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PARASITOIDS  ATTACKING  SYNANTHROPIC  FLIES

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Aleochara spp.

Alysia spp.

Aphaereta spp.

Australencyrtus Johnston & Tiegs

Australomalotylus Risbec, 1956

Braconidae

Chalcididae

Cynipidae

Diapriidae

Dicyclus pallinervosus Walker

Dirhinoides Masi

Dirhinoides mathuri Mani & Dubey

Dirhinus

Dirhinus anthracia Walker

Dirhinus bakeri (Crawford)

Dirhinus banksi Rohwer

Dirhinus frequens Masi

Dirhinus himalayanus Westwood

Dirhinus intermedius Mani & Dubey

Dirhinus luciliae Rohwer

Dirhinus luzonensis Rohwer

Dirhinus pachycerus Masi

Dirhinus sarcophagae Froggatt

Dirhinus vlasovi Nikolskaya

Dirrhinoidea Girault

Dirrhinus crythroceras Cameron

Dirrhinus Dalman

Dirrhinus ruficornis Cameron

Encyrtidae

Encyrtus vindemiae Rondani

Eniaca Kirby

Eniacella Girault

Eniacella bicornuticeps Girault

Eniacella rufricornis Girault

Eniacomorpha Girault

 

 

 

Eucoilidae

Figites spp.

Figitidae

Halizoa Förster

Halizoa rufipes Ashmead

Hexacola (= Trybliographa) spp

Ichneumonidae

Mormoniella Ashmead

Mormoniella brevicornis Ashmead

Muscidifurax Girault & Sanders

Muscidifurax raptor Girault & Sanders

Muscidifurax raptorellus Kogan & Legner

Muscidifurax raptoroides Kogan & Legner

Muscidifurax uniraptor Kogan & Legner

Muscidifurax zaraptor Kogan & Legner

Nasonia Ashmead

Nasonia brevicornis Ashmead

Nasonia vitripennis (Walker)

Pachycrepoideus vindemiae Rondani

Pareniaca Crawford

Pareniaca bakeri Crawford

Pareniaca trichophthalma Masi

Phaenopria spp.

Phygadeuon spp.

Platymesopus macellariae Brèthes

Prospalangia Brèthes

Prospalangia platensis Brèthes

Pteromalidae

Pteromalus abnormis Boheman

Pteromalus muscarum Hartig

Pteromalus subgen. Halizous Thomson

Pteromalus vitripennis Walker

Spalangia Latreille

?Spalangia abenabooi Girault

Spalangia astuta Förster

Spalangia atherigonae Risbec

Spalangia cameroni Perkins

 

 

Spalangia drosophilae Ashmead

Spalangia endius Walker

Spalangia formicaria Kieffer

Spalangia gemina Bouek

Spalangia hirta Haliday

Spalangia homalaspis Förster

Spalangia hyaloptera Förster

Spalangia longepetiolata Boucek

Spalangia melanogastra Masi

?Spalangia mors Girault

Spalangia muscae Howard

Spalangia muscarum Girault

Spalangia muscidarum Richardson

Spalangia muscidarum var. stomoxysiae Girault

Spalangia muscidarum var. texensis Girault

Spalangia nigra Latreille

Spalangia nigripes Curtis

Spalangia nigroaenea Curtis

Spalangia orientalis L. F. Graham

Spalangia philippinensis Fullaway

Spalangia rugosicollis Ashmead

Spalangia rugulosa Förster

Spalangia stomoxysiae Peck

Spalangia sundaica L. F. Graham

Sphegigaster spp.

Staphylinidae

Stictonotus insuetus Walker

Stilpnus spp.

Tachinaephagus Ashmead

Tachinaephagus javensis Subba-Rao

Tachinaephagus stomoxicida Subba-Rao

Tachinaephagus zealandicus Ashmead

Trichopria spp.

Trybliographa spp

Urolepis rufipes (Ashmead)

Urolepis Walker

 

 

 


DESCRIPTION OF SPECIES

 

 

Parasitic Staphylinidae (Coleoptera)

 

The front wings are without veins, hard or leathery, short, exposing much of abdomen (Fig. 49a-A).  The first visible abdominal sternite is not interrupted by hind coxal cavities.  Outer lobe or galea of maxillae not a segmented process.  Maxillary palpi are much shorter than antennae.  Hind tarsi with at least as many segments as front and middle tarsi.  Two important genera are Aleochara and Anotylus.  Key references are Thompson (1944), Peschke & Fuldner (1977), Seevers (1978), Borror et al. (1981), Peschke et al. (1987a), and Omar et al. (1991a)

 

1.  Aleochara spp. (Coleoptera: Staphylinidae: Aleocharinae)  (PHOTO)

 

These parasitoids are characterized by the forewings being developed into leathery shields (elytra) under which the hind wings (the organs of flight) are folded when at rest.  The elytra are short, leaving much of the abdomen exposed (Fig. 50a-A & B).  The abdomen is highly flexible as in Aleochara taeniata Erichson (Fig. 50a-C).  The genus Aleochara differs from other staphylinids in that the antennae are inserted on the face between the anterior margins of the eyes, the tarsi have 5 segments, the maxillary palpi have 5 segments and the labial palpi have 4 segments.  The two terminal segments of the palpus are much narrower than the preceding with the last segment minute.  All species of this genus in which life histories are known are solitary ectophagous parasitoids on the pupae of muscoid flies within the puparium (Kemner 1926, Legner et al. 1976, Lesne & Mercier 1922, Jones (1967), Moore & Legner 1971, 1973, 1974a,b; 1975, White & Legner 1966). 

 

Additional key references are  Coquillett (1891), Wadsworth (1915), Scott (1920), Kemner (1926), Soring (1927), Zorin (1927), Scheerpeltz (1933, 1934), Burks (1952), Read (1962), Fuldner (1963, 1964, 1968, 1971, 1973), Drea (1966), Horion (1967), Wingo et al. (1967), Allee (1969), Adashkevich (1970), Adaskevich & Perekrest (1973, 1974), Hünten (1971), Riegel (1971), Schneider (1971), Heller (1974, 1976), Heller & Treece (1976), Lohse (1974), Pfenning (1975), Schulz (1975), Watts & Combs (1975), Peschke & Fuldner (1977), Peschke & Metzler (1987),  Ienistea & Fabritius (1978, 1982), Kirknel (1978), Peschke (1978a,b,c; 1983, 1985a,b; 1986, 1987), Peschke et al. (1987b), Ursu & Sperantia (1978), Tawfik et al. (1980), Hertveldt et al. (1984a,b), Klimaszewski (1984), Klimaszewski & Blume (1986), Klimaszewski & Cervenka (1986), Hunter et al. (1985), Samsoe-Peteresen (1985, 1987), Whistlecraft et al. (1985), Gordon & Cornect (1986), Scott & Rutz (1988), and  Wright & Müller (1989).

 

Family Braconidae (Hymenoptera: Ichneumonoidea)

 

The fore wing costal cell is absent.  There is either one recurrent vein (Fig. 45b-A) or none.  Ventral abdominal segments are soft and membranous, with a median fold.  Abdomen is not much elongated and the propodeum is not prolonged beyond hind coxae.  These parasitoids are less than 12 mm long (Fig. 45b-B)  Two important genera are Alysia and Phaenocarpa.  Key references are Marsh & Altson (1920), Lima (1960, 1962), Tobias (1962, 1963), Riek (1970), Prince (1976), Richards (1977), Borror et al. (1981), Zhao (1984), Wharton (1976, 1986, 1987), Krombein et al. (1979), Gauld & Bolton (1988), and Goulet & Huber (1993).

 

2.  Alysia spp. (Hymenoptera: Braconidae)

 

The parasitoids of this genus are common in areas of moderate to high rainfall (Myers 1927, 1929; Laing 1937, Griffiths 1964, 1966; Fischer 1970, 1971; Wharton 1986).  A common species in Europe, A. manducator Panzer, was successfully established in Australia and New Zealand (Miller 1927, Newman 1928, Morgan 1929, Holdaway & Evans 1930, Holdaway & Smith 1932.)  The nearctic Alysia ridibunda Say has been found active on Calliphoridae (Lindquist 1932, 1940; Marsh 1968).  Species of Alysia may be distinguished from those of Aphaereta by their larger size (ca. 2X as long), their shining black body color and the uneven thickness of antennal funicular segments.  They are solitary endophagous larval parasitoids of muscoid flies. 

 

Additional key references are Altson (1920), Caudri (1941), Likovský (1965, 1973), Burgess & Wingo (1968), Vinogradova & Zinovjeva (1972), Zinovjeva (1976, 1978, 1981, 1985, 1987, 1988), Chernoguz (1984, 1986), Chernoguz & Reznik (1987), Chernoz & Vaghina (1987), and Chernoguz et al. (1987).

 

3.  Aphaereta spp. (Hymenoptera: Braconidae)

 

These parasitoids are found primarily in climates with substantial rainfall.  Aphaereta pallipes (Say) is most frequently encountered in the Holarctic (McComb 1958, Salkeld 1959, House & Barlow 1961, Griffiths 1964, 1966; Lange 1964, Lange & Bronskill 1964, Houser 1966, Houser & Wingo 1967a, Marsh 1969, Fischer 1970, 1971; Garry & Wingo 1971, Figg et al 1983a,b, Whistlecraft et al. 1984, Rueda & Axtell 1985b).  Aphaereta aotea Hughes & Woolcock is found in Australia (Hughes & Woolcock 1976, 1978, Hughes et al 1974).  Species of Aphaereta may be readily distinguished from those of Alysia by their smaller size (ca. 1/2 as long), their reddish-brown body color and uniform thickness of funicular antennal segments.  They are solitary endophagous larval parasitoids of muscoid flies.  Key references are Fischer (1966), Zinovjeva (1974), and Gherasin & Lacatusu (1977).

 

Family Ichneumonidae (Hymenoptera: Ichneumonoidea)

 

Forewing venation is complete, not reduced, with at least one complete cell present.  Stigma and costal cell are absent.  There are 2 recurrent veins usually present (Fig. 45a-A & B).  Ventral abdominal segments are soft and membranous, with a median fold.   Two important genera are Phygadeuon and Stilpnus.  Key references are Thompson (1944), Salt (1952), Townes & Townes (1966, 1973), Riek (1970) Richards (1977), Krombein et al. (1979), Borror et al. (1981), Subba-Rao & Hayat (1985), Pisicà & Fabritius (1986), Blanchot (1988, 1991a,b), Gauld & Bolton (1988), Rollard (1988) and Goulet & Huber (1993).

 

4.  Phygadeuon spp. (Hymenoptera: Ichneumonidae)

 

This genus and Stilpnus can be distinguished from other parasitoids noted herein by their complete wing venation.  Both the fore wings and hind wings have venation closed to form several cells (Fig. 46a-A).  This genus is unique here in having antennae of 22 segments with the first two segments short and the third segment longer than the first two combined.  The antennae are inserted in the middle of the face between the eyes.  The species are solitary internal larval parasitoids most often found in humid higher Holarctic latitudes (Legner 1966, Legner & Olton 1968, Legner et al. 1976).  Additional key references are Monteith (1956), Horstmann (1967, 1972, 1975, 1986), Müller (1971), Frilli (1973), Plattner (1975, 1979), Plattner & Naton (1975), and Naton (1983).

 

5.  Stilpnus spp. (Hymenoptera: Ichneumonidae)

 

The wing venation is complete as in Phygadeuon, there being complete cells in both the fore wings and hind wings.  Antennae have 16 segments with the first two segments short, the first being shorter than the next two together.  The color is shining metallic black.  They are solitary endophagous larval parasitoids, apparently restricted to the genus Fannia in accumulated organic wastes (Legner & Olton 1971, Legner et al. 1976, Loomis et al. 1968).  They vary greatly in size with males being about 1/2 that of females.  A common species is S. anthomyiidiperda (Viereck).

 

Family Chalcididae (Hymenoptera: Chalcidoidea)

 

Hind femora are greatly enlarged, ventrally toothed (either a few large or many small teeth) (Fig. 18a-A).  The prepectus is reduced or fused, not triangular (Fig. 18a-B).  The frons is projected into 2 horns (around antennae) when viewed dorsally.  They are solitary parasitoids which attack mature host larvae or young pupae.  Their general appearance is shown in Fig. 18a-E.  Two important genera are Brachymeria and Dirhinus.  Key references are Ashmead (1899), Schmiederknecht (1907, 1909), Froggatt (1916), Thompson (1944), Schmitz (1946), Boucek (1956), Nikolskaya (1960), Riek (1970),  Richards (1977), Krombein et al. (1979), Subba-Rao & Hayat (1985), Fabritius & Andriescu (1987), Xue (1988a), Xue et al. (1987c), Gauld & Bolton (1988), and Goulet & Huber (1993).

 

Genus Dirhinus Dalman, 1818 (Hymenoptera: Chalcididae)

 

(Dirrhinus Dalman, 1923; Eniaca Kirby, 1883; Dirrhinoidea Girault, 1912; Pareniaca Crawford, 1913; Eniacella Girault, 1913; Eniacomorpha Girault, 1915; Dirhinoides Masi, 1947).

 

Parasitoids in this genus possess a pair of horns on the head which in some species bear a tooth (Figs 42a-B & D).  They have an elongated body which is somwhat depressed dorsally (Fig. 42a-E).  The mandibles are long and narrow, almost straight.  The genae are very large and punctured.  They parasitize various brachycerous Diptera, seeking out full-grown larvae or pupae in the soil.  The key reference is Boucek & Narendran (1981).

 

6.  Dirhinus anthracia Walker, 1846 (Hymenoptera: Chalcididae)

 

(Dirrhinus ruficornis Cameron, 1905; Eniacella rufricornis Girault, 1913; Eniacella bicornuticeps Girault, 1915; Dirhinus sarcophagae Froggatt, 1919; Dirhinus frequens Masi, 1933; Dirhinus intermedius Mani & Dubey, 1974).

 

Female head densely punctate, in dorsal view with eyes longer than temples, these converging, slightly convex (Fig. 42a-A).  Antennae of female reddish with yellow or white hairs.  Each horn has a deep external apical notch with subparallel ridged (= carinate) sides, the inner ridge being laminate but almost regular (Fig. 42a-B & C).  Horn in middle is usually distinctly broader than the gap between antennal sockets, in dorsal view.  The ocellar area is somewhat elevated; in lateral view the head is less than 0.6 as wide as long, in dorsal outline from the horn edge to occiput it is strongly convex.  However, the vertex appears as almost one plane.  Length of gena (from eye to submandibular corner) is obviously greater than the short diameter of eye.  Labrum bears scattered hairs.  Antennal scape is thickened toward its base.  Pronotum bears small lateral patches of thicker hairs, and the punctuation in center is dense.  Abdominal petiole is transverse, area of 4 dorsal ridges about 2X as broad as long, anteriorly frequently margined.  In male the horns dorsally in middle are narrower than the gap.   Size varies with host size.  The general appearance is as in Fig. 42a-D & E.  There is a broad host range including Muscidae, Sarcophagidae, Calliphoridae, Tephritidae and Tachinidae (Boucek & Narendran (1981).  Original distribution India, Burma, Sri Lanka.  Additional key references are Walker (1846), Froggatt (1919, 1921), Lever (1938), Masi (1947), Dresner (1954), Boucek (1956), and Mani et al. (1974).

 

7.  Dirhinus bakeri (Crawford, 1914)  (Hymenoptera: Chalcididae)

 

(Pareniaca bakeri Crawford, 1914; Pareniaca trichophthalma Masi, 1927).

 

This is a small species, 2.5-4.1 mm long, with antennae usually black, but at times with pedicel and flagellar base in female reddish.  Facial edge of each horn with a distinct additional tooth (viewed laterally) (Fig. 40b-A & B).  Tip of horn reaches much farther from eye than frontal tooth (Fig. 40b-A).  Fore and mid femorae and tibiae usually are black, and wings usually whitish.  Abdominal petiole is rather closely joined with abdomen.  Gena of both sexes is longer than the maximum diameter of eye (Fig. 40b-A).  Hosts include Musca domestica and species of Stratiomyiidae and Tachinidae (Boucek & Narendran 1981).  Original distribution India, Sri Lanka, Malaysia, Philippines, Japan.  Additional key references are Masi (1947), Habu (1960), Baltazar (1966), and Geetha-Bai & Sankaran (1977).

 

8.  Dirhinus banksi Rohwer, 1923 (Hymenoptera: Chalcididae)

 

Female body black including most of legs and flagella.  Scapes, pedicels, tarsi and joints of legs are testaceous.  Thorax is much flattened dorsally, especially scutellum completely flat and with broad smooth area separated by single row of punctuations from hind margin (Fig. 42b-A & B). Lateral head at least 1.5X as long as wide, with facial outline convex, and receding near horn tips.  In dorsal view head width is ca. 1.67X the minimum distance between eyes, horns appearing wide and nearly with parallel sides in basal half (Fig. 42b-C).  Flagellum plus pedicel 1.6-1.7 X as long as head is wide in lateral view, and clava a bit shorter than 3 preceding segments combined.  Pronotum medially without smooth longitudinal strip (Fig. 42b-B).  Mesosternum with impunctate areas behind the fore coxae frequently are well delimited from the posterior punctate part and not extending quite 1/2 to mid coxae.  Petiole with area of 4 ridges slightly to moderately transverse.   In male antennae are paler or darker yellow, slightly less clavate than in female.  Eyes are relatively small, in lateral view the height of eye is only about 1.2X of the height of horn projecting above it.  Length of female 2.5-3.7 mm., male 2.6-3.1 mm.  The only known host is Lucilia sp. (Boucek & Narendran 1981).  Original distribution India, Sri Lanka, Thailand, Cambodia, Malaysia.   Additional key references are Rohwer (1923), Masi (1947), and Habu (1976).

 

9.  Dirhinus himalayanus Westwood, 1836 (Hymenoptera: Chalcididae)

 

(Dirrhinus crythroceras Cameron, 1906; Dirhinus luzonensis Rohwer, 1923; Dirhinus luciliae Rohwer, 1923; Dirhinus pachycerus Masi, 1927; Dirhinus vlasovi Nikolskaya, 1952; Dirhinoides mathuri Mani & Dubey, 1972).

 

Body is very black, with sparse coarse punctuation on thorax; wings are clear without distinct hairs in female.  The apex of each horn viewed dorsally is almost rounded, without a notch (Fig. 41a-A & B).  Distal 1/2 of hind tibia with another shallow groove outside tarsal sulcus, the groove being outlined by some extra ridges.  Clava of female symmetrical, with a broad conical apex (Fig. 41a-C).  Hind tibia have a  conspicuous external ridge.   Length of female 2.4-4.9 mm. (the longest of all species noted here).  Host range is broad in Diptera in carcasses and excrement (Bouek & Narendran 1981).  Original distribution Arabia, India, Malaysia, Sumatra, Japan, Philippines & Hawaii.   Additional key references are Cameron (1906), Rohwer (1923), Ferrière (1935), Roy & Siddons (1939), Roy et al. (1940, 1950), Stearn (1943), Nikolskaya (1952, 1960), Dresner (1954), Habu (1960), Mani & Dubey (1972), and Sankaran (1977, 1985),  Srinivasan & Panicker (1988), Xue (1989), Geetha-Bai (1990), Geetha-Bai & Sankaran (1977, 1985),

 

Family Encyrtidae (Hymenoptera: Chalcidoidea)

 

Fore wing venation greatly reduced, with a single vein along margin and very short spur.  Fore wing with uncus well separated from postmarginal vein.  Marginal vein indistinct, somewhat punctiform.  All tarsi with 4 segments.  Abdominal petiole at most 1-segmented.  Eggs are dumbbell-shaped.  Endophagous, gregarious larval parasitoids.  The principal genus is Tachinaephagus.  Key references are Thompson (1944), Wilson & Woolcock (1960), Legner & Bay (1965a), Riek (1970), Ho et al. (1974), Richards (1977), Trjapitzin (1978), Krombein et al. (1979), Prinsloo & Annecke (1979), Noyes (1980), Noyes & Hayat (1984), Borror et al. (1981), Subba-Rao & Hayat (1985), Gauld & Bolton (1988) and Goulet & Huber (1993).

 

Genus Tachinaephagus Ashmead, 1904 (Hymenoptera: Encyrtidae)

 

(Tachinaephagus Girault, 1917; Australencyrtus Johnston & Tiegs, 1921; Australomalotylus Risbec, 1956)

 

There are three species known to parasitize synanthropic Diptera in decomposing organic wastes, although the whole genus is exclusively parasitic on the larvae or pupae of many Diptera.  Vertex of head is almost 1/3rd to 1/2 the width of head, frons is broad.  Eyes are hairy with long dense setae.  Antennae inserted just below the lower eye level with toruli (= sockets) widely separated, scape cylindrical, the funicle segments 1-3X longer than broad (quadrate) (as in Fig. 38a-A).  Mandibles have three sharp teeth.  Scutellum long, shining but bears long setae which arise from microscopic punctures.  Legs are hairy.  Basal area of fore wing is evenly setose (Fig. 39a-A).  Abdomen is either slightly shorter or longer than thorax and flat above in dead specimens.  Digitus of male bears 3 teeth.  The eggs are encyrtiform (dumbbell-shaped).  They are endophagous gregarious larval parasitoids.  Size varies with host size and number of individuals developing on one host.  Key references are Tachikawa (1963), Olton & Legner (1975) and Subba-Rao (1978).  Other references are Ashmead (1904a,b), Girault & Sanders (1909, 1910a), Girault (1917), Dodd (1921), Froggatt (1921), Johnston & Tiegs (1921), Hardy (1924), Gourlay (1930a,b), Newman & Andrewartha (1930), Ferrière (1933), Gahan (1938), Risbec (1956), Ghesquière (1960), Olton (1971), and Subba-Rao (1972, 1976).

 

10.  Tachinaephagus javensis Subba-Rao, 1978 (Hymenoptera: Encyrtidae)

 

Body is uniformly dark brown.  Legs, coxae and antennal scape are testaceous yellow, funicle and club brown; venation and discal cilia of forewing brown.  Head vertex is broader than 1/2 the head width; ocelli are large in an equilateral triangle (Fig. 38a-C), the posterior part separated from the ocular border by about their own diameter.  Antennal scape is cylindrical, its pedicel only slightly longer than the 1st funicle segment, 2nd and 3rd segments shorter than 1st, 4-6 about equal but shorter than 3rd; club apically rounded, the joints not well separated (Fig. 38a-A); thorax is moderately convex; scutellum rugose; Fore wings with caudal cell moderately wide, not parallel with submarginal vein (Fig. 38a-B); marginal fringe short; abdomen longer than thorax, tergites shining and smooth.  Males have not been found.  Hosts include Haematobia and species of Musca (Subba-Rao 1978).  Original distribution Indonesia.  The key reference is Subba-Rao (1978).

 

11.  Tachinaephagus stomoxicida Subba-Rao, 1978 (Hymenoptera: Encyrtidae)

 

Body is almost black; head vertex with faint metallic green reflections; coxae are dark brown, rest of legs brown with somewhat darker tarsi; scape brown, funicle and club dark brown.  Fore wings are slightly hairy (Fig. 39a-A); head vertex more than 1/2 width of head; ocelli in an equilateral triangle, the posterior pair separated from ocular borders by a little more than their own diameter.  Antennal socket is broad, semicircular and shallow.  Malar sulcus impressed deeply only basally.  Antennal scape cylindrical, slightly dilated above, funicle segments well separated, the club segments deep and club apex angular (Fig. 39a-C).  Thorax is only slightly convex with scutellum almost flat; mesoscutum scaly, scutellum smooth and shining except for tiny pits bearing long black setae.  Fore wings are long and narrow, costal cell very narrow and parallel with submarginal vein (Fig. 39a-A & B).  Discal ciliation is coarse and dense.  Abdomen is slightly longer than thorax, almost quadrate, tergites smooth and shining.  Males resemble female except for their antennae.  The only known host is Stomoxys nigra (Subba-Rao 1978).  Original distribution in Uganda but established in Mauritius.  Key references are Subba-Rao (1978) and Greathead (1986).

 

12.  Tachinaephagus zealandicus Ashmead, 1904 (Hymenoptera: Encyrtidae)

 

(Tachinaephagus australiensis Girault, 1917; Stenosterys fulvoventralis Dodd, 1921; Australencyrtus giraulti Johnston & Tiegs, 1921; Australomalotylus rageaui Risbec, 1956).

 

The wing venation is greatly reduced with a single vein along the margin and a very short spur, the stigmal vein, near its center (Fig. 39b-B).  Costal cell of forewing wide, its border not parallel with submarginal vein (Fig. 39b-B & C).  The antennae are located in the middle of the face between the eyes.  They are of less than 14 segments with the first segment elongated, longer than the next two combined.  As in Muscidifurax, the pronotal disc is finely reticulate and almost imperceptibly punctured.  The color is shining black with the underside of the thorax and the legs testaceous (Fig. 39b-D).  Hosts include Calliphoridae, Fannia canicularis (L.) and Musca domestica L.  Original distribution Australasia.  Key references are Tachikawa (1963), Olton (1971), Olton & Legner (1974, 1975) and Subba-Rao (1978).  Other references are Ashmead (1904), Froggatt (1921), Johnston & Tiegs (1922), Hardy (1924), Gourlay (1930a,b), Newman & Andrewartha (1930), Ferrière (1933), Gahan (1938, Risbec (1956), and Legner & Olton (1968b).

 

Family Pteromalidae (Hymenoptera: Chalcidoidea)

 

Forewing marginal vein is less than twice as long as stigmal (Fig. 19b-A).  Antennae have 6 or fewer funicle segments.  Hind tarsus with at least 5 segments.  Abdominal petiole often conspicuous and with dorsal carinae.  They are ectophagous, pupal parasitoids inside the host puparium.  Important genera are Muscidifurax, Nasonia, Pachycrepoideus, Spalangia, Sphegigaster, and Urolepis.  Key references are Latrielle (1805), Dalman (1820), Walker (1836, 1839), Curtis (1839), Förster (1841, 1956), Ashmead (1896b), Dalla-Torre (1898), Perkins (1910), Waterston (1915), Fortsetzung (1916), Girault (1916, 1921), Parker (1924), Parker & Thompson (1928), Ceballos (1941), Thompson (1944), Delucchi (1955), Boucek (1963), Peck (1963), Baltazar (1966) DeSantis (1967, 1979, 1980), Graham (1969), Riek (1970), Abraham (1975, 1978a), Wylie (1976b), Richards (1977), Burks (1979), Gordh et al. (1979), Krombein et al. (1979), Barlin & Vinson (1981), Borror et al. (1981), Yoshimoto (1984), Subba-Rao & Hayat (1985), Gauld & Bolton (1988), Hoebeke & Rutz (1988), Delvare & Aberlenc (1989), Grissell & Schauff (1990), Goulet & Huber (1993).

 

Genus Muscidifurax Girault & Sanders, 1910 (Hymenoptera: Pteromalidae)

 

Wing venation is incomplete and the marginal vein is about twice as long as the stigmal vein (Fig. 23a-A).  Antennal insertions are in the middle of the face between the eyes.  The first antennal segment is longer than the next two combined, and there are less than 14 antennal segments.  Females have one ring segment and 7 funicular segments (Fig. 23a-B), males have 2 and 6, respectively.  The disc of the pronotum and the head are finely reticulate, without coarse punctures.  The several species are very similar in appearance but have good behavioral characters distinguishing them (Legner 1969a,b; Legner et al. 1976, Kogan & Legner 1970), and they are electrophoretically distinct (Kawooya 1983).  Females are black; males black with translucent testaceous spots on the first, second and third ventral abdominal segments.  The eggs are hymenopteriform, covered with small tubercles that distinguishes them from those of Spalangia (Gerling 1967) and with size differences for some species.  The species may be either solitary or gregarious.  The average mass of solitary species of this genus is relatively fixed, as host size does not appreciably affect them (Legner 1969a).  They are ectophagous pupal parasitoids.  The key reference is Kogan & Legner (1970).  Van den Assem & Povel (1973) discussed courtship behavior patterns that are specific.  Markwick (1974) and Markwick et al. 1989 gave biological characteristics that distinguish M. raptor and M. zaraptor and these species from Spalangia endius. 

 

Other references referring to distribution, identity, biology and genetics of species of this genus are Frison (1927), Anonymous (1938), Nikolskaya (1952), Dresner (1954), Legner (1967b, 1969a,b; 1972, 1987a,b,c,d,e; 1988a, 1988b, 1988c,d; 1989a, 1990, 1991a, 1991b, 1993), Legner & Dietrick (1974), Legner & Gerling (1967), Legner et al. (1967), McCoy (1967), Wylie (1967, 1971a,b, 1972b), Berry & Speicher (1972), Broadbent (1972), Kotschetova & Tjutjunkova (1973), Ables & Shepard (1974), Kawooya (1983), van den Assem (1985), Propp (1986), Mandeville & Mulles (1990b,c), Mandeville et al. (1990), Mann et al. (1990b), Wilhoit et al. (1991a).

 

13.  Muscidifurax raptor Girault & Sanders, 1910 (Hymenoptera: Pteromalidae) (PHOTO)

 

The fringe of setae (or their sockets) is well developed on the posterio-apical margin of the fore wing  (Fig. 23a-A).  The stigma forms an abrupt enlargement at the end of the stigmal vein, usually subquadrangular and distally acuminate where a hair is often encountered (Fig. 27a-A).  The uncus is directed distally.  The frontal grooves are parallel (Fig. 27a-B).  The median area of the propodeum of female is closed behind (Fig 27a-C).  The male digitus is subtrapezoidal, broadest distally and usually with 3 apical processes (Fig. 27a-D).  Hind legs with one tibial spur (Fig. 27a-E).  Hind leg with tarsal claw bearing less than 11 setae (Fig. 27a-F).  Length of female is 2.33 mm, male 1.73 mm.  The species is biparental and solitary.  An almost cosmopolitan species which has not been collected in the Oriental region nor most of Asia.  Key references are Girault & Sanders (1910a,c), McCoy (1963, 1967), Legner (1969a,b; 1976a), Legner & Gerling (1967), Kogan & Legner (1970), Morgan & Patterson (1975a), Fabritius (1978, 1981b,c,d; 1983a, 1984, 1986c) and Kawooya (1983). 

 

Additional references are DeBach (1943), Sharma (1967, 1971), Burton & Turner (1968), Shibles (1969), Wylie (1970), Victorov & Azizov (1972), Markwick (1974), Tingle & Mitchell (1975), Podoler & Mendel (1977), Fabritius (1979b, 1980a,b), Morgan et al. (1979b), Capehart et al. (1981), Coch (1981), Klunker (1981, 1982), Merritt et al. (1981), Rutz & Axtell (1981), Shepard & Kissam (1981), Propp & Morgan (1983b, 1984a, 1985a), and Geden et al. (1992a,c).

 

14.  Muscidifurax raptorellus Kogan & Legner, 1970 (Hymenoptera: Pteromalidae)

 

The fringe of setae (or their sockets) is absent from the posterio-apical margin of forewing, as in M. zaraptor (Fig. 26b-A).  The stigma may or may not be elongated, sometimes being roundly clubbed.  The uncus is directed towards of the apex of the wing.  The pedicel of female antennae is not slender proximally (Fig. 26b-B), and frontal grooves are usually convergent (Fig. 26b-C).  The median area of the propodeum of female is usually open behind with the lateral and median plicae not fused in the middle (Fig. 26b-D).  The male digitus is subtrapezoidal, broader distally and usually with only 3 distal processes (Fig. 26b-E).  Length of female is 2.11 mm, male 1.82 mm. The Peru race averages female 2.33 mm, male 1.72 mm although they may assume the smaller size of the Chilean race when deprived of host feeding.  This South American species occurs as several races demonstrating various degrees of gregarious development and number of eggs deposited into a single host puparium.  The Peruvian race is usually solitary in development, but seasonal variations in larval cannibalism vary.  The Chilean race has shown the greatest degree of gregarious development (> 80%) and is quite thigmotactic.  All known races are biparental.  Key references are Kogan & Legner (1970), Kawooya (1983), and Legner (1987b,e; 1988a,b,c; 1989a,b,c,d; 1990, 1991a,b; 1993).

 

          Superparasitism (= insertion of more than one parasitoid egg per host) occurs in both the Peruvian and Chilean race, and subsequent cannibalism by hatched larvae always follows.  The Peruvian race deposits a lower number of eggs per host than the Chilean race but cannibalism intensity is higher.  Therefore, it is difficult to determine the exact number of eggs initially deposited by both species.  The number of adult parasitoids that survive is always less in the Peruvian race and usually averages about one.  On the contrary, more adult survivors usually occur in the Chilean race, averaging about seven at a host density of 20 per 24 hrs.  A standardization of host density, size, age and duration of exposure to parasitization is essential in experiments as they influence the number of eggs deposited and the rate of cannibalism.

 

 

15.  Muscidifurax raptoroides Kogan & Legner, 1970 (Hymenoptera: Pteromalidae)

 

The fringe of setae (or their sockets) is well developed on the posterio-apical margin of forewing.  The stigma is formed as a gradual dilation of the tip of the stigmal vein, and the uncus is directed distally (Fig. 28b-B).  The frontal grooves are convergent.  The male digitus is subrectangular and usually with 4 apical processes (Fig. 28b-C).  Spiracle of the female propodeum is not remote from the lateral plica, i.e., the spiracular ridge is shorter than the largest diameter of the spiracle (Fig. 28b-A).  Length of female 2.31 mm, male 1.78 mm.  This species is biparental and solitary.  It was originally known from Costa Rica and southern Mexico.  Key references are Kogan & Legner (1970), and Legner et al. (1976).

 

16.  Muscidifurax uniraptor Kogan & Legner, 1970 (Hymenoptera: Pteromalidae)

 

The fringe of setae (or their sockets) is well developed on the posterio-apical margin of forewing.  The stigma is formed as a gradual dilation of the tip of the stigmal vein, and the uncus is directed distally (Fig. 28a-B).  The frontal grooves are convergent.  When male are present, the digitus is subrectangular and usually with 4 distal processes (Fig. 28a-C).  The spiracle of the female propodeum is remote from the lateral plica, i.e., the spiracular ridge is as long as the longest diameter of the spiracle (Fig. 28a-A).  Length of female 2.15 mm, male 1.68 mm, the occasional male are the smallest in this genus.  This species is uniparental and solitary in development.  It was originally known only from Puerto Rico where it occurs with the biparental M. raptor.  Key references are Kogan & Legner (1970), Legner (1985a,b; 1987a,d; 1988), and Kawooya (1983).

 

17.  Muscidifurax zaraptor Kogan & Legner, 1970 (Hymenoptera: Pteromalidae)

 

The fringe of setae (or their sockets) is absent from the posterio-apical margin of the forewing as in M. raptorellus.  The stigma is small, elongated, suboval, often acuminate at the internal angle where a hair is implanted (Fig. 26a-A).  The uncus is usually directed toward the anterior margin of the wing.  The antennal pedicel of female is obviously slender proximally (Fig. 26a-B), and frontal grooves are usually parallel (Fig. 21a-C).  The median area of the propodeum in female is closed behind by the fusion of the lateral and median plicae (Fig. 26a-D).  The male digitus is subrectangular with 4-5 apical processes (Fig. 26a-E).  Hind leg with tarsal claw bearing more than 11 setae (Fig. 26a-G).  Length of female is 2.,84 mm, male 2.18 mm.  This species is biparental, solitary and occurs naturally in the western Nearctic.  Key references are Kogan & Legner (1970), Wylie (1971b, 1979), Legner 1977, 1979a,b,c) and Kawooya (1983).  <PHOTO>

 

Additional references are Coats (1976), Petersen & Meyer (1983a,b), Petersen & Matthews (1984), Petersen & Pawson (1988a,b; 1991), Petersen et al. (1986, 1992), Pawson et al. 1987, Mandeville (1988), Mandeville & Mullens (1990a), and Mandeville et al. (1988),

 

Genus Nasonia Ashmead (Hymenoptera: Pteromalidae)

 

18.  Nasonia vitripennis (Walker, 1836) (Hymenoptera: Pteromalidae)

 

(Nasonia Ashmead, 1904; Mormoniella Ashmead, 1904; Pteromalus vitripennis Walker, 1836; Pteromalus muscarum Hartig, 1838; Pteromalus abnormis Boheman, 1856; Dicyclus pallinervosus Walker, 1872; Stictonotus insuetus Walker, 1872; Mormoniella brevicornis Ashmead, 1904; Nasonia brevicornis Ashmead, 1904; Platymesopus macellariae Brèthes, 1913di).

 

Head reticulate, wider than broad.  Upper margin of antennal sockets is slightly above level of the lower margin of compound eye.  A distinct dorsocephalic ridge exists.  The malar sulcus is not as pronounced as in Spalangia endius (see Fig. 37a-D).  Antennae are stout with two ring segments and 6 funicular segments (Fig. 24a-B); the pedicel is ca. 2 1/2X longer than first funicular segment.  The pronotal collar bears an anterior ridge and is strongly reticulate ventrolaterally.  The mesosternum and scutellum is strongly reticulate, the frenal groove is distinct (Fig. 24a-A).  Propodeum has a median plica which lacks lateral branches.  The prosternum bears a median groove.  Fore wing with marginal veins of uniform width and as wide as post marginal vein.  Middle leg bears one tibial spur and dense bristles over its entire length.  The spur is ca. 1/3rd as long as first tarsomere.  The hind leg bears one tibial spur with a bristle-like structure (Fig. 24a-C).  Tergite 9 of abdomen with a pair of slightly projecting sensory structures, each with 5 long setae.  Length of female 1.10-1.90 mm, male 1.00-1.50 mm.  Hosts are primarily blowflies in animal carcasses, although occasionally parasitization occurs in muscoid Diptera found in manure and accumulated organic wastes.  The species is cosmopolitan.  General appearance (Fig. 24a-E).  Key references are Rueda & Axtell (1985b). 

 

Other key references to this most extensively studied (and undoubtedly the least beneficial) fly parasitoid include Froggatt (1914a,b; 1918), Froggatt & McCarthy (1914), Robaud (1917), Altson (1920), Miller (1922, 1972, Miller & Tsao 1974), Tiegs (1922), Hardy (1924, 1925), Séguy (1929), W. Davies (1930), Dautert Willimzik (1931), Cousín (1930, 1933), Evans (1933), Smirnov (1934), Smirnov & Kuzina (1933), Smirnov & Vladimirow (1934), Vladimirova & Smirnov (1934), Robaud & Colias-Belcour (1935), Fukuda (1939), Jacobi (1939), DeBach (1940), DeBach & Smith (1941a,b; 1947), DeBach (1943), van der Merwe (1943), Geršenson (1946), Moursi (1946), Ray (1948, 1953, 1955a,b; 1957, 1958), Kayhardt & Whiting (1949), Kayhardt (1956), Ray & Whiting (1954), Ray et al. (1954), Ullyett (1949, 1950), Ullyett & DeVries (1940), Friedler & Ray (1951), Whiting (1951, 1954a,b; 1955a,b,c,d; 1956a,b; 1957, 1958, 1960, 1965, 1967), Wylie (1958, 1962, 1963, 1964a,b; 1965a,b,c; 1966a,b,c; 1970, 1973b, 1976a), Whiting & Busa (1959), Edwards (1954a,b; 1955a,b; 1961), Saul (1954, 1955, 1957, 1960, 1981), Saul & Kayhart (1956), Barrass (1961), Saul et al. (1965, 1967), Cutler (1955), Firschel & Wolsky (1956), Fluke (1957), Rohner & Wolsky (1957), Varley & Edwards (1957), Pennypacker (1958), Schneiderman & Horwitz (1958), Schneiderman et al. (1956a,b,c), Ohgushi (1959a,b; 1960, 1961), Ohgushi & Kato (1959), Barrass (1960a,b; 1961, 1962, 1965, 1969, 1976a,b), King (1961a,b; 1962a,b,c,d; 1963), King & Hopkins (1963), King & Rafai (1970), King & Ratcliffe (1969), King & Richards (1968), King et al. (1968, 1969), Ankersmit et al. (1962), Nagel (1962), Nagel & Pimentel (1963, 1964), Saunders (1962, 1965a,b,c; 1966a,b; 1967, 1968, 1973, 1974a,b; 1978, 1981), Saunders & Sutton (1969), Saunders et al. (1970), Mortimer & van Borstel (1963), Beard (1964b, 1972), Hopkins & King (1964), Madden (1964), Madden & Pimentel (1965), Hair & Turner (1965), Velthuis et al. (1965), Pimentel (1966, 1984), Pimentel & Al-Hafidh (1963, 1965), Pimentel et al. (1963, 1978), Azab et al. (1967a,b), Chabora (1967, 1970a,b,c; 1972, 1980), Chabora & Chabora (1971), Chabora & Pimentel (1966, 1970), Clark & Cole (1967), Clark & Kidwell (1967), Legner (1967b,c), Legner & Gerling (1967), Ratcliffe & King (1967, 1968, 1969a,b,c; 1970), Takahashi & Pimentel (1967), Walker (1967), Walker & Pimentel (1966), Rabinovich (1969), Richards (1969), Slifer (1969), Smith (1969), Smith & Pimentel (1969), Sanger & King (1971), Barash & Ryder (1972), Copland & King (1972), Holmes (1972, 1974a,b) Rafai & King (1972), Grant et al. (1974, 1980), Olson & Pimentel (1974), van den Assem (1974), van den Assem et al. (1981), Cassida (1975), I. Davies (1975), I. Davies & King (1975), Sagan & Fashing (1977), van den Assem (1977, 1985), van den Assem & Feuth-DeBruijn (1977), van den Assem & Putters (1980), van den Assem & Vernel (1979), van den Assem & Visser (1976), van den Assem et al. (1980a,b,; 1981, 1984), White & Grant (1977), Abraham (1978b, 1985), Abraham & König (1977),Best (1978), Cornell & Pimentel (1978), DeLoof et al. (1979a,b), Fashing & Sagan (1979), Smith & Cornell (1979), Zareh et al. (1980), Bull (1982), Skinner (1982, 1983, 1985), Jachmann (1983), Werren (1984), Werren et al. (1981, 1986), Wibel et al. (1984), Xue (1984a, 1986d, 1987b, 1988c), Xue et al. (1987b, 1988), Huger et al. (1985), Parker & Orzack (1985), Orzack et al. (1986), Preutu (1986), Schmidt (1986), Fried (1987), Fried & Pimentel (1986, 1990), Jones & Turner (1987), Omar (1987a,b), and Darling & Werren (1990).

 

Genus Pachycrepoideus Rondani, 1875 (Hymenoptera: Pteromalidae)

 

19.  Pachycrepoideus vindemiae Rondani, 1875 (Hymenoptera: Pteromalidae) (Encyrtus vindemiae Rondani, 1975).

 

This species is similar in appearance to species of Muscidifurax but is distinguished by the short marginal vein which is not longer than the stigmal vein.  The wing venation is incomplete.  The antennae arise from the middle of the face between the eyes with the first segment longer than the next two combined.  Antennae have 3 anelli (Fig. 22a-D) and less than 14 funicular segments.  The head and the disc of the pronotum are finely reticulate without obvious punctures.   The pronotal collar is without a transverse ridge on its anterior margin (Fig. 22a-A).  Propodeum without median folds (Fig. 22a-B).  Mesosternum has a pair of anterior tubercles (Fig. 22a-C).  Digitus of male with only 2 apical processes (Fig. 22a-E).  The species is ectophagous, usually solitary on the pupa within the puparium.  It is cosmopolitan in distribution (Girault & Sanders 1910, Jaynes 1930, Crandall 1939, Nostvik 1954, Steve 1959, Legner et al. 1967, Legner & Olton 1968, van den Assem 1974, Rueda & Axtell 1985b). 

 

Additional key references are Pickens et al. (1975), Pickens & Miller (1978),  Morgan (1980b), Morgan & Patterson (1975a), Morgan et al. (1978b), Pickens (1981), van Alphen & Thunnissen (1983), Thompson (1981a,b), Thompson et al. (1983), and Panicker & Srinivasan (1986).

 

Genus Spalangia Latreille, 1805 (Hymenoptera: Pteromalidae) (Prospalangia Brèthes, 1915)

 

Species of Spalangia have incomplete wing venation, with the marginal vein about 10X as long as the stigmal vein (Fig. 21b-A).  The antennae are situated at the front margin of the head (Fig. 21b-B).  The first antennal segment is longer than the next two combined (Fig. 21b-C).  There are less than 14 funicular segments.  The pronotal disc is coarsely punctured with polished interspaces (e.g., Figs. 29b-A & 33a-C).  The eggs are hymenopteriform and smooth with the size being variable according to the species (Gerling 1967).  Host size does not much affect the size of solitary species of this genus (Legner 1969a,c).  They are usually solitary ectophagous pupal parasitoids.  The 9 species treated here are easily distinguished among themselves.  Key references are Boucek (1963, 1965, 1988).  Markwick (1974) gives biological characteristics of S. endius and distinguishes this species from Muscidifurax. 

 

Other references include Cameron (1881), Ashmead (1896a), Pinkus (1913), Richardson (1913), Brethes (1915), Fullaway (1915), Vandenburg (1928, 1931), Simmonds (1929a,b), Graham (1932), Handschin (1934), McCoy (1963), Legner (1965a, 1967a,b), Gerling & Legner (1968), Burks (1969, 1979), Mourier (1971a,b), Azizov (1972), Kotschetova & Azizov (1972), Wylie (1972a), Chu (1984), Rueda & Axtell (1985b), Propp (1986), Mandeville (1988), Mandeville & Mullens (1990c), Mandeville et al. (1990), Huang (1990), Mann et al. (1990a,b), Omar et al. (1991b).

 

20.  Spalangia cameroni Perkins, 1910 (Hymenoptera: Pteromalidae)

 

(Spalangia melanogastra Masi, 1940; Spalangia atherigonae Risbec, 1951).

 

The disc of the head between the eyes is sparsely punctured with the punctures mostly separated by more than their diameters.  The disc of the pronotum has an isolated wavy crossline consisting of large closely spaced punctures near to and parallel to the posterior margin (Fig. 33a-C).  The pronotal collar is rounded anteriorly, without a distinct ridge (Fig. 33a-C, D & E).  The anterio-lateral surface of the pronotum is rugose or crowdedly rugosely punctured.  The length of the gena is longer than that of the eye (Fig 33a-A).  The ratio of abdominal petiole length to the narrowest width is 1.8 in femaleand 2.5 in male  (Fig. 33a-F & G), and there are no lateral hairs present.  The length of the extended body is 2.5-3.3 mm in femaleand 2.4-3.0 mm in male.  A cosmopolitan species.  Key references are Perkins (1910), Simmonds (1929b), Masi (1940), Risbec (1951), Boucek (1963), Legner & Brydon (1966), Legner & Gerling (1967), Legner & Olton (1968, 1979), Legner (1969a), Legner & Greathead (1969), Legner et al. (1967, 1976, 1990a,b), Gerling & Legner (1968), Wylie (1972a), Markwick (1974), Rutz & Axtell (1980a), Moon et al. (1982), Scott et al. (1988), Morgan et al. (1989), and Maini & Bellini (1990a,b).

 

21.  Spalangia drosophilae Ashmead, 1887 (Hymenoptera: Pteromalidae)

 

Head thicker than wide, with hairy punctures sparsely distributed, and ocellar line complete (Fig 29a-D).  Malar sulcus distinct.  Antennae with reticulated scape, pedicel ca. 2 1/4 X as long as first funicular segment.  Pronotal collar without anterior ridge, hairy punctures scarce or absent anterolaterally and transverse line of hairy punctures absent posteriorly (Fig. 29a-A & B).  Mesoscutum and scutellum with very few punctures (Fig. 29a-C).  Frenal groove indistinct.  Middle of propodeum with deep punctures forming a Y-shaped row (Fig. 29a-C).  Mesopleuron with set of punctures.  Mesosternum with short median longitudinal groove lacking punctures anteriorly.  Middle leg with one tibial spur with few bristles, bitial spur about 1/2 to 3/4 length of first tarsomere.  Hind leg with one smooth unbristled tibial spur.  Tergite 9 of abdomen with pair of slightly flattened sensory structures, each with 5 long setae and densely surrounded by short spines and setae.  Petiole rugosely punctuated dorsally and ventrally.  Length of female 0.90-1.60 mm, male 0.80-1.25 mm.  Hosts are primarily tiny Diptera breeding in a variety of habitats (e.g., Hippelates spp. in soil, Phoridae in dung, etc.).  Key references are Boucek (1963) and Rueda & Axtell (1985b). 

 

Additional references are Ashmead (1887), Lindquist (1936), Simmonds (1944, 1946, 1947a,b; 1952, 1953a,b; 1954, 1956), Legner & Bay (1964a,b, 1965b,c), Legner et al. (1966a,b,c,), Legner (1967a, 1968, 1969a), Capehart et al (1981), Marshakov (1983).

 

22.  Spalangia endius Walker, 1839 (Hymenoptera: Pteromalidae)

 

(Spalangia muscidarum var. stomoxysiae Girault, 1916; Spalangia philippinensis Fullaway, 1917; Spalangia muscidarum var. texensis Girault, 1920; Spalangia orientalis L. F. Graham, 1932; Spalangia stomoxysiae Peck, 1951).

 

The disc of the head between the eyes is sparsely punctured with the punctures mostly separated by more than their diameters (Fig. 37a-C).  The disc of the pronotum has an isolated wavy crossline consisting of large closely placed punctures in front of and parallel with the posterior margin (Fig. 37a-A & B).  The pronotal collar is rounded anteriorly, without a distinct ridge.  The anterio-lateral surface of the pronotum is umbilicately punctured with the interspaces smooth, not rugose (Fig. 37a-A & B).  The length of the gena is about equal to that of the eye (Fig. 37a-C).  The malar sulcus is distinct (Fig. 37a-D).  The ratio of abdominal petiole length to the narrowest width is 1.7 in female and 2.0 in male, with lateral hairs rare (Fig. 37a-E & F).  The length of the extended body is 2.0-3.0 mm in female and 1.9-2.6 mm in male.  A cosmopolitan species.  Key references are Boucek (1963), Legner (1965a, 1967b, 1979b,c), Legner et al. (1965, 1976), Legner & Brydon (1966), Legner & Greathead (1969),  Ables & Shepard (1974b), Morgan et al. (1975a,b; 1976a,b; 1977, 1978a, 1979a, 1981a,b; 1986), Morgan & Patterson (1975b, 1977, 1989), Morgan (1981b), Rueda & Axtell (1985). 

 

Additional references are Walker (1839, 1846), Girault (1915, 1916, 1920), Fullaway (1917), Graham (1932), Handschin (1932), Peck (1951), Dresner (1954), Shibles (1969), Yust (1970), Tingle & Mitchell (1975), Ables & Shepard (1976b), Ables et al. (1976), Weidhaas et al. (1977), Ienistea & Fabritius (1978), Schmidt & Morgan (1978), Thornberry & Cole (1978), Rutz & Axtell (1980a), Morgan (1980a,b; 1981b, 1985), Morgan et al. (1981b, 1986), Merritt et al. (1981), Petersen et al. (1983), Propp & Morgan (1983a,b; 1984a,b; 1985a,b), Donaldson & Walter (1984), Stafford et al. (1984), Bathon & Fabritius (1985), Bloomcamp (1985), Cabrales et al. (1985), Arellano & Rueda (1888), Xue (1988c), Xue et al. (1987a, 1989).

 

23.  Spalangia gemina Boucek 1963 (Hymenoptera: Pteromalidae)

 

Head facial view subtriangular, almost as wide as long, with large moderately prominent eyes and genae converging in a straight line (Fig. 33b-A).  Frons densely punctuated, interspaces very narrow to about as wide as punctures (Fig. 33b-A).  Antennal sockets quite deep, transversely ribbed in deepest parts, smooth in the middle except fine rugosity between antennal sockets.  Side view of head ca. 2X as long as thick.  Antennae short, slightly thickened at tip; scape dull, granulated and slender, at least as long as 5 following segments combined (Fig. 33b-B).  Pronotum with transverse ridge, collar not margined anteriorly, with very deep, arched, coarsely crenate cross line off hind margin.  All surface before this furrow coarsely reticulated rugose save for a low triangle just at cross line.  Hairy punctures elevated.  Abdominal petiole about 1.4X as long as broad, with almost parallel sides, bare with distinct ridges.  The body is black.  Tarsi are pale testaceous except the dark claw segments.  Length of female  3.1-4.1 mm, male 2.6-3.5 mm.  Original distribution Mauritius, South Asia, Fiji & tropical South America [probably established in Florida].  Key references are Boucek1963, 1965), and Morgan et al. (1991).

 

24.  Spalangia longepetiolata Boucek, 1963 (Hymenoptera: Pteromalidae)

 

The disc of the head between the eyes is sparsely punctured, the punctures being separated mostly by more than their diameters (Fig. 32b-A).  The disc of the pronotum has a distinct wavy coarse line of large crowded punctures parallel to and just anterior to the posterior margin (Fig. 32b-B & C).  The pronotal collar is rounded at the anterior margin.  The anterio-lateral surface of the pronotum is rugose or crowdedly rugosely punctured (Fig. 32b-B & C).  The length of the gena is greater than that of the eye.  The ratio of abdominal petiole length to the narrowest width is 1.5 in female and 3.5 in male (Fig. 32b-D & E).  Thus, female do not have the long petiole which is found in male and which generated the name of this species (Boucek 1963).  Lateral hairs absent on petiole.  The length of the extended body is ca. 2.2-3.1 mm in female  and 2.0-2.8 mm in male.  The species was originally known from Central Africa.  Additional key references are Boucek (1965), and Legner & Olton (1969b).

 

25.  Spalangia nigra Latreille, 1805 (Hymenoptera: Pteromalidae)

 

(Spalangia hirta Haliday, 1833; Spalangia rugosicollis Ashmead, 1894; Spalangia muscae Howard, 1911).

 

The disc of the head between the eyes is very densely crowdedly punctured, the punctures being separated by less than their diameters (Fig. 36b-B).  The pronotum is without a distinct wavy line of large punctures parallel to the posterior margin, and the pronotal collar is rounded anteriorly without a distinct ridge (Fig. 36b-C & D).  The anterio-lateral surface of the pronotum is rugose or crowdedly ruguosely punctured.  The length of the gena is greater than that of the eye (Fig. 36b-B).  The malar sulcus is indistinct (Fig. 36b-E).  The ratio of abdominal petiole length to the narrowest width is 2.0 in female and 2.2 in male (Fig. 36b-G).  There are at least 12 lateral hairs present on each side of petiole.  The length of the extended body is 3.0-4.5 mm in female and 2.5-3.7 mm in male.  This was originally a Holarctic species.  General appearance in Fig. 36b-H & I.  Key references are Latreille (1805, 1809), Dalman (1820), Haliday (1833), Bouché (1834), Nees (1834), Curtis (1839), Howard (1911), Richardson (1913b), Graham-Smith (1919), Girault (1920), Clausen et al. (1927), Parker & Thompson (1928), Perron (1954), Boucek (1963), Legner (1969a), Rutz & Axtell (1980a), Hall & Fischer (1988).

 

26.  Spalangia nigripes Curtis, 1839 (Hymenoptera: Pteromalidae)

 

(Spalangia nigripes Curtis, 1839; Spalangia hyaloptera Förster, 1850; Spalangia formicaria Kieffer, 1905; Spalangia muscarum Girault, 1920).

 

The disc of the head between the eyes is sparsely punctured, the punctures being separated mostly by more than their own diameters (Fig. 37b-B).  The disc of the pronotum is without a distinct wavy cross-line of large crowded punctures parallel to and just anterior to the posterior margin (Fig. 37b-A).  The pronotal collar is rounded at the anterior margin (Fig. 37b-A).  The anterio-lateral surface of the pronotum is umbilicately punctured with the interspaces smooth.  The length of the gena is less than that of the eye (Fig. 37b-B).  Th ratio of abdominal petiole length to the narrowest width is 1.6 in female (Fig. 37b-C).  The length of the extended body is 2.5-3.7 mm in female and 2.0-3.1 mm in male.  This species was originally Holarctic in distribution.  General appearance in Fig. 37b-D.  Key references are Curtis (1839), Walker (1848), Förster (1850), Dalla-Torre (1898), Kieffer (1905), Girault (1920), and Boucek (1963).

 

27.  Spalangia nigroaenea Curtis, 1839 (Hymenoptera: Pteromalidae)

 

(Spalangia homalaspis Förster, 1850; Spalangia astuta Förster, 1851; Spalangia muscidarum Richardson, 1913; Prospalangia platensis Brèthes, 1915; ?Spalangia abenabooi Girault, 1932; Spalangia sundaica L. F. Graham, 1932; ?Spalangia mors Girault, 1933).

 

 

The disc of the head between the eyes is sparsely punctured with the punctures separated mostly by about their diameters.  The disc of the pronotum has an isolated wavy crossline consisting of closely spaced large punctures in front of and parallel to the posterior margin, and the pronotal collar is bordered anteriorly by a narrow groove setting off the ridge-like margin (Fig. 34a-A & B).  The anterio-lateral surface of the pronotum is rugose or crowdedly rugulosely punctured.  The length of the gena is less than that of the eye (Fig. 34a-C).  Middle of propodeum with parallel rows of punctures that diverge posteriorly (Fig. 34a-D).  The ratio of abdominal petiole length to the narrowest width is 1.7 in female and 2.2 in male (Fig. 34a-E & F).  The length of the extended body is 2.9-3.8 mm in female and 2.5-3.5 mm in male.  The species is cosmopolitan.  Key references are Curtis (1839), Förster (1850, 1851), Johnston & Bancroft (1912), Pinkus (1913), Richardson (1913a,b), Parker (1924), Parker & Thompson (1928), Girault (1932, 1933), Graham (1932), Handschin (1932), Lindquist (1936), Peck (1951), Boucek (1963), Legner (1965a), Legner & Brydon (1966), Legner & Olton (1968b), Legner et al. (1965, 1967), Hoelscher & Combs (1969), Azizov (1972), Kochetova 7 Azizov (1972),Rutz & Axtell (1980a,b), and Siafacas (1980).

 

28.  Spalangia rugulosa Förster, 1850 (Hymenoptera: Pteromalidae)

 

Head, excluding antennal sockets, irregularly coarsely puncuated (even on temples).  Genae converge and eyes large.  Eye longer than gena (Fig. 36a-B).  Head in lateral view thickened with temple about 2X narrower than eye.  Antennal scape dull, granulated, slender, as long as 5.5 following segments combined; 1st funicular segment longer than broad, the 2nd and 3rd quadrate, distal segments slightly transverse, clava as long as 2.7 preceding segments (Fig. 36a-A).  Pronotum somewhat globose, with a margin, irregularly crowdedly punctured except for a narrow strip at hind margin (Fig. 36a-C).  Punctures of this smooth strip are deeper as a wavy line.  Propodeum with complete median carina.  Abdominal petiole ca. 1.8X as long as wide, deeply incised submedially on anterior margin.  Third abdominal tergite 3X as long as the 2nd at meson.   Dense hairs of body black; body black, sometimes with a faint metallic hue on head and thorax; tarsi also black, anterior basitarsus brown.  Length of female 3.1-3.8 mm, male 3.1 mm. Principal host Muscina stabulans Fallen.  Original distribution Europe and Central Asia.  Key references are Förster (1850) and Boucek (1963).

 

Genus Sphegigaster (Hymenoptera: Pteromalidae)

 

29.  Sphegigaster spp. (Hymenoptera: Pteromalidae)

 

The wing venation is greatly reduced with a single vein along the anterior margin and a spur, the stigmal vein near its apex.  The antennae are located on the middle of the face between the anterior margins of the eyes, not on a platform.  They are of less than 14 segments with the first segment elognated, longer than the next two combined.  The pronotal disc is finely densely punctured.  The color is shining black with the legs testaceous.  The abdomen is smaller than the thorax in contrast to other genera treated here (Fig. 20b-A).

 

A Sphegigaster sp. was originally found active on Musca domestica and Stomoxys calcitrans only in East and South Africa (Legner & Olton 1968, Legner & Greathead 1969).  However, the genus is well represented by several species in the Palearctic (Nicholskaya 1952, Graham 1969).

 

Genus Urolepis Ashmead 1896 (Hymenoptera: Pteromalidae)

 

30.  Urolepis rufipes (Ashmead, 1896) (Hymenoptera: Pteromalidae)

 

(Urolepis Walker, 1946; Halizoa Förster, 1956;  Pteromalus subgen. Halizous Thomson, 1878; Halizoa rufipes Ashmead, 1896).

 

Head reticulate, wider than broad, lower margin of antennal sockets well above level of lower margin of eye.  Dorsocephalic ridges distinct.  Malar sulcus slightly distinct.  Antennae with last distal funicular segments and clava broad, two ring segments present and pedicel about 1 1/2X longer than first funicular segment (Fig. 24b-B).  Pronotal collar with anterior ridge and coarsely reticulate ventrolaterally.  Mesoscutum and scutellum coarsely reticulate (Fig. 24b-A).  Frenal groove usually not clearly apparent.  Propodeum with median plicae without lateral branches and bearing dense fine setae along lateral margin.  Prosternum with median groove.  Fore wing with marginal vein of uniform width, slightly wider than post marginal vein and apical margin with fringe setae.  Middle leg with one tibial spur bearing sparse bristles along proximal 2/3rds of entire length, spur ca. 1/2X as long as first tarsomere.  Hind leg with one tibial spur lacking bristles (Fig. 24b-C).  Tarsal claw bears 5 setae (Fig. 24b-D).  Tergum 9 of abdomen with a pair of projecting sensory structures each with 5 long setae.  Key reference is Rueda & Axtell (1985b).  Additional key references are Ashmead (1896b), Nabrotsky et al. (1974), Deonier & Regensburg (1978), Burks (1979), Petersen et al. (1985), Smith & Rutz (1985, 1986, 1987, 1991b), and Pawson et al. (1987).

 

Families Cynipidae, Figitidae & Eucoilidae (Hymenoptera: Cynipoidea)

 

Antennae are never elbowed, the scape only slightly longer than broad and slightly shorter than first flagellar segment.  Abdomen may be laterally compressed, the first abdominal segment is cylindrical or annular, or minute, indistinct, fused dorsally with tergite 2.  Parasitoids of synanthropic Diptera are rare among members of this family.  Key references are Keilin & Baume-Pluvinel (1913), Girault (1921), James (1928), Huzimatsu (1940), Thompson (1944), Jenni (1951), Weld (1952), Walker (1959), Hadorn & Walker (1960), Belizin (1963), Bakker et al. (1967, 1972), Streams (1968), Eijsackers & van Lenteren (1970), Nappi & Streams (1970), Riek (1970), van Lenteren (1972, 1976), van Lenteren & Bakker (1972, 1975, 1978), Samson-Boshuizen et al. (1974), Richards (1977), Quinlan (1978), Chabora & Smolin (1979), Chabora et al. (1979), Krombein et al. (1979), Diaz (1980), Veerkamp (1980), Vet & van der Hoeven (1984), Vet & van Alphen (1985), Subba-Rao & Hayat (1985), Quinlana (1986), Gauld & Bolton (1988), and Goulet & Huber (1993).

 

31.  Figites spp. (Hymenoptera: Cynipoidea: Figitidae)

 

Largest segment of abdomen (in lateral view) tergites II or III, and never more than 1 short tergite preceding the large tergite (Fig. 44a-A).  Mostly small insects; tergite II not forming 1/2 the abdomen (Fig. 44a-A).  They are solitary endophagous larval parasitoids with separate species occurring in different geographic, but rarely demonstrating high parasitization rates.  Key references are Legner & Olton (1968b), and Legner et al. (1967).

 

                 32.  Hexacola (= Trybliographa) spp. (Hymenoptera: Cynipoidea: Eucoilidae)

 

Scutellum with dorsal "cup"; pronotum pronounced, extending forward into an anterior plate with a strong posterior margin (Fig. 43a-A); separate species in different geographic areas.  Attack smaller species of Diptera (e.g., Chloropidae) at low parasitization rates.  Key references are James (1928), Roberts (1935), Wishart & Monteith (1947), Jenni (1951), Simmonds (1952), Wishart & Monteith (1954), Jorgenssen (1957), Mulla (1962), Makarenko (1963, 1965), Legner & Bay (1964, 1965), Legner & Olton (1969), Legner et al. (1966), Clausen et al (1965), Eskafi & Legner (1974a,b,c), and Jones & Hassell (1988).

 

Family Diapriidae (Hymenoptera: Proctotrupoidea)

 

Antennae inserted on middle of face (Fig. 48a-A & C).  Fore wings without apparent stigma, venation reduced with only a short marginal vein (Fig. 48a-B).  The principal genera are Trichopria and Phaenopria.  Phaenopria occidentalis Fouts (Fig. 48a-E) is parasitic on Hippelates spp. in California, while Trichopria spp. are parasitic on Hippelates spp. in the West Indies.. Key references are Thompson (1944), Sundholm (1964), Riek (1970), Richards (1977), Fabritius (1979a), Krombein et al. (1979), Teodorescu & Ursu (1979), Nixon (1980), Borror et al. (1981), Subba-Rao & Hayat (1985), Gauld & Bolton (1988) and Goulet & Huber (1993).

 

                                   33.  Trichopria spp. & Phaenopria spp. (Hymenoptera: Diapriidae)

 

Wings do not have veins except for a short marginal vein and with a stigma so near the margin that the stigmal vein is almost absent (Fig. 48a-A & B).  The antennae usually arise from a small platform in the middle of the face between the eyes (Fig. 48a-C).  The femora and apical 1/2 of the tibiae are swollen.  The body is polished black without ground sculpture and with scattered long setae.  The size varies with that of the host.  Species of this genus are usually solitary endophagous larval parasitoids.  Key references are Muesebeck (1961), , Legner et al. (1967); Legner & Olton (1968b).  Additional references are Cros (1935), Bailey (1947), Magnarelli & Anderson (1980), Bradley et al. (1984), Morgan et al. (1990), and Blanchot (1992a).

 

 

REFERENCES:  [Please refer to File:  <fly-par.ref.htm>;  & also to MELVYL Library ]

 

 

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NOTE:

 

     Illustrations were simplified, corrected and/or updated to suit the present key. 

          However, please refer to cited authors for greater detail.