPERS IN TENNESSEE DURING 1932. Jour. Econ. Ent. 26: 300-301. (320) STONE, A., SABROSKY, C. W., WIRTH, W. W., and others. 1965. A CATALOG OF THE DÍPTERA OF AMERICA NORTH OF MEXICO. U.S. Dept. Agr., Agr. Handb. 276, 1696 pp. (321) STORER, T. I. 1943. GENERAL ZOOLOGY. 798 pp. New York and London. 282 TECHNICAL BULLETIN NO. 14Ó0, U. S. DEPT. OF AGRICULTURE

(322) STRANDTMANN, R. W. 1945. OBSERVATIONS ON THE NESTING HABITS OF SOME DIGGER WASPS (SPHECIDAE; HYMENOPTERA) . Ent. Soc. Amer. Ann. 38: 305-313. (323) SWEETMAN, H. L. 1958. THE PRINCIPLES OF BIOLOGICAL CONTROL. 560 pp. Dubuque, lowa. (324) TAYLOR, G. W. 1886. THE ENTOMOLOGY OF VANCOUVER ISLAND. NOTES ON SEVENTY-SIX SPECIES OF CICINDELIDAE AND CARABIDAE COLLECTED NEAR VICTORIA, VANCOUVER ISLAND. Canad. Ent. 18: 34-37. (325) THOMPSON, W. R. 1943. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 1. PARASITE HOST CATALOGUE. PT. 2. PARASITES OF THE DER- MAPTERA AND DÍPTERA. 99 pp. Imp. Agr. Bur., Inst. Ent., Belle- ville, Ontario. (326) 1951. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 2. HOST PARASITE CATALOGUE. PT. 1. HOSTS OF THE COLEÓP- TERA AND DíPTERA. 147 pp. Commonw^ealth Agr. Bur., Common- wealth Inst. Biol. Control, Belleville, Ontario. (327) 1953. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 2. HOST PARASITE CATALOGUE. PT. 2 HOSTS OF THE HYMENOP- TERA (AGAONIDAE TO BRACONIDAE). 190 pp. Commonwealth Agr. Bur., Commonwealth Inst. Biol. Control, Belleville, Ontario. (328) 1955. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 2. HOST PARASITE CATALOGUE. PT. 3 HOSTS OF THE HYMENOP- TERA (CALLICERATID TO EVANIID). 332 pp. Commonwealth Agr. Bur., Commonwealth Inst. Biol. Control, Belleville, Ontario. (329) 1958. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 2. HOST PARASITE CATALOGUE. PT. 5 HOSTS OF THE HYMENOP- TERA (MISCOGASTERIDAE TO TRIGONALIDAE) LEPIDOPTERA AND STREP- SIPTERA. 698 pp. Commonwealth Agr. Bur., Commonwealth Inst. Biol. Control, Belleville, Ontario. (330) and SIMMONDS, F. J. 1964. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 3. HOST PREDATOR CATALOGUE. 204 pp. Commonwealth Agr. Bur., Commonwealth Inst. Biol. Control, Belleville, Ontario. (331) and SIMMONDS, F. J. 1965. A CATALOGUE OF THE PARASITES AND PREDATORS OF INSECT PESTS. SECT. 4. HOST PREDATOR CATALOGUE. 198 pp. Commonwealth Agr. Bur., Commonwealth Inst. Biol. Control, Belleville, Ontario. (332) THORNE, G. 1940. THE HAIRWORM, GORDIUS ROBUSTUS LEIDY, AS A PARASITE OF THE MORMON CRICKET, ANABRUS SIMPLEX HALDEMAN. Wash. Acad. Sci. Jour. 30: 219-231. (333) TORRE-BUENO, J. R. DE LA. 1962. A GLOSSARY OF ENTOMOLOGY. 336 pp. Brooklyn Ent. Soc, New York. ARTHROPOD AND NEMATODE PARASITES IN AMERICA 283

(334) TOWNSEND, C. H. T. 1892. A DEXiiD PARASITE OF A LONGICORN BEETLE. Jamaica Inst. Jour. 1: 105-106. (335) _ 1892. A SARCOPHAGID PARASITE OF ciMBEX AMERICANA. Canad. Ent. 24: 126-127. (336) TREHERNE, R. C, and BUCKELL, E. R. 1924. THE GRASSHOPPERS OF BRITISH COLUMBIA WITH PARTICULAR REFER- ENCE TO THE INFLUENCE OF INJURIOUS SPECIES ON THE RANGELANDS OF THE PROVINCE. Canada Dept. Agr. Bui. 39 (n.s.), 47 pp. (337) TUCKER, E. S. 1907. SOME RESULTS OF DESULTORY COLLECTING OF INSECTS OF KANSAS AND COLORADO. Kans. Univ. Sei. Bul. 4, pp. 51-112. (338) U.S. DEPARTMENT OF AGRICULTURE, AGRICULTURAL RESEARCH SERVICE. 1960. PARASITES CONTROL GRASSHOPPERS. U.S. Dept. Agr., Agr. Res. 8, No. 11, p. 10. (339) 1961. A GLOSSARY OF SOME FOREIGN-LANGUAGE TERMS IN ENTOMOLOGY. U.S. Dept. Agr., Agr. Handb. 218, 59 pp. (340) VAN DEN BOSCH, R., and TELFORD, A. D. 1964. ENVIRONMENTAL MODIFICATION AND BIOLOGICAL CONTROL. In De- Bach, P., Biological Control of Insect Pests and Weeds, pp. 459- 488. New York. (341) VAN DYKE, E. C. 1928. A RECLASSIFICATION OF THE GENERA OF NORTH AMERICAN MELOIDAE

(COLEóPTERA) AND A REVISION OF THE GENERA AND SPECIES FOR-

MALLY PLACED IN THE TRIBE MELOINI, FOUND IN AMERICA NORTH OF MEXICO, TOGETHER WITH DESCRIPTIONS OF NEW SPECIES. CALIF. Univ. Pub. Ent. 4, pp. 395-474. (342) VAURIE, P. 1950. THE BLISTER BEETLES OF NORTH CENTRAL MEXICO (COLEÓPTERA, MELOIDAE). Amer. Mus. Nat. Hist. Amer. Mus. Novitates 1477, 68 pp. (343) VIERECK, H. L. 1904. A HANDSOME SPECIES OF TACHYSPHEX FROM ARIZONA. Ent. News 15: 87-88. (344) WADE, J. S. 1935. A CONTRIBUTION TO A BIBLIOGRAPHY OF THE DESCRIBED IMMATURE STAGES OF NORTH AMERICAN COLEÓPTERA. 114 pp. U.S. Dept. Agr. Bur. Ent. and Plant Quar., Washington, D.C. (345) WAKELAND, C. 1958. THE HIGH PLAINS GRASSHOPPER. A COMPILATION OF FACTS ABOUT ITS OCCURRENCE AND CONTROL. U.S. Dept. Agr. Tech. Bul. 1167, 168 pp. (346) WALKER, F. 1836. MONOGRAPHIA CHALCIDITUM. Ent. Mag. 3: 465-496. (347) 1849. LIST OF THE SPECIMENS OF DIPTEROUS INSECTS IN THE COLLECTION OF THE BRITISH MUSEUM. V. 2, pp. 231-484. London. 284 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

(348) 1849. LIST OF THE SPECIMENS OF DIPTEROUS INSECTS IN THE COLLECTION OF THE BRITISH MUSEUM. V. 4, pp. 689-1172. London. (349) 1866. DESCRIPTIONS OF COLEÓPTERA. In Lord, J. K., The Naturalist in Vancouver Island and British Columbia, v. 11, 375 pp. London. (350) WALLIS, J. B. 1961. THE CICINDELIDAE OF CANADA. 74 pp. Toronto. (351) WALSH, B. D. 1866. [No title.] Pract. Ent. 1, p. 126. (352) WEBER, N. A. 1935. THE BIOLOGY OF THE THATCHING ANT, FORMICA RUFA OBSCURIPES FOREL, IN NORTH DAKOTA. Ecol. Monog. 5: 165-206. (353) 1939. DESCRIPTIONS OF NEW NORTH AMERICAN SPECIES AND SUBSPECIES OF MYRMiCA LATREiLLE (HYM: FORMICIDAE) . Lloydia 2: 144-152. (354) 1941. NOTES ON THE EFFECT OF DROUGHT UPON THE NESTING HABITS OF ANTS. Canad. Ent. 73: 140-141. (355) WELCH, H. E. 1965. ENTOMOPHiLic NEMATODES. Ann. Rev. Ent. 10: 275-302. (356) WERNER, F. G. 1943. SOME NEW NORTH AMERICAN SPECIES OF EPICAUTA (COLEOP. MEL- OIDAE). Psyche 50: 65-73. (357) 1945. A REVISION OF THE GENUS EPICAUTA IN AMERICA NORTH OF MEXICO (COLEÓPTERA: MELOIDAE). Harvard Mus. Compar. Zool. Bui. 95, pp. 421-517. (358) ENNS, W. R., and PARKER, F. H. 1966. THE MELOIDAE OF ARIZONA. Ariz. Agr. Expt. Sta. Tech. Bui. 175, 96 pp. (359) WERNER, W. V. 1903. [A NOTE ON SCELIO.] Wash. Ent. Soc. Proc. 5: 308-309. (360) WHEELER, W. 1905. NEW SPECIES OF FORMICA. Amer. Mus. Nat. Hist. Bui. 21, pp. 267- 274. (361) 1908. THE ANTS OF TEXAS, NEW MEXICO AND ARIZONA. I. Amer. Mus. Nat. Hist. BuL 24, pp. 399-485. (362) 1913. A REVISION OF THE ANTS OF THE GENUS FORMICA (LINNÉ) MAYR. Harvard Mus. Compar. ZooL BuL 53, pp. 379-565. (363) WiCKHAM, H. F. 1894. THE COLEÓPTERA OF CANADA. 1. THE CICINDELIDAE OF ONTARIO AND QUEBEC. Canad. Ent. 26: 149-154. (364) 1896. PRELIMINARY HANDBOOK OF THE COLEÓPTERA OF NORTHEASTERN AMERICA. N.Y. Ent. Soc. Jour. 4: 33-49. (365) WiEDEMANN, C. R. W. 1821. DíPTERA EXOTICA. Ed. 2, 244 pp. Kiliae (Kiel). ARTHROPOD AND NEMATODE PARASITES IN AMERICA 285

(366) 1828. AUSSEREUROPAISCHE ZWEIFLÜGELIGE INSEKTEN. V. I, 608 pp. Hamm. (367) WiLCOX, J. 1966. EFFERIA COQ. IN AMERICA NORTH OF MEXICO (DÍPTERA: ASILIDAE). Calif. Acad. Sei. Proc. 3, pp. 85-234. (368) WILLIAMS, F. X. 1913. MONOGRAPH OF THE LARRIDAE OF KANSAS. Kans. Univ. Sci. Bul. 8: 121-213. (369) 1913. NOTES ON THE HABITS OF SOME SOLITARY WASPS THAT OCCUR IN KANSAS, WITH THE DESCRIPTION OF A NEW SPECIES. KanS. Univ. Sei. Bul. 8: 223-230. (370) 1932. TACHYSPHEX Fusus FOX. Hawaiian Ent. See. Free. 8, p. 17. . (371) 1950. THE WASP OF THE GENUS SOLIERELLA IN CALIFORNIA (HYMENOPTERA, SPHECIDAE, LARRINAE). Calif. Aead. Sei. Proe. 26, pp. 355-417. (372) and HUNGERFORD, H. B. 1914. NOTES ON COLEÓPTERA FROM WESTERN KANSAS. Ent. NeWS 25: 1-9. (373) WiLLISTON, S. W. 1880. SOME INTERESTING NEW DÍPTERA. Conn. Aead. Arts and Sei. Trans. 4, pp. 243-246. (374) 1883. THE NORTH AMERICAN SPECIES OF NEMESTRINIDAE. Canad. Ent. 15: 69-72. (375) 1885. ON THE NORTH AMERICAN ASILIDAE (PT. II ). Amer. Ent. Soe. Trans. 12: 53-76. (376) 1889. THE DIPTEROUS PARASITES OF NORTH AMERICAN BUTTERFLIES. In Seudder, S. H., Butterflies of the Eastern United States and Canada, (q.v.) 3: 1912-1924. Cambridge, Mass. (377) 1893. LIST OF DÍPTERA OF THE DEATH VALLEY EXPEDITION. In Riley, C. V., The Death Valley Expedition. A Biological Survey of Parts of California, Nevada, Arizona, and Utah. Pt. II. 4. Report on a Small Collection of Insects Made During the Death Valley Ex- pedition. North Amer. Fauna 7, pp. 253-259. (378) 1894. [A note, no title.] Ent. News 5: 47-48. (379) WILSON, C. C. 1936. NOTES ON THE WARRIOR GRASSHOPPER CAMNULA PELLUCIDA (SCUD- DER) AND ITS EGG PARASITE APHOEBANTUS HIRSUTUS COQUILLETT, IN NORTHERN CALIFORNIA, 1928-29. Jour. Econ. Ent. 29: 413-416. (380) and KEIFER, H. H. 1941. A STUDY OF THE MARGÍNATE GRASSHOPPER WITH NOTES ON ASSOCI- ATED CONGENERIC SPECIES. Calif. Dept. Agr. Bul. 30: 291-306. (381) WULP, F. M. VANDER. 1895. FAMILY MUSCIDAE. In Zoologia-Insecta-Diptera, v. 2, 489 pp. London. 286 TECHNICAL BULLETIN NO. 14Ó0, U. S. DEPT. OF AGRICULTURE

(382) YORK, G. T. 1955. NOTES ON PARASITIZATION OF GRASSHOPPERS BY NEMESTRINIDS. Jour. Econ. Ent. 48: 328. (383) and PRESCOTT, H. W. 1952. NEMESTRiNiD PARASITES OF GRASSHOPPERS. Jour. Econ. Ent. 45 I 5- 10.

UNPUBLISHED MATERIAL'

(384) BANFILL, J. C. 1970. Personal correspondence (Dec. 8, 1970). (385) CAMPBELL, J. E. 1964. A BIOLOGICAL STUDY OF THE GRASSHOPPER MITE, EUTROMBIDIUM LO- CUSTARUM (WALSH) (ACARIÑA: TROMBIDIIDAE). [Unpublished master's thesis. Copy on file Dept. Ent., Univ. Mo., Columbia.] (386) HUGGANS, J. L. 1960. A STUDY ON THE ECOLOGY AND BIOLOGY OF THE GRASSHOPPER MITE, EUTROMBIDIUM LOCUSTARUM WALSH IN MISSOURI (TROMBIDIIDAE). [Unpublished master's thesis. Copy on file Dept. Ent., Univ. Mo., Columbia.] (387) POINAR, G. 1967. Personal correspondence (Nov. 4, 1969). (388) PRESCOTT, H. W. 1969. Personal correspondence (Feb.-Aug. 1969). (389) U.S. DEPARTMENT OF AGRICULTURE, AGRICULTURAL RESEARCH SERVICE. 1917-70. Bozeman, Mont., Grasshopper Investigations Laboratory un- published records. (390) 1970. Entomology Research Division, Insect Identification and Parasite Introduction Research Branch, personal correspondence (May 1, 1970). ^ Unless otherwise indicated, copy on file Grasshopper Invest. Lab., Agr. Res. Serv., U.S. Dept. Agr., Bozeman, Mont.

INDEX

Page Page abdominalis (Say), Tachytes 120 anonyma (Riley), Lespesia 74 aculeata (Aldrich), Blaesoxipha __.. 29 arida (Williston), Efferia 220 aenea (Say), Pomphopoea 212 atlanis (Aldrich), Blaesoxipha 33 aerarius Patton, Sphex 97 atratus LePeletier, Sphex 98 aestuans (Linnaeus), Efferia 217 auripes (Provancher), Aphaereta 129 albibarbis (Macquart), Efferia 219 albidae (Say), Epicauta 178 barbatus Osten Sacken, aldrichii Coquillett, Psilocephala.— 251 Anastoechus 242 americana Weber, Myrmica bicaudata (Hine), Efferia 221 sabuleti 257 bilineatus (Say), Cantharis 175 angustifrons (Aldrich), bisulcus (Ashmead), Scelio 79 Blaesoxipha 31 angustifrons (Meigen), Hylemya .. 213 californiens Chaudoir, angustipennis Loew, Diogmites 216 Pasimachus 158 ARTHROPOD AND NEMATODE PARASITES IN AMERICA 287

Pasre Page californicus DeJean, haemorrhoidalis (Macquart), Anisodactylus 148 Psilocephala 251 caliginosus (Fabricius), harpax Pandelle, Sarcophaga 60 Harpalus 155 harveyi (Hine), Efferia 224 callosa LeConte, Epicauta 179 helenae (Bromley), Efferia 225 calopteni Riley, Scelio 81 helvolus (Loew), Stenopogon 237 cana (Hiñe), Efferia 222 hesperia Casey, Amara 146 canadensis Snyder, Acridomyia 10 heteroneura (Meigen), cessator (Aldrich), Blaesoxipha __„ 35 Taxigramma 142 cimbicis (Townsend), hirsutus Coquillett, Aphoebantus 243 Boettcheria 54 hunteri (Hough), Blaesoxipha 40 cinérea (Forster), Epicauta 181 hyalinipennis Ashmead, Scelio 81 circumpicta LaFerté-Sénectère, hyalinus Say, Perilampus 135 Cicindela 165 claripennis (Macquart), ichneumoneus ichneumoneus Euphorocera 72 (Linnaeus), Sphex 102 clausa (Osten Sacken), Immaculata (Say), Epicauta 190 Trichopsidae 20 impar (Aldrich), Blaesoxipha 43 coloradensis ( Aldrich ), impuncticollis (Say), Amara 146 Blaesoxipha 36 infidelis Fall, Lytta 210 coloradensis (Cresson), inquinatus Loew, Stenopogon 238 Brachymeria 131 interrupta (Macquart), Efferia.— 226 comma (Fabricius), Agonoderus 145 conferta (Say), Epicauta 183 kellyi (Aldrich), Blaesoxipha 45 coyote Bromley, Stenopogon 236 cyanipennis LeConte, Lytta 209 laeviventris Cresson, Sphex 103 cylindriformis (Say), lateralis Adams, Psilocephala 252 Amblycheila 163 latipennis Horn, Callisthenes 149 latipennis (Loew), Ospriocerus 231 decaudata Cobb, Steiner, and lemniscata Fabricius, Epicauta 192 Christie, Agamermis 86 locustarum (Walsh), dentata (Coquillett), Ceracia 70 Eutrombidium 127 desertus LeConte, Harpalus 156 longus (Wiedemann), Proctacanthus 232 elongatus LeConte, Pasimachus _. 159 luggeri Riley, Scelio 82 elongatus elongatus Cresson, Tachytes 120 maculata (Say), Epicauta 194 exornatus Fox, Tachytes 122 melanopli Christie, externum (Say), Calosoma 150 Agamospirura —: 87 milbertii Macquart, fabricii (LeConte), Epicauta 185 Proctacanthus 233 falciformis (Aldrich), molesta validiuscula Emery, Blaesoxipha 38 Solenopsis 258 femorata (Macquart), Efferia 224 murina (LeConte), Epicauta 196 ferruginea (Say), Epicauta . 187 mus mus (Osten Sacken), fonscolombei (Dufour), Aphoebantus 243 Brachymeria 132 formosa Say, Cicindela 165 neglectus Bromley, Stenopogon 239 fossor (Rohwer), Solierella 95 nivalus Horn, Anisodactylus 149 fraternus LeConte, Harpalus 156 normalis Werner, Epicauta 197 frenchii (Williston), Lespesia 75 nuttalli Say, Lytta 210 fulgida Say, Cicindela 166 fulviventris fulviventris Cresson, obductus Fox, Tachytes 123 Tachytes 122 obesa (Say), Amara 147 fumisquama Snyder, Acridomyia ._. 12 oblita LeConte, Epicauta 198 funebris Horn, Epicauta 188 obscuripes Forel, Formica rufa 255 fusus Fox, Tachysphex 108 obscuriventris Loew, Stenopogon—. 240 obtusopilosa Emery, Formica 254 gavia (Aldrich), Blaesoxipha 40 occidentalis (DeJean), Pterostichus 160 haemorrhoidalis ( Fallen ), occidentalis (Hine), Machimus 231 Sarcophaga 58 oedipodae Ashmead, Scelio 83 288 TECHNICAL BULLETIN NO. 1460, U. S. DEPT. OF AGRICULTURE

Page Page oestriforme Brauer and sarcophagae Gahan, Aphaereta 130 Bergenstamm, Hemithrixion 73 sarraceniae Riley, Sarcophaga 61 oophaga Sabrosky, Goniopsita 249 sarracenioides Aldrich, opifera (Coquillett), Blaesoxipha.,. 47 Sarcophaga 62 oreas Osten Sacken, Systoechus 245 sayi Banks, Tachytes 125 oregona Horn, Epicauta 199 scrutator (Fabricius), Calosoma—. 151 ovivorus (Riley), Scelio 84 scutellaris Say, Cicindela 172 segmenta (Say), Epicauta 203 pallidula (Hine), Efferia 227 semirufus (Cresson), pedicularius DeJean, Tachysphex 112 Selenophorus 161 sericans LeConte, Epicauta 205 pennsylvanica (DeGeer), sexguttata Fabricius, Cicindela 173 Epicauta 200 shermani Parker, Sarcophaga 64 pennsylvanicus (DeGeer), similis (Bridwell), Solierella Harpalus 157 plenoculoides 96 pepticus (Say), Tachytes 124 similis Rohwer, Tachysphex 113 picticornis (Loew), Stenopogon 240 simplex (Fallen), Leucostoma 75 platura (Meigen), Hylemya ^ 214 sinuata Meigen, Sarcophaga 65 plenoculiformis Williams, snowi (Hine), Efferia 229 Tachysphex 109 solitus (Walker), Systoechus 246 plenoculoides similis (Bridwell), somnulentis DeJean, Harpalus 158 Solierella 96 sphaerioollis (Say), Linsleya 208 plumipes (Fabricius), splendida Hentz, Cicindela 174 Trichopoda 76 staminea (Williston), Efferia 230 propinquus Viereck, Tachysphex 110 sternodontis Townsend, pubidorsus (Costa), Sphex 104 Sarcodexia 57 pulchra Say, Cicindela 167 striatipes (Ashmead), Solierella...- 97 punctatus Haldeman, striatus (Smith), Sphex 106 Pasimachus 160 subglabra (Fall), Epicauta 206 puncticollis ( Mannerheim ), subnigrescens Cobb, Mermis 87 Epicauta 202 punctulata Oliver, Cicindela 168 taediosa (Aldrich), Blaesoxipha— 52 purpurea Oliver, Cicindela 169 tarsatus (Say), Tachysphex 114 tegularis (Cresson), quadrinotatus Say, Oxybelus 140 Brachymeria 134 quebecensis ( Provancher), tenuipunctus Fox, Tachysphex 115 Tachysphex 111 tepidum LeConte, Calosoma 152 terminatus (Smith), Tachysphex.... 117 rapax (Osten Sacken), Efferia 228 texanus (Cresson), Tachysphex _._. 119 rapax (Walker), Helicobia 56 thomae Fabricius, Sphex 107 repanda DeJean, Cicindela 170 tibialis Coquillett, Acemya 68 reversa (Aldrich), Blaesoxipha 50 tranquebarica Herbst, Cicindela— 174 rodecki James, Proctacanthus 234 rubriventris Macquart, uncata (Wulp), Blaesoxipha 53 Senotainia 141 unicinctus (Say), Stizoides 140 rufa obscuripes Forel, Formica 255 rufiventris rufiventris Cresson, Sphex 105 validiuscula Emery, Solenopsis rufofasciatus Cresson, Tachytes 125 molesta 258 ruscarius Say, Elaphrus 154 vertebratus (Say), Promachus 234 vicina Robineau-Desvoidy, sabuleti americana Weber, Calliphora 13 Myrmica 257 vitripennis (Walker), Nasonia 138 sackenii (Williston), vittata (Fabricius), Epicauta 207 Neorhynchocephalus 16 vulgaris Loew, Systoechus 247 sarcophaga Gahan, Eupteromalus 138 wilcoxi LeConte, Calosoma 153

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