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1924] The Biology of Trichopoda pennipes Fab. 57

THE BIOLOGY OF TRICHOPODA PENNIPES FAB.(DIPTERA, TACHINID2E), A PARASITE OF THE

COMMON SQUASH BUG.*

BY HARLAN N. WORTHLEY.

Massachusetts Agricultural Experiment Station, Amherst,Mass.

PART II.

MORPHOLOGY

In the study of the adult anatomy, pinned dried specimenswere used. For the definition of the mouth parts, sclerites of thethorax, and the genitalia, however, it was found necessary torelax the parts and examine them in liquid. For this purpose,specimens were soaked for about an hour in a cold 10 per centsolution of caustic potash (boiling often causes distortion of theparts) washed in water and treated with weak acetic acid tostop the action of the caustic potash. They were then placed in70 per cent alcohol.

The parts were examined under a Zeiss binocular micro-scope, at magnifications varying from sixteen to sixty-fivediameters. Many structures were obscure except under thebrightest illumination, and therefore most of the examinationswere made in the rays from a powerful lamp. A Ford headlightwas mounted on a ringstand and connected through a trans-former with the ordinary one hundred and ten volt circuit.This lamp proved to be quite satisfactory, since it was placed onthe desk at a distance of two feet from the binocular, allowingplenty of room to work. A lamp of this kind, focussed upon the.microscope stage by means of the set-screw in the lamp, throws’.little light into the eyes and develops little heat, while the objectunder observation is brought into strong relief.

*The first portion of this article appeared in Psyche, vol. 31, pp. 7-16,February, 1924.

58 Psyche [April

Adult.

The adult fly is about the size of the common house fly’but it is much more gay in appearance. It may be seen on sunnydays hovering about squash plants, or resting with half-spreadwings upon the foliage of squash and upon certain wild flowersas well. It is strikingly colored, with deep reddish brown eyeson a head marked with black, gold and silver. The thorax isgolden in front, with four longitudinal black stripes, clear blackbehind, and gray at the sides. The abdomen is of a brilliantorange color, except at the extreme tip, which is darker. Theconspicuous abdomen, and the fringe of feather-like sete alongthe outer side of the hind tibiee, immediately catch the eye of theobserver, and serve to make this species one of the most strikingamong Tachinid flies.

A discussion of the adult anatomy is complicated by thediversity of terms which may be applied to the different struc-tures. Taxonomists have applied names which, in many cases,are morphologically inaccurate, and morphologists ,themselveshave differed both in the nomenclature and in the interpretationof parts. The source of the terms used in this paper is indicatedin the text of the different sections, and in many cases duplicatenames for the various structures are given in the list of ab-breviations used in the figures.

Head. P1. 1, figs. 1 and 2. In describing the head, theterms used are those of Peterson (1916), except the chetotaxy,which follows Coquillet (1897) and Walton (1909).

Viewed from in front, the head is elliptical in outline, andbroader than deep (3.2 mm. by 2.4 mm.). Its most conspicuousfeature is perhaps the frontal suture (fs), which extends in adark, shining, inverted U-shaped band from just above theinsertion of the antennm to a point midway between the vibrissee(vib) and the curve of the compound eyes (ce), where it tapersout. Within the curve of the frontal suture lies the fronto-clypeus (fc), termed by Coquillett the "facial depression" and

Letters in parenthesis are those used ih labeling the figures, and areexplained in the list of abbreviations preceding the plates at the end of thispaper.


by Walton the "facial plate." The tentorial thickenings (tt)arising at each side near the oral margin and running upwardnearly to the insertion of the untenne, are easily seen, lyingjust within the facial or vibrissal ridges, which are not pro-nounced. The vertex (v) is 11 that portion of the head, viewedis the ocellar triangle, bearing on its raised surface three ocelli(oc). From the region of the ocelli to the frontal suture runs umedian broad velvety-black bund or "vittu" (mv), which isdemarked from the rest of the vertex only by its color, whichstrongly contrasts with the golden-yellow tomentum of thelateral portions of the vertex. The genee (ge) are those portionsof the vertex lying below the ends of the frontal suture, andbetween the oral margin and the eyes. Their color is silvery-gray, which shades into the gold of the rest of the vertex above,and into the brownish-yellow of the fronto-clypeus.

Viewed from the side the hed is quadrate in shape. Thepostgene (pge) are those regions behind the genre and extendingbackward and upward along the curve of the compound eyesto u point midway between the oral margin and the ocellartriangle. The occiput is designated as that portion of the caudalaspect of the head extending from line drawn midway acrossthe occipital foramen upward to the vertex. The edge of thisarea can be seen from the side (ocp).

Chetotaxy of the Head. On either side of the median vittis a row of frontal bristles (fb) which, since they bend inwardacross the vitta, may be called "transfrontals."

On the ocellar triangle, just behind the anterior ocellus, liethe great ocellar pair (ob), while behind these, and passingbetween the two lateral ocelli, follow three or more pairs of"lesser ocellar" bristles, which in T. pennipes are very small.

Here is one instance of the confusion of terms mentioned at the beginningof this paper. The area bearing the frontal bristles, although it has beencalled the "front" by taxonomists, is morphologically not the front at all,but the vertex. The true front, which lies below the antennae and is fusedwith the clypeus, also bears a double row of macrochaetae which, to one not aspecialist, might readily be mistaken for the frontal bristles. The writer doesnot recommend here any reconciliation between the terms of the morpho-logists and the usage of the taxonomists, but merely wshes to point out thetrue relation of parts. To his mlnd, any attempt to modify the terminologyother than by concerted action among taxonomic workers and morphologistswould only result in "confusion worse confounded."

60 Psyche [April

Behind the ocelli nd on the edge of the occiput is s transverserow of four mcrochsetse. The inner, lrger pir re the post-vertical bristles (pvt), nd the smller, outer ones, the innervertical bristles (ivt). The outer verticals, present in someforms, re not represented in this species. The fronto-orbitls,which lie between the frontal bristles nd the curve of the com-pound eyes, re lso bsent.

On the fronto-clypeus, disposed long ech fcil ridge,is short row of fcil bristles, the vibrissl row. The uppermost.pir re the vibrissse (vib) which in T. pennipes do not ssumefrom the front, which lies between the compound eyes, nd be-tween these nd the frontsl suture. At the very top of the vertextheir typical position immediately sbove the oral mrgin, butre shifted upward to lie hlfwy between the oral mrgin ndthe tips of the ntennse. The smller bristles ccompnyingthe vibrissm extend on either side in single line long the oralmargin to the region of the postgense, where they mingle withthe silvery-white beard which depends from this region. Asingle row of short mcrochmtse extends around the edge of theocciput from the inner vertical bristles downwsrd to the regionof the postgense. These are clled the cili of the posteriororbit (cpo).

Appendages of the Head.

Antenna. The ntennse (nt) rech halfway between thebse of the frontal suture nd the oral mrgin. The first twosegments sre velvety-black in color, with silvery sheen. Thesecond segment bers few mcrochsetse. The third segment,which is much lrger titan the other two, is ben-shped ndvries from blck to mouse-colored: with the bse sometimesslightly tawny. This segment bers the rist (sr), lrgebristle which is inserted on the outer edge bout one third thedistance from bse to tip of the segment. The rist is prac-tically bsre, hving but few very tiny hirs nesr its base.

Proboscis. The proboscis (pb), usually folded well bck inthe oral cvity, is very much modified structure, the psrts ofwhich it is very difficult to homologize with the mouth prts of


generalized insects. The work of Peterson (1916) on the mouthparts of Diptera was very thorough, and his figures correct andintelligible, but since he derives his hypothetical dipterousmouth parts from a consideration of Orthoptera while Crampton(1921, p. 91)would evolve Diptera from ancestors like Mecop-tera, the homologies of dipterous mouth parts still constitute adisputed question.

The membrane of the basiproboscis (bpb) is largely composedof the mentum and sub-mentum, according to Peterson. Themaxillary palpi; (mxp) lie on this membrane in front. Above themaxillary palpi lie the exposed portions of the tormm (to), andbelow lie the external plates of the stipes (st). The galem (ga)lie on the surface, and are continuous with the lower ends ofthe ental portions of the stipes. The large chitinous internalstructure of the basiproboscis is the fulcrum (ful) and is com-posed of the basipharnyx, or united po.rtions of the epipharynxand hypopharynx, and the ental portions of the tormm. At thedistal end of the basipharynx lies the hyoid (hy), which articulateswith the distal portion of the hypopharynx as well, and throughwhich passes the alimentary canal.

The mediproboscis (mpb)bears the chitinized plate, thetheca (the), on its caudal aspect, and the hypopharynx (hyp)and labrum-epipharynx (lep) lie in a chitinized groove on theupper surface of the labium.

The distiproboscis (dpb) is composed of a pair of lob,:-s orlabella, which Peterson interprets as the paraglossm (pg).Crampton, however, calls them the united labial palpi. Variousother structures can be seen in the distiproboscis, such as aY-shaped plate called the fl_lrcu (fu) and the structures calledpseudotrachm (pst).

Thorax. P1. I, figs. 3 and 4. The structure of the thoraxin Trichopoda pennipes is typical of the order Diptera as a whole,in which the mesothorax, which is the only wing-bearing seg-ment, is greatly enlarged and distorted, evidently for the purposeof accomodating the great wing muscles. The prothorax (P)is very small, and the metathorax, which bears the halteres(ha),is very much reduced.

62 Psyche [April

In naming the sclerites of the thorax the terminology usedby Young (1921), which is largely based on Crampton (1914),is employed.

The dorsal aspect of the thorax is completely covered bythe notum of the msothorax, as defined by Snodgrass (1909a),or the mesonotum. This is divided by two transverse suturesinto three sclerites, the prescutum (psc), scutum (sc) andscutellum (sly). The prescutum, including the humeral calli(hc) is yellow in color, with four longitudinal bands of velvety-black. In the males the yellow coloration extends backwardonto the scutum where it merges with the black of that sclerite.The scutellum of both sexes appears black to the naked eye, butunder the binocular most specimens show a faint tinge of verydark orange. The scutum is produced laterally into an anteriorwing process, the suralare (sur), and a posterior wing process,the adanale (ad). The scutellar bridge df Walton (sb) is seen asa lateral overlapping of the scutellum onto the scutum. Belowthis is the axillary cord (axc) of Snodgrass (1909a), which isproduced to form the margin of the calypteres. A posttergite(pt) is demarked behind the scutellum. The pseudonotum orpostnotum of Snodgrass, which he would recommend callingthe "postscutellum", in this case is located ventrad of the scu-tellum, and cannot be seen from above. It is divided into amedian plate, the meditergite (mt), and two pairs of lateralplates, the anapleurotergites (aplt) and the katapleurotergites(* Mention may logically be made here of the characterrecently reported by Malloch (1923) for differentiating muscoidflies. In Malloch’s own words, "It is invariably possible todistinguish between the Sarcophagidm, Muscidm and Calli-phoridm on one hand, and the Tachinidee and Dexiidm on theother, by the shape of the metanotum. In the last two this isbiconvex in profile, there being a small but distinct convexityjust below the scutellum which is absent in the other threefamilies known to me." The use of the term "metanotum" byMalloch follows the usage of older taxonomic workers, and ismorphologically inaccurate. It is really the meditergite (mt)of the postscutellum which is meant, and the "biconvexity"


apparent in Tachinidm and Dexiidse is conditioned by thepresence of the posttergite (pt), which, as a glance at the figurewill show, lies iust below the scutellum. An examination offigures 38, 39, 40 and 41 of Young bears out this point.

The pleural region of the mesothorax is pollinose gray incolor, and is much distorted. The pleural suture, which ingeneralized insects runs a nearly straight course from the coxalcavity to the wing base, thus dividing the pleuron into an an-terior episternum and a posterior epimeron, is here bent twiceat right angles, so that while the two ends are nearly vertical,the middle is horizontal. In addition a portion of the anepister-hum (aes) has been split off from the rest by a secondary in-vasion of membrane, and has become closely associated withthe anepimeron or pteropleurite (ptp.) The katepisternum hasfused with the sternum to form the sternopleurite (stp). It isthe enlargement of this sclerite which has evidently caused thebending of the pleural suture, and has crowded the meropleurite(inept), which is .composed of katepimeron plus meron, backagainst he pleuron of the metathorax.

The numerous small plates which lie in the membrane sur-rounding the base of the wing are very difficult to see, but areeasily identified with those sclerites outlined by Crampton (1914)in his ground plan of a typical thoracic segment in winged insects.The tegula (tg) lies in the angle between the scutum and theanepisternum. The notale (n) is a detached portion of thescutum lying just above .the base of the wing. The basalarplates are two in number, the anterior one (aba) not demarkedfrom the posterior portion of the anepisernum, the posteriorone (pba) very small and lying between it and the pleural wingprocess (wp). The subalar plates are two in number, the an-terior one (asa) lying behind the wing process and above thepteropleurite, the posterior one (psa) which is much smallerlying just below a posterior lateral process of the scutum. Thesebasalar and subalar plates are the pre and post paraptera ofSnodgrass (1909a).

The tergum of the metathorax, or the metanotum (n) isreduced to a narrow band connecting the halteres (ha), andvisible only at the sides where it is produced to form points of

64 Psyche [April

attachment for the abdomen. The pleuron of this segment isdivided into metaepisternum (es3) and metaepimeron (em).A spiracle (sp) is present just before the metaepisternum, asbefore the mesoepisternum. The region around the base of thehaltere is so modified that it is impossible to tell whether preand post alar bridges connect the metanotum with the meta-pleuron.

Some of the terms used above are different from those incommon use among taxonomists. The mesoanepisternum (aes)has been called by dipterists the mesopleura. The mesosterno-pleurite (stp) is equivalent to the sternopleura of authors,while the meropleurite (mep) plus metapleuron plus meta-sternum equals the hypopleurite, so-called.Chaetotaxy of the Thorax. The thorax of T. pennipes is notheavily armed with macrochmte. However, representatives ofmost of the groups mentioned by Walton are present. Twohumerals (hu) adorn each humeral callus. Posthumerals arewanting, as are anterior acrosticals. The anterior dorsocentralrows are represented by two very variable bristles (adc) placednear the hinder margin of the prescutum, while at each rearcorner of this sclerite are borne two notopleural bristles (np).On each side, between the notopleurals and the anterior dorso-centrals, lies a single bristle, the presutural (psu).

On either side of the scutum a single bristle (sa) representsthe supra-alar row, and another (ia) each intra-alar row. Twopost alars (pa) are present, and each of the posterior dorso-central (pdc) and posterior acrostical (pac) rows is representedby a single bristle. It will be seen that these last four bristlesform a transverse row near the hind margin of the scutum. Thisis called the prescutellar row.

On the scutellum an anterior bristle and a posterior bristlemark the position of the marginal scutellar row (ms). Theanterior bristle was seen to be accompanied by a smaller one inone or two specimens. No discal scutellars are present.

The mesoanepisternum bears a vertical row of bristlescalled the mesopleural row (mr), situated just before the mem-brane which divides it. Below the anterior spiracle are two


bristles, oae on the prothorax, the propleural bristle (pp), andone on the steraopleurite, which the writer has called the sub-stigmal bristle (ss). The sternopleurite bears typically twosterno-pleurals (stb), although a third was found to be presenton some individuals. A curved row of three to five hypopleurls(hp) is located on the meropleurite. A single pteropleuralbristle (ptb) was present in some specimens examined, whileothers bore as many as four.

Appendages of the Thorax.Legs. P1. I, fig. 4; P1. II, figs. 7 and 8. The coxa (cx) is tawny

in color, with a grayish bloom, while the trochanter (tr) and theproximal portion of the femur (fe) are yellowish. The distal por-tion of the femur and the tibia (tb) and tarsus (ta) are black. Theclaws are yellowish tipped with black, and are fringed with veryfine light-colored hairs. There is a bristle-like empodium (ep)which, since it is a prolongation of a ventral plate, is u true em-podium ccording to Crampton (1923). The pulvilli (pv) arebuff-colored, and in the male are quite large and conspicuous.The first two pairs of legs display no features of particularinterest. The tibim of the hind legs, however, exhibit on theouter side peculiar row of black, feather-like setm, which standnearly erect, and the longest of which are at least one third thelength of the tibia itself. This row is in reality double, since arow of smaller scales is appressed to the larger ones on theoutside. The hind tibia also bears on its inner face singlebristle of a size noticeably larger than any of the surroundinghairs.

Wings. P1. II, figs 5 and 6. The wings of the female aredusky, with the posterior margin sub-hyaline. Those of all themales examined bear u somewhat variable yellowish area in theforepart of the wing, the extent of which is indicated in figure 5.According to Coquillett (1897) this character is not constant.

The figure of the wing of the female (fig. 6) explains thevenation of the wings, while the cells are labeled in the figure ofthe wing of the mule. The chief point of interest in the wingvenation of T. pennipes is that M is bullate or weakenedbasally, making M appear as a stub sticking up from Cu.

66 Psyche [April

Abdomen. P1. II, figs. 9-14. P1. III, fig. 15. The abdomenin both sexes is of a bright orange color and is destitute of macro-chete. It is sparsely clothed, however, with short black hairs.Seven pairs of spiracles (sp) are present, borne at the lateralmargins of the tergites (tl, t2), etc.). Those of the sixth andseventh segments are hidden beneath the posterior edge of thefifth tergite (ts). The tergites of the first and second segmentsare fused, the fusion being denoted by an area of weaker chitin,.which is demarked in the figures by a pair of dotted lines betweentl and t. The adventitious suture (us) in the first tergite, men-tioned by Young, is readily seen.

The tip of the abdomen in the female is wholly black, thiscoloration including the fifth tergite and in some individuals ex-tending further forward to include part of the fourth tergite.The terminal abdominal segments of the male in spccimensexamined by the writer were in no case wholly black, althought5 and t6 were darker than those preceding.

Genitalia.1 Pl. II, figs. 13-14; P1. III, fig. 15. In both sexes thesegments beyond the fifth abdominal may truly be called genitasegments. In the male these segments curve downward and cometo lie beneath the fifth tergite. In the female those beyond thefifth are telescoped when at rest, being extended for oviposition.

In the male the fused tenth and eleventh tergites,whichare ventral in position, act as a cover for the cedeagus (oe),being tucked beneath the edge of the fifth sternite (ss) when atrest. When the cedagus is extruded, however, this flap liftsup, allowing the ninth sternite (s.) to push forth. This lattersegment is very much modified. Its fused cerci are median inposition and form the cedagus, a very complicated structurewhich encloses the znembranous penis. At the base of thecedagus are seen two pairs of lateral projections, called gono-pophyses (go), the inner pair of which are hyaline. They arewell-chitinized, however, feeling hard to the touch of a dis-secting needle. At the base of the cedagus the ninth sternite is

1The writer has based his description of the genitalia largely on thecondition of these structures in generalized insects. It is apparent that thestudy of a series of dipterous genitalia may reverse some of his decisions re-garding the true character of the parts.


rather more heavily chitinized than elsewhere, resulting in theappearance of a chitinized box (chb) from which the mdeagusprotrudes and on which the gonopophyses are borne. Thischitinized box also bears a median dorsal hook-like projection,called by the writer the genital prong (gp). The styli of theninth segment, which in some insects function as outer claspers,are here much reduced in size and are apparently non-functional,since when the genitalia are extruded they barely appear beyondthe posterior edge of the eighth tergite. A peculiar structure,which the writer is at a loss to homologize with any genitalappendage of generalized insects, appears in the "genital furca"(gf). This is a fork-like chitinized rod which lies between thesides of the ninth sternite, to which it is connected by muscles.It splits at the base of the cedugus, one arm extending to eitherside of the latter organ. Its function is quite evidently that ofguiding the movements of the cedagus.

In the female the eighth segment is nrrow ring, bearingbelow the median ventral vlve (vv) of the ovipositor nd lterallythe two inner vlves (iv). Dorsally this segment seemed tober median dorsal valve (dv), but this may prove to bemodified portion of the ninth segment, which is supposed tobear the dorsal valve. This point could not be definitely de-termined from the dried mteril at the writer’s disposal, evenafter soaking in custic potash and gently extending the ovi-positor by pushing from within by means of blunt needle.

Secondary Sexual Characters. The foregoing account ofthe external anatomy of Trichopoda pennipes contains scatteredreferences to certain differences which were apparent betweenthe two sexes. These differences were constant in series ofeight males and seven females. Scarcely any difference in sizecould be noticed, the males veraging 8.6 mm. in length, thefemales 8 mm. Both the largest and the smallest were males,the one 10 mm. long, the other measuring 7 mm. To a certainextent the size of the adult fly is affected by the abundance offood availuble to the larva which preceded it, nd when con-tained in keys for the identification of species may be foundmisleading.

68 Psyche [April

Two characters were found by which the sex of living fliescan be determined without undue handling. These are theferrugineous spot in the wing of the male as against the evenlydusky wing of the female, and the black tip of the female ab-domen as against the dark orange of that of the male. A minordifference was in the size of the pulvilli, these being shorter thanthe last tarsal segment in the females, and inconspicuous, inthe males the pulvilli were longer than the last tarsal segment,nnd quite broad and conspicuous. This is a character, however,that is not readily noticed unless a male and a female are examinedat the same time, and it is therefore of little practical use, in ataxonomic sense.

EGG. P1. III, fig. 16.

The eggs of Trichopoda pennipes vary in color from clearshining white to dirty gray, the coloration seeming not to dependon the age of the egg. The individual egg is ovate in outline,being slightly larger at one end. It is strongly convex, and isflattened on the side next the body surface of the host. Thisflattened surface is covered by a colorless cement, by which theegg is affixed to the body of the host. The egg measures .56 mm.in length by .37 mm. in breadth, and its greatest height is .25mm. The surface of the chorion appears sm6oth except underhigh magnification, when it is seen to be faintly reticulate intiny hexugons. The chorion is comparatively thick and "leather-y", and remains rigid after hatching. The micropyle appearsto be borne on a small papilla at the smaller end of the egg.Eggs which have hatched show a circular hole on the flattenedside near the broader end. Since it is this flattened side which ispressed against the host, it is impossible to tell if an egg hashatched without first removing it from the body surface of thehost.

LArvA. P1. III, figs. 17 and 18.

The larva has not been examined in all instars. Whenfullgrown, it is a dead-white maggot, with black hook-like

*Drake (1920) published recognizable photographs of both sexes, but hisdesignations are erroneous. Osten Sacken, in a foot-note to the work of Say(1829), also has confused the sexes.


rasping mouth parts (mh) and a pair of black anal stigmata. (ans)It is quite robust, and although its greatest circumference isabout midway of its length, it can hardly be called fusiform,since it tapers away to a point in front, while the anal end isblunt. It is about 10 mm. long by 3.5 mm. in diameter, a sur-prising size when one considers that the adult host measuresbut 15 mm. in length.

The structure of the cephalo-pharnygeal skeleton, and thearrangement of the slits in the anal stigmata vary in the dif-ferent species, and figures of these organs are therefore includedin the plates. No sign of the parastomal sclerites mentionedby Banks (1912) as occurring in certain muscoid larvm could befound in the cephalo-pharyngeal skeleton of T. pennipes.

PuPARIUM. P1. III, figs. 19, 20, 21.

The pupa itself has not been observed. The pupariumwhich encloses it, however, ih of a deep reddish-black color,cylindrical in shape, and rounded at both ends. It is formedfrom the skin of the mature larva, and upon it the anal stigmataappear as twin tubefcies at the posterior end. The pupariaaverage about 7.5 mm. in length and 3.5 mm. in diameter. Atthe anterior end, before the emergence of the adult fly, a trans-verse split occurs, reaching backward nearly a quarter of thelength of the puparium. The split then extends around thecircumference, this resulting in the formation of two flaps whichare pushed aside by the ptilinum of the emerging adult.

Some time after the examinations of the puparium had beenfinished by the writer, the work of Greene (1922) on the pupariaof muscoid flies came to hand. The puparium of T. pennipesis there figured and discussed, and significant characters com-pared with those of the puparia of other species.

Aldrich, J. M.1905.

1915.

BIBLIOGRAPHY.

A catalogue of North American Diptera. Smith-sonian Misc. Coll., vol. 16, no. 1444, p. 425.Collecting in Tachinidm. Ann. Ent. Soc. America,vol. 8, p. 83. (Distribution of T. pennipes.)

70 Psyche [April

Banks, Nathan.1912. On the Structure of Certain Dipterous Larvm,

with Particular Reference to Those in HumanFoods. U. S. Dept. Agric., Bur. Ent., Tech.Set., Bull. 22.

Brauer, F. and Bergenstamm, J. E. V.1891. Die Zweifliigler des Kaiserlichen Museums, vol. 5,

p. 412 (calls T. pennipes the male, and T. pyrrho-gaster and T. ciliata of Wiedemann the female).

Chittenden, F. H.1899. Some Insects Injurious to Garden and Orchard

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1908. The Common Squash Bug. U. S. Dept. Agric.,Bur. Ent., Circ. 39, 2nd ed. p. 9. (Mentionsparasitism by T. pennipes.)

Cook, A. J.1889. A squash Bug Parasite. 2nd Ann. Rept.

Michigan Agric. Exp. Sta.--Rept. of Entomologistpp. 88-103. Also in Ann. Rept. of the Sec. ofthe Michigan State Bd. Agric., p. 151. (Givesfirst account of parasitism which names T.pennipes.)

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The Biology of Trichopoda pennipes Fab. 71

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1923. Preliminary note on the Terminology Appliedto the Parts of an Insect’s Leg. Canad. Ent.,vol. 55, no. 6, p. 130.

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Quart. Bull. State Plant Bd. Florida, vol. 4,pp. 41-94. (Treats of T. pennipes on pp. 67-74,87-88).

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nipes, p. 219.9, Thereva hirtipes; p. 315.9, Ocypteraciliata, later declared synonyms; and p. 327.5,Dictya pennipes, change of genus from Musca.)

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Sci., Torino, (2) vol. 44, p. 6 and 7. (T. pyrr-hogaster and T. pennipes.)

Girault, A. A.1904. Anasa tristis De Geer; History of Confined Adults.

In Ent. News, Vol. XV, p. 335. (Recbrds breedingT. pennipes.)

Greene, Chas. T.1922. An illustrated Synopsis of the Puparia of 100

Muscoid Flies (Diptera). Proc. U. S. Nat.Mus. vol. 60, Art. 10, pp. 37, figs. 99.

2 Psyche [April

Howard, L. O.1904. Insect Book. Plate XV, figs 25. (Color illustra-

tion of T. pennipes, female.)

Jones, Thos. H.1918. The Southern Green Plant Bug. U. S. Dept.

Agric., Bull. 689, p. 22. (Records T. pennipes asparasitic upon Nezara viridula.)

Latreille, P. A.1829. Cuvier’s Regne Animale, vol. 5, p. 512. (Erection

of genus Trichopoda.)

Malloch, J. R.1923. A New Character for Differentiating the Families

of Muscoidea. In En. News. vol. XXXIV,pp. 57-58.

Morrill, A. W.1910. Plant Bugs Injurious to Coton Bolls. U.S.

Dept. Agric., Bull. 86, p. 92. (Reared T. pennipesfrom Leptoglossus oppositus.)

Osten Sacken, C. R.1878. Catalogue of the Described Diptera of North

America, 2nd edit. Smithsonian Misc. Coll., vol.16, no. 270. (T. pennipes, p. 146.).

Packard A. S.1875. Tachina Parasite of the Squash Bug. In Am-

erican :Natural vol. 9, p. 513. (Evidently theearliest account of the habits of T. pennipes.)

Peterson, Alvah.1916. The Head Capsule and Mouthpurts of Dipteru.

Illinois Biol. Monog. vol. 3, No. 2, pp. 110;plate 25.


Robineau-Desvoidy, J. B.1830. Essai sur les Myodaires. (P. 283.1, change of

genus to Trichopoda; p. 284.2, T. flavicornis;and p. 285.7, T. haitensis, later declared synonyms.)

Say, Thomas.1829. Description of North American Dipterous Insects.

In Jour. Acad. Sci. Philadelphia vol. 6, p. 172.Complete Works, vol. 2, p. 364. (Phasia jugatoria,synonym of T. pennipes.)

Snodgrass, R. E.1909a. The Thoracic Tergum of Insects. In Ent. News,

vol. 20, pp. 97-103.1909b. The Thorax of Insects and the Articulation of the

Wings. In Proc. U. S. Nat. Mus., vol. 36, pp.511-595, plates 40-69.

Thompson, W. R.1910. Notes on the Pupation and Hibernation

Tachinid Parasites. Journ. Econ. Ent.,III, pp. 283-295.

ofvol.

Townsend, C.1893.

1897.

1908.

I-Io ToOn the Geographic Range and Distribution ofthe Genus Trichopoda. Ent. News, vol. 4,pp. 69-71.On a Collection of Diptera from the Lowlands ofRio Nautla in the State of Vera Cruz. Ann.and Mag. Nat. Hist. (6), vol. 20, p. 279.(Records T. pennipes.)A Record of Results from Rearings and Dissec-tions of Tachinidee. U. S. Dept. Agric., Bur.Ent., Tech. Ser. 12, part VI.

Van der Wulp, F. M.1888. Biologia

p. 434.Centrali-Americana. Dipt., Vol. 2,

T. pennipes.

74 Psyche [April

Walton, W. R.1909. An illustrated Glossary of Chmtotaxy and Ana-

tomical Terms used in Describing Diptera.Ent. News, vol. 20, pp. 307-319, plates XIII-XVI..

Watson, J. R.1918. Insects of a Citrus Grove. Univ. of Florida

Agric. Exp. Sta., Bull. 148, p. 2gl. (Records T.pennipes as parasitic upon Nezara viridula.)

Weed, C. M. and Conradi, A. F.1902. The Squash Bug. New Hampshire Agric. Exp.

Sta., Bull. 89. (An account of the parasitichabit of T. pennipes.)

Wiedemann, C. R. W.1830. Aussereuropaische Zweiflugelige Insekten, vol. 2,.

(p. 272.6; T. pyrrhogaster; p. 273.8, T. ciliata;p. 274.9, T. pennipes.)

Williston, S. W.1896. On the Diptera of St. Vincent (W. I.). Trans.

Ent. Soc. London for 1896, p. 352. (Records T.pennipes from St. Vincent.)

Wilson, C. E.1923. Insect Pests of Cotton in St. Croix and Means of

Combating Them. Virgin Islands Agric. Exp.Sta., Bull. No. 3. (Record of T. pennipes onp. 14).

Worthley, It. N.1923. The Squash Bug in Massachusetts. Jour Econ.

Ent. vol. 16, p. 78. (Chart showing parallelseasonal histories of T. pennipes and Anasatristis.)


Young, B. P.1921. Attachment of the Abdomen to the Thorax in

Diptera. Cornell Univ. Agric. Exp. Sta. Mere. 44,pp. 251-306, figs. 76.

EXPLANATION OF FIGURES.

Plate I.

Fig. 1.Fig. 2.Fig. 3.Fig. 4.

Head of male--front view.Head of malewside view, showing mouth parts.Dorsum of male thorax.Thorax of male--side view.

Plate II.

Fig. 5.

Fig. 8.Fig. 9.Fig. 10.Fig. 11.Fig. 12.Fig. 13.Fig. 14.

Wing of male, showing extent of ferrugineous spot.Cells labeled.

Wing of female. Veins labeled.Tibia of metathoracic leg, showing fringe of feather-

barbed setm.Terminal segments of tarsus of male.Abdomen of male--side view.Abdomen of female--side view.Abdomen of male--ventral view.Abdomen of female--ventral view.Male genitalia.Female genitalia.

Plate III

Fig. 15.Fig. 16.

Fig. 17.Fig. 18.

Ninth abdominal sternite of male.Egg. a, outline from side; b, from top; c, showing

hole in ventral surface after hatching.Mature larva.Cephalo-pharyngeal skeleton of larva, a and b, of

second stage (?) larva, side and top views; c andd, mature of larva, side and top views.

76

Fig. 19.Fig. 0.Fig. 21.

Psyche

Puparium, from the top.Empty puparium, from the side.Anal stigmata of puparium.

Plate IV

Dorsal view of male fly.

[April

1st A

abaadadc

aeaes

aexalansantaplt

arasasaAxCaxe

bpbbu

CCCcechbcpoCuC

CuCcx

dpbdv

emeps

ABBREVIATIONS

Nanal or 6th longitudinalvein

Nanterior basalar plate--adanaleanterior dorso-central

bristle--oedagus--mesoanepisternum or

mesopleuraaxillary excision--axillary lobe--anal stigmata--antennamanapleurotergite of post-

scutellum--aristaadventitious suture--anterior subalar plateaxillary (or anal) cellmaxillary cord

--basiproboscismbutton

--costal veinmcostal cell--compound eye--chitinized box--cili of posterior orbitkcubital (3rd basal or

anal) cell--3rd posterior cell--coxa

---distiproboscis---dorsal valve (?) of ovi-

positor

--metaepimeron---empodiumNmetaepisternum

fbfc

fefsfuful

gagegfgogPhahchcvhphuhyhyphys

ia

kplt

leplp

M

Ml/2M+CuMC1MC2MC

frontal bristles-fronto-clypeus (facial de-

pression, facial plate)--femur--frontal suturemfurca--fulcrum

--galeaNgena (cheek)Ngenita! furcagonopophyses--genital prong--halteremhumeral callus--humeral cross-vein--hypopleural bristles--humeral bristles--hyoidmhypopharynx--hypostomal sclerite

mintra-alar bristleinner valve of ovipositorminner vertical bristle

--katapleurotergite of post-scutellum

mlabrum-epipharynx--lateral plate

mmedla--medial (posterior)cross-

vein--4th longitudinal vein--Sth longitudinal vein--medial (2nd basal) cell--discal cellk2nd posterior cell

PSYClAE x924 PLATE

WORTHLEYBioIogy of Trichopoda pennipes

PsYca 924 PLA_TE II

Fig.Fig"

7i

PSYCHE 1924 PLATE III

Fig. 18

Fig.20

allS

Fig.21

a

WORTI-ILEY---Biology of Trichopoda pennipes

PSYCHE, 924 PLATE IV

WORTHLEYBiology of Trichopoda pennipes

The Biology of Trichopoda pennipes Fab. 77

md

mepmh

mpbmrmsmt

mv

nnnp

obOCocp

Ppapac

pbpbapde

PgpgePPpsapscpstpsupt"

ptbptp

pvpvt

umandibles (great hooks)of larva R+a

--meropleurite R4+5mmouth hooks (or man-RC

dibles) of larvaumediproboscis R.C--mesopleural row RaC--marginal scutellar bristles--meditergite of postscu-r-m

tellum--median vitta of vertex--maxillary palpus

notale sammetanotum !sb--notopleural bristles SC

scocellar bristles SCC--ocelli slocciput sp

spv--prothorax ss--postalar bristles st--posterior acrostical stb

bristle stpproboscisposterior basalar platesty--posterior dorso-central sur

bristle--paraglossaepostgenaepropleural bristle--posterior subalar platemesoprescutumpseudotrachaepresutural bristleposttergite of mesoscu-

tellumpteropleural bristlempteropleurite or mesoan-

epimeron---pulvillusposterior vertical bristles

--lst longitudinal vein---2nd longitudinal vein3rd longitudinal veinradial (1st basal )cellmarginal cellsubmarginal celllst posterior or apical

cellradio-medial (anterior)

cross-vein

s slitsl, s, etc. abdominal sternites

supra-alar bristlescutellar bridgesubcosta (auxiliary vein)mesoscutummsubcostal cellmesoscutellum--spiracleuspurious veinsubstigmal bristlestipessternopleural bristlesmesosternopleurite

(sternopleura)stylususuralare

t, t, etc. abdominal tergitesta tarsustb tibiatg tegulathe thecato tormaetr trochantertt tentorial thickenings

v vertexvib vibrissaevv ventral valve of ovi-

positor

wp wing processR --radius

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