Systematic Entomology (1986) 11, 377-387

Parhadrestiinae, a new subfamily for Parhadrestia James and Cretaceogaster Teskey (Diptera: )

NORMAN E. WOODLEY Systematic Entomology Laboratory, USDA

ABSTRACT. A new subfamily of Stratiomyidae is proposed for Parhadrestia James and Cretaceogaster Teskey (fossil from Upper Cre- taceous Canadian amber). Evidence is delimited that indicates that this subfamily is the sister-group to all other known stratiomyids. Taxa in the subfamily are systematically described, including a new species, Parhadrestia curico, from Chile.

Introduction character states has been determined by James (1975) described the stratiomyid genus outgroup comparisons. Elsewhere (Woodley, in Parhadrestia based on a single species he had preparation) I have shown that the discovered from Chile, P.atava. He placed the are clearly the sister-group to the Stratiomyidae, genus in the presumably because it so they have been used as the functional possesses an abdomen with seven larger seg- outgroup in this study. In the same work I have ments, a plesiomorphic feature in the family. In discussed the autopomorphic character states connection with a study of the world genera of used to characterize stratiomyids, so they are not Beridinae, I have re-examined the genus, and repeated here. Also in that work I discuss the have concluded that it is actually the sister-group characters which support the cladistic placement to all known extant stratiomyids. Also, of the , Beridinae, and remaining re-examination of the only known specimen of subfamilies of Stratiomyidae as shown in Fig. 1, Cretaceogaster pygmaeus Teskey (1971), a fossil so this argumentation is also omitted in the pre- from the Canadian amber (Upper ), sent paper and these characters do not bear indicates that it is cladistically related to numbers in the figure. In the following discus- Parhadrestia. Following a discussion of these sion, the initial statement is the apomorphic relationships, I propose a new subfamily for character state for each character in question. these two genera. Terminology of morphologi- (1) Hypandrium at least partly fused to cal features follows that of McAlpine (1981). gonocoxites. In all known stratiomyids except Parhadrestia and Cretaceogaster, the hypandrium (ninth abdominal sternite) is at Cladistic relationships least partly fused to the gonocoxites. Only in Fig. 1 presents a cladogram showing the charac- some Chiromyzinae and Beridinae is it partly ters used to support the hypothesized relation- free; in other stratiomyids it is completely fused ships between Parhadrestia, Cretaceogaster, and to the gonocoxites. In Parhadrestia (Figs. 13, other known stratiomyids. The polarity of the 18), the hypandrium is completely free from the gonocoxites, and is apparently free in Cre- taceogaster. Teskey (1971: Fig. 4) has illustrated Correspondence: Dr Norman E. Woodley, Sys- tematic Entomology Laboratory, IIBIII, ARS, the male genitalia of the fossil, and a view of USDA, c/o U.S. National Museum of Natural Parhadrestia from the same angle is remarkably History, NHB 168, Washington, DC 20560, U.S.A. similar. The structure of the hypandrium is quite

377 378 Norman E. Woodtey

midtibial spur lost 9A 1 midtibial spur 9 1 flagellomere aA 2 flagellomeres 8 Mg absent 7 palpi one-segmented 6 AS6.7 reduced tergal grooves + larval setae tufted ? ? aedeagus trifid ? scutellar spines mouthparts reduced cell cup truncate 5 R4 8R5 subparallel 4 8AS reducedllost 3 parameres reduced 2 hypandrium fused ii i

FIG. 1. Cladogram showing relationships of the tww genera of Parhadrestiinae to other . Character statements at thc left indicate the apomorphic character state. Unnumbered characters are not discussed in the text (see Woodley, in preparation). Question marks indicate characters which cannot be viewed in the fossil holotype of Cretaceogaster. Open squares indicate plesiomorphic character states, filled squares indicate apomorphic character states, and half-filled squares indicate that both character states occur in that lineage. Purhudrestiinae, primitive Stratiomyidae 379 different In the Xylomyidae, because their geni- nite, which is often modified in various ways talia are highly modified from the general such that its shape is often species specific (e.g. plesiomorphic form found in ‘lower Webb, 1984: 251). ’. Figures in Nagatomi & Tanaka (4) Wing veins R4 and R5 nearly parallel (1971) indicate that the hypandrium is quite apically. This character state is unique to reduced in size and is fused to the gonocoxites at Parhudrestia and Cretaceogaster. In its anterior corners in xylomyids. Fusion of the Parhadrestia (Fig. 8) the branches of R4+s are hypandrium to the gonocoxites is clearly very long, and are parallel for much of their apomorphic, as noted by Stuckenberg (1973). length. In Cretaceogaster (Fig. 9) the position of (2) Fused parameres reduced in size. In the fork is quite similar, but the branches are Purhadrestia, the parameres of the aedeagal shorter and parallel only near their apices. In all complex have apparently become fused and other stratiomyids and in Xylomyidae, R4 is form a dorsal hood over the aedeagus (Figs. 15- shorter to much shorter than R5, and is strongly 17), much like that found in (see divergent from it, ending considerably anterior McAlpine, 1981, for a discussion of parameres to the wing apex. within the Diptera). This fusion is evidently (5) Apical section of wing vein CuA, (closing independent of that found in Tabanoidea, as the cell cup) short and straight. In both Parhadrestia parameres are not fused in Xylomyidae (see and Cretaceogaster (Figs. 8, 9), vein CuA, is RozkoSny, 1973: Figs. 21, 25; McAlpine, 1981: short and straight, forming a truncate closure of Fig. 2.122). In addition, they are not mem- the anal cell (cell cup). In all other known branous in Purhadrestia. In the remaining stratiomyids and xylomyids, vein CuA, is stratiomyids this structure has become markedly slightly to strongly arcuate, so that the end of cell reduced in size and essentially forms the small cup appears rounded, or sometimes acute, but structure by which the aedeagal complex is never truncate. This is perhaps the strongest attached to the gonocoxal apodemes. It seems evidence for the close relationship of unlikely that the lateral lobes of the trifid aedea- Parhadrestiu and Cretaceogaster. gal complex found in many stratiomyids are (6) Palpi one-segmented. In Parhudrestia, the homologous with parameres as thought by palpi are much reduced in size and are one- RozkoSny (1982), because neither Parhudrestia segmented. This structure cannot be seen in Cre- or Chiromyzinae have a trifid aedeagal complex. taceogaster. In most other stratiomyids and In the Chiromyzinae the aedeagus is apparently xylomyids, the palpi are large and clearly two- a simple tube (see Nagatomi & Yukawa, 1969), segmented. They are one-segmented in although the apex may be vaguely lobed Chiromyzinae, but in members of this subfamily (Nagatomi & Yukawa, 1969: Fig. 10). Much I have examined they are strongly ovoid to remains to be learned about the male genitalia of nearly spherical, not small and peg-like as in Chiromyzinae, and this knowledge will be Purhadrestia. Presumably this is not syn- important in interpreting the structures of the apomorphic in these two primitive groups, but aedeagal complex in more apomorphic more knowledge of the poorly known stratiomyids. Chiromyzinae is needed before this can be eluci- (3) Eighth abdominal sternite in male very dated. In a few other scattered taxa of reduced or lost. In nearly all stratiomyids, the stratiomyids a one-segmented palpus is present, eighth abdominal sternite of the male is com- but these are clearly derived from related genera pletely unsclerotized, only membrane remaining which have two-segmented palpi. In one genus in its place. In some Beridinae and a few of Xylomyidae, Xyfomya,the palpi are one-seg- Chiromyzinae, small vestiges of sclerotization mented. They are not reduced in size. Presuma- are still present medially, but in other members bly this state is autapomorphic, for the genus, as of these subfamilies the sternite is lost. In other xylomyids have two-segmented palpi. Parhadrestia the eighth sternite is quite large, (7) Wing vein M3 absent beyond discal cell. In although somewhat smaller than the eighth both Parhadrestiu and Cretaceogaster (Figs. 8, tergite. In relative size, it is much larger than in 9), vein M3is absent. In most other stratiomyids any other known stratiomyids. This region of the and in all xylornyids M3 is present. It is absent abdomen cannot be seen in Cretaceogaster. from some Chiromyzinae and Beridinae, and all Xylomyidae have a relatively large eighth ster- (which are grouped in ‘other 380 Norman E. Woodley

subfamilies’ in Fig. 1). In the first two sub- tibia. A single spur is found only in a few families, intermediate states can be seen in Chiromyzinae and in scattered Beridinae, as which Mi is shortened. This character is well as all known Antissinae and in the enigmatic apparently quite plastic, and loss of M3 has evi- genus Rondani. dently occurred independently several times. (8) Antennal flagellum strongly fused, two flagellomeres present. Character state distribu- PARHADRESTIINAE new subfamily tion discussed under 8A below. (8A) Antennal flagellum composed of a single Diagnosis. Small, drab stratiomyids with a flagellomere. In Cretaceogaster, the antennal short abdomen and rather long wings. Defining flagellum is largely fused, with a large basal autapomorphic states (numbered as above) are flagellomere and a short, style-like apical (6) one-segmented palpi, (7) wing vein M3 flagellomere. Teskey (1971: Fig. 1) showed the absent beyond discal cell, (4) fork of wing vein flagellum as a single flagellomere, but my obser- R4+5nearly parallel apically, with R4 ending vations indicate that two flagellomeres are pre- nearer and the wing apex than in other sent. Parhadrestia (Figs. 2, 4, 6) has the KS Stratiomyoidea, (5) wing vein CuAz straight, flagellum entirely fused into a single thus the end of cell cup truncate, and (8) the tlagellomere. James (1975: 21) claimed that at largely fused antennal flagellum (two or fewer least several were present (‘flagellomeres well antennal flagellomeres). For purposes of routine fused, difficult to differentiate, only the terminal identification, characters 4 and 5, each unique to 2 more discrete but short and forming at most a the subfamily, will serve to diagnose specimens. feebly differentiated style’), but I have cleared Other characters, although plesiomorphic, use- antennae of both sexes, and the flagellum shows ful in identification include the completely free no evidence of subdivisions. Most stratiomyids hypandrium, the fused parameres forming a dor- (and many other , including all sal hood over the aedeagus, the unarmed Xylomyidae) have eight antennal flagellomeres. scutellum, the presence of one or two apical Within the Stratiomyidae, only in a few spurs on the middle tibia, and the abdomen with Chiromyzinae and in Draymonia Aubertin seven large, undifferentiated segments. (Beridinae) are there fewer than five The type-genus of the subfamily is flagellomeres in the antennal flagellum. As far Parhadrestia James (1975: Zl), the only known as I am aware, Walker (Chiromyzinae) is extant genus in the subfamily. Also included is the only other stratiomyid that has a completely Cretaceogaster Teskey (1971: 1660) from Upper fused flagellum, which has presumably arisen Cretaceous Canadian amber. In the following independently from that found in Parhadrestia, section I am presenting redescriptions of both as a less fused flagellum is found in other genera to conform to my descriptions of genera chiromyzines. of Beridinae (Woodley, in preparation). I also (9) One apical spur present on middle tibia. describe a second species of Parhadrestia. Character state distribution discussed under 9A below. (9A) Both spurs lost on middle tibia. PARHADRESTIA James Parhadrestia has a single spur on the middle tibia, not reported by James (1975). Cre- Parhadrestiu James, 1975: 21. Type-species, taceogaster has two spurs on the middle tibia. P.atava James, by original designation. Teskey (1971) stated that none were present, but I have seen them upon re-examination of the Diagnosis. The only extant genus of holotype fossil. They are small and difficult to Parhadrestiinae. Defining autapomorphic char- see, as they are in many extant stratiomyids that acter states include the antennal flagellum com- possess them. The only other stratiomyid genus posed of a single flagellomere and the presence with two tibia1 spurs is Beridops Enderlein of only one midtibial spur. Other character (Beridinae), although this character state is states which will distinguish Parhadrestia from widespread in Xylomyidae and in the extant stratiomyids include the large, fused Xylophagoidea. Most stratiomyids exhibit a parameres over the aedeagus; the completely further loss in having no spurs on the middle free hypandrium; the very long, parallel Parhndrestiinae, primitive Strationiyidae 381

FIGS. 2-7. Headsof Purhudrestiu species: 2,3, male of P.curico, new species; 4,5, male of P.atava James; 6, 7, female of P.utuvu James. branches of the fork of vein R4+s; and the prominent in male, nearly flat in female, (6) straight vein CuA, which closes cell cup in a lower frons short pilose, faintly tomentose just truncate manner. above antennae in female only, (7) upper from Description. Head (Figs. 2-7): (1) slightly evenly pilose, (8) face wide in both sexes, diverg- higher than long (cf: 1.2, 0 1.3), (2) wider than ing ventrally, strongly receding in male, less so in high (1.4), (3) lower frons of male small, triangu- female, (9) short pilose, rather evenly but spar- lar; very wide in female, slightly convergent sely tomentose, however with densely tomen- toward vertex, (4) upper frons of male very tose margins along eyes, very narrow in male, reduced; wide in female, convergent toward ver- wider and more conspicuous ventrally in female, tex, (5) ocellar tubercle relatively large, slightly (10) upper occiput convex and rounded, more 382 Norman E. Woodley

9

FIGS. 8-12. 8, Wing of Parhadrestia alava James. 9, Wing of Cretaceogaster pygmaeus Teskey (after Teskey. 1971: Fig. 3). 10. Abdomen of female of Parhadrestio afava James, cleared, in dorsal view. 11, Female spermatheca of P.atava James. 12. Female genital furca of P.utava James, ventral view. Parhadrestiinae, primitive Stratiomyidae 383 prominent in female, (11) short pilose, (12) eighth relatively undifferentiated, (54) shallow lower occiput rounded and prominent in male, subapical tergal grooves present on segments more so in female, (13) eyes of male holoptic, 2-6, (55) first sternite without erect hairs. Male sharply divided into an upper field of larger genitalia (Figs. 13-18) with (56) capsule quitc ommatidia and a lower field with smaller rounded, (57) gonocoxal apodemes slender, ommatidia; widely dichoptic in female, with moderately long, extending anterior to basal uniform ommatidia, (14) posterior eye margin margin of capsule, (58) posterior margin of rounded in both sexes, (15) eyes pilose, the hairs hypandrium rounded, (59) hypandrium (Figs. moderately to very dense, (16) antennae 13, 18) completely free of gonocoxites, (59A) inserted slightly below middle of head in profile posteroventral margins of gonocoxites in male (0.58), slightly above middle in female triangularly produced below gonostyli, (60) (0.46), (17) 0.49-0.67 length of head, (18) gonostyli slender, simple, somewhat club- flagellum 2.57-2.67 times length of shaped apically, (61) aedeagal membrane scape+pedicel, (19) completely fused, formed apparently absent (62, 63) aedeagal complcx of a single flagellomere, (20) conically tapered, a aparently simple, not trifid, (64) fused para- little more cylindrical in female, (21) with usual meres large, forming a hood-like shield over velvety vestiture, without longer hairs, (22, 23) acdeagus (Figs. 15-17), (65) eighth sternite palpi one-segmented, (24) reduced, short large, over one-half as wide as eighth tergite and cylindrical, (25) with a few apical hairs. Thorax: fully as long, slightly narrowed posteriorly, (66) (26) generally pilose dorsally, pleura in large epandrium (Fig. 14) very short, quite broad and part bare, with little tomentum, (27) wide, (66A) cerci short, ovoid. Female genitalia anepimeron bare, (28) meron+katepimeron with (67) furca (Fig. 12) longer than wide, (68) with a few hairs at extreme anteroventral posterior bridge unsclerotized, apparently lost, margin, (29) subscutellum mostly tomentose, (69) median aperture very large, (70) post- (30) scutellum unarmed, (31) middle tibia with a erolateral processes absent, (71) ninth tergite single short apical spur, (32) hind femora 1.2- narrow, poorly defined, only narrowly attached 1.3 times as long as middle femora, (33) slender, to furca, (72) spermathecae three, along with thc thickest medially, (34) without structural simple ducts completely unpigmented (Fig. 1 I), modifications, (35) hind tibiae very slender, (73) cerci two-segmented, (74) short, first seg- cylindrical, (36) first tarsomere of hind leg 0.86- ment much thicker than second. 0.89 length of tarsomeres two to five, (37) Distribution. Known only from Coquimbo slender, not at all swollen, (38) wings (Fig. 8) not and Maule Provinces, Chile. sexually dimorphic, (39) hyaline, (40) com- Remarks. Little is known about this genus pletely set with microtrichia, (41) subcosta beyond the few collected specimens. James straight, complete, (42) cell r, 8.5-10.0 times (1975) placed the genus in the Beridinae based longer than broad, unusual shape corrolated upon plesiomorphic character states. Two with insertion and shape of RZ+3,(43) R2+,aris- species are known, one of which is described ing far proximal to r-m and somewhat proximal below as new. It is interesting to note that the to base of cell dm, (44) R,,, straight, (45) R4 prosternum is small and completely separate beyond fork strongly curved basally, long and from the proepisternum. Fusion of the proster- subparallel to R5apically, (46) MI and M2 separ- num and proepisternum was implied by Hennig ate to contiguous and short petiolate at discal (1967) to be synapomorphic between cell, (47) M3absent beyond discal cell, (48) base Xylomyidae and Stratiomyidae. of CuA, longer than base of M3, (49) apex of CuAl slightly arcuate, (50) alula ovoid, rather broad, posterior margin evenly rounded, inci- Key to species of Parhadrestia sion present but not sharp, completely set with microtrichia. Abdomen: (51) short and small, 1 Antennal flagellum longer, more gradually attenu- ate apically (Figs. 4, 6), dark in male (may be about 1.7 times longer than broad, (52) broadest yellowish in female); male face wide (Fig. 5); near base, tapering apically, tergites and pilosity of eyes (Fig. 4) sparser than in P.curico especially sternites reduced in female (Fig. lo), aluva pleural and intersegmental membranes quite Antennal flagellum shorter, more strongly attenu- extensive, (53) seven larger segments, even ate apically (Fig. 2), entirely yellowish; male face 384 Norman E. Woodley

FIGS. 13-18. Male genitalia of Parhadrestia species. 13-16, Parhadrestia atava James. 13, Genital capsule, dorsal view. 14, Epandrium and post-genital segments, dorsal vicw. 15, Aedeagal complex, lateral vicw. 16, Aedeagal complex, dorsal view. 17-18, P.curico, new species. 17, Aedeagal complex, dorsal view. 18, Genital capsule, dorsal view. Abbreviations: ard, aedeagus; cerc, cercus: epand. epandriuni; Roncx, gonocoxites; goncx apod, gonocoxal apodeme; gonst, gonostylus: hypd, hypandrium; pm, paramcrcs.

narrower than in P.urava (Fig. 3); pilosity of eyes characters are best appreciated when both (Fig. 2) denser than in P.atava ...... curico species can be examined and compared, Description. Male. Head (Figs. 4, 5) black, vertex 0.16 width of head; frons very reduced, Parhadrestia atava James lower frons slightly impressed medially ; face very wide; tomentum of head whitish grey. Parhadrestia atava James, 1975: 21. dense along narrow facial margins, sparse on Type material. Holotype male labelled: rest of face and much of occiput; pilosity of head ‘CHILE, Coquimbo Prov. Hacienda lllapel, Rio whitish yellow, short, longest on genae and Illapel, 600-900 m, X-19-1966, E. I. Schlinger, lower occiput, where it is slightly longer than M. E. Irwin/Parhadrestia atava James antennal segments 1+2, short and moderately HOLOTYPE’ (temporarily in California dense on eyes; antennae (Fig. 4) short, 0.55- Academy of Sciences). The spccinien is in 0.61 length of head, ratio of segments (holotype) excellent condition. 3 :5: 17, black, apex of second segment very nar- Diagnosis. The key characters will separate rowly yellowish; flagellum gradually attenuate this species from P.curico. The longer, dark toward apex; basal segments with a few short, antennal flagellum of the male is the best charac- pale hairs and some pale, velvety vestiture, ter for identification, as the other diagnostic especially on second segment, flagellum with Parhadrestiinae, primitive Stratiomyidae 385 dense velvety vestiture; palpi brownish black, branous areas between sclerites; tergites with small and difficult to see without dissection; pro- little tomentum on basal areas. Female geni- boscis small and compact, brownish. Thorax talia (Figs. 11, 12) as in generic description. black, but posterior part of post-pronotal lobe, Length, 3.8-4.0 mm. postalar callus, and parts of pleural sclerites Specimens examined. Holotype 6,and 16, below wing base brownish; mesonotum and 29 paratypes, and one additional 6,all with scutellum finely, densely punctate; mostly with- same data as holotype (Washington State out tomentum, but silvery white tomentum University). present on prosternum, posterior part of metakatepisternum, laterotergite, and most of subscutellum; mesonotum and scutellum Parhadrestia curico sp.n. evenly, densely pilose with whitish yellow, sub- appressed hairs; pleural areas mostly bare and Type material. Holotype male labelled: shiny, with short pale pilosity present on pro- ‘CHILE: Curico Prov. [now part of Maule Pro- episternum, posterior half of anepisternum, vince] Estero la Jaula, I. 1964. L. Pena upper margin of katepisternum, posterior NothofaguslHOLOTYPE 6 Purhudrestia margin of metakatepisternum, and most of curico N. E. Woodley.’ (Canadian National Col- katatergite; legs blackish brown but yellowish on lection, Ottawa, Canada). apices of femora, bases and extreme apices of Diagnosis. Very similar to P.atava, best iden- tibiae, and bases of first tarsomeres; legs evenly tified by its shorter, yellowish antenna1 set with pale hairs, slightly longer and more erect flagellum. It is also slightly smaller in size, but on femora; wings hyaline, cell r, yellowish with the few known specimens of the genus this brown in apical part, veins brownish; veins MI could lack significance. and Mzseparate at discal cell; halter dark brown, Description. Male. Differs from P.atava as stem more yellowish. Abdomen brownish black, follows: Head (Figs. 2, 3) with vertex narrower, cerci and apical portions of genitalia yellowish; 0.13 width of head; face narrower; pilosity of tergites except laterally and entire sternites with head slightly shorter and sparser, but very dense pale tomentum of moderate density, more evi- on eyes, especially dorsally; occiput narrower dent on sternites; lateral margins of tergites with viewed in profile; antennae (Fig. 2) shorter, 0.49 moderately long, erect, yellowish white hairs, length of head, ratio of segments 2:4: 15, first sternites with thinly scattered, pale pilosity. segment and basal half of second brownish Male genitalia (Figs. 13-16) as in generic black, rest yellowish with flagellum gradually description, capsule longer in relation to its but slightly darker apically; flagellum more width than in P.ciirico, aedeagal complex also ovoid and strongly attenuate apically. Thorax slightly longer and narrower, with proximal por- with mesonotal pilosity somewhat shorter and tion of aedeagus narrower than in P.curico. denser; wing with Mi and M, short petiolate at Length, 3.1-3.2 mm. discal cell. Malegenitalia (Figs. 17, 18) with cap- Female. Differs from male as follows: Head sule shorter in relation to its width than in (Figs. 6,7) with fronsverywide, 0.41-0.42 width P.atava, and aedeagal complex slightly shorter of head, lower part slightly concave medially and broader, with proximal part of aedeagus toward antennae; tomentum of facial margin wider. Length, 2.7 mm. wider, more restricted to ventral areas; anten- Female. Unknown. nae 0.65-0.67 length of head, slightly longer Remarks. The new species epithet, a noun in than in male, relative lengths of segments apposition, refers to the province in which the 3 : 4.5 : 21, flagellum (Fig. 6) less strongly holotype was collected. tapered toward apex, may be extensively This species is extremely . similar to yellowish on basal portion. Thorax with Parhadrestia atava. However, the structure and brownish areas as in male, but may be slightly coloration of the antennae are distinctly paler and more noticeable; pilosity as in male different. The much denser pilosity of the eyes is but slightly shorter; first tarsomeres more dis- also distinctive, but is best appreciated if both tinctly yellowish basally. Abdomen (Fig. 10) species can be compared. which is also true of irregularly brownish yellow ventrally; relatively the narrower face. The differences in the male larger than in male, with more extensive mern- genitalia are slight, and when more material of 386 Norman E. Woodley both species is available, it may be found that the mal to r-m (44) R4+5straight (45) R4beyond fork differences seen here are obscured by variation short, subparallel to R5 near their apices (46) MI in each. The genitalia of these small are and Mzseparate at discal cell (47) M3absent (48) difficult to examine, especially the structures of CuAl widely fused to discal cell (49) CuA, the aedeagal complex. Also, the petiolate nature straight apically. Abdomen (52) short, tapered of veins M, and M2at the discal cell may not be a posteriorly (53) apparently with seven constant characteristic of the species. undifferentiated segments (54) with some evi- dence of subapical tergal grooves. Male genitalia with (58) hypandrium rounded posteriorly (59) CRETACEOGASTER Teskey apparently free from gonocoxites (59A) post- eroventral margins of gonocoxites triangularly Cretaceogaster Teskey, 1971: 1660. Type- produced below gonostyli (60) gonostyli, or at a species, C.pygmaeus Teskey, by original least part of them, sharp apically, (66A) cerci designation. rather short. Distribution. Known only from the type Diagnosis. Similar in general appearance to locality: Cedar Lake, Manitoba, Canada. Parhadrestia, but known only as a fossil. Defin- Remarks. Known only from the type-species, ing autapomorphic character states not deter- Cretaceogaster pygmaeus Teskey, which is mined, but may be distinguished from known from a unique fossil specimen. This Parhadrestia by the shorter fork of R4+5and by specimen is the oldest known fossil stratiomyid the differently shaped antennal flagellum which which can be placed in the family with certainty has a style-like second flagellomere. It also has (Teskey, 1971). It was originally placed in the two apical spurs on the middle tibia. Cre- subfamily Pachygastrinae, but characters used iaceogasier is presumably extinct, known only to support this placement are mostly not from Upper Cretaceous Canadian amber. autapomorphic for that subfamily. Fusion of Description. Based upon the holotype male. antennal flagellomeres is found in Characters with numbers omitted cannot be pachygastrines, but not with reduction to only observed in the fossil. Head (1, 2) globose, one or two flagellomeres. This is only found in nearly hemispherical (3, 4) male frons reduced some Chiromyzinae, another primitive group of by enlarged eyes, (5) ocellar tubercle fairly stratiomyids. The unarmed scutellum and fusion large, slightly prominent (8) face short and wide, of vein CuA, to the discal cell are plesiomorphic sides diverging ventrally, receding (9) short for the entire family (Woodley, in preparation). pilose (10) upper occiput with narrowly rounded Large eyes are widespread throughout large por- margins, apparently nearly flat (13) eyes of male tions of the family. Only the loss of vein M3 can holoptic, upper ommatidia larger than lower be considered an autapomorphic feature of the ones (17) antennae shorter than head (18) Pachygastrinae, but this character state is also flagellum between 1.5 and 2.0 times as long as found in other stratiomyids, including scape+pedicel (19) apparently with two Parhadrestia and some Chiromyzinae and flagellomeres (20) first flagellomere short-coni- Beridinae. The latter two groups are, aside from cal, second short, gradually tapered to a rather the parhadrestiines, the two most plesiomorphic sharp apex. Thorax (26) generally with short, groups of stratiomyids. Teskey’s statement that dense pilosity (30) scutellum simple, unarmed the larger sixth and seventh segments of the (31) middle tibia with two very short apical spurs abdomen ‘suggests an association with the (32) hind femora 1.36 times length of middle Beridinae’ is correct, as the Pachygastrinae have femora (33) slender, thickest medially (34) sim- only five larger abdominal segments. This is a ple, without modifications (35) hind tibiae very synapomorphic feature for all stratiomyids slender, cylindrical (36) first tarsomere of hind excluding Parhadrestiinae, Chiromyzinae leg 0.82 length of tarsomeres 2-5 (37) slender, and Beridinae [i.e. ‘other subfamilies’ in Fig. 11 not inflated, wings (Fig. 9) (39) hyaline (40) (Hennig, 1973). It would have been difficult to probably evenly set with microtrichia (only some realize the significance of this fossil in 1971, as are visible), (41) subcosta straight, complete Parhadrestia had not yet been discovered. The (42) cell rl 5.6 times longer than wide, elongate accuracy of the original figures were instrumen- triangular in shape (43) R2+3arising well proxi- tal in my recognition of Cretaceogaster as being a Parlradrestiinae, primitive Stratiomyidae 387 close relative of Parhadrestia. The overall Comparative Zoology, Harvard University, similarity in habitus between the actual fossil of Cambridge, Massachusetts, kindly made the Cretaceogaster and Parhadrestia is striking. The fossil specimen of Cretaceogaster available for relative plesiomorphic nature of the antennae study. I am very grateful to Art Borkent of the and midtibial spurs in Cretaceogaster are possi- Biosystematics Research Institute for his careful bly an indication that it is ancestral to review of the manuscript. Parhadrestia. Some notes on a few characters not yet dis- References cussed are worthwhile. The abdominal tergites are unfortunately not well preserved in the Hennig, W. (1967) Die sogennanten ‘niederen fossil, but show some indication that preapical Brachycera’ im Baltischen Bernstein (Diptera: , Xylomyidae, , tergal grooves may be present, as they are in Tabanidae). Stuitgarter Beitrage zur Naturkunde, Parhudrestia. The spine-like processes which 174, 1-51. Teskey surmised were part of the aedeagus are Hennig, W. (1973) Ordnung Diptera (Zweifliigler). intriguing. If they are indeed part of the aedea- Handhuch der Zoologie, 4(2) 2/31 (Lieferung 20), 1-337. gal complex, this must differ considerably from James, M.T. (1975) A preliminary review of the that found in Parhadrestia. It is unfortunate that Stratiomyidae of Chile. Part 11. Melanderia, 20, the aedeagus is not extruded further, such that i-iv, 1-28. the nature of the parameres could be ascer- McAlpine, J .F. (1981) Morphology and Terminology- Adults (chapter), pp. 9-63. In: McAlpine, J.F. et tained. Finally, the weakened state of the post- al. (eds.) Manual of Nearctic Diptera, Vol. 1. erior veins (essentially posterior to R,) is Research Branch, Agriculture Canada, Ottawa. interesting, as those of Parhadrestia are rather Monograph 27, 674pp. strong. This could be autapomorphic for Cre- Nagatomi, A. & Tanaka, A. (1971) The Solvidae of taceogaster. I suspect that vein MI separating the Japan (Diptera). Mushi, 45,101-146. Nagatomi, A. &Yukawa, J. (1969) The Chiromyzinae basal radial cell from the basal medial cell is from New Guinea (Diptera: Stratiomyidae). actually present (not shown by Teskey, 1971: Pacijic , 11, 633-643, Fig. 3), but quite faint and not easily seen in the RozkoSny, R. (1973) The Stratiomyioidea (Diptera) fossil. It is little more than a fold in Parhadrestia, of Fennoscandia and Denmark. In: Fauna Entomologica Scandinavica, Volume 1. Scandina- but is present. vian Science Press, Gadstrup. 140pp.+catalogue. RozkoSny, R. (1982) A Biosystematic Study of the European Stratiomyidae (Dipiera), Volume 1. Acknowledgments Introduction, Beridinae, and Stratio- rnyinae. Dr W. Junk, The Hague, Boston and Lon- don. viii+4Olpp. I wish to thank Paul H. Arnaud, Jr, of the Stuckenberg, B.R. (1973) The , a new California Academy of Sciences, San Francisco, family in the lower Brachycera (Diptera). Annals California, for the loan of the holotype of of the Natal Mweum, 21,649-673. Parhadrestia atava, temporarily in his care. Her- Teskey, H.J. (1971) A new soldier from Canadian amber (Diptera: Stratiornyidae). Canadian bert J. Teskey of the Biosystematics Research Entomologist, 103, 1659-1661. Institute, Ottawa, Canada, loaned the specimen Webb, D.W. (1984) A revision of the Nearcticspecies of Parhadrestia curico, and William J. Turner of of the family Solvidae Insecta: Diptera). Transac- the James Entomological Collection, tions of the American Entomological Society, 110, Washington State Univerisity, Pullman, 245-293. Washington, loaned additional material of P.ntava. Frank M. Carpenter of the Museum of Accepted 8 August 1985

27