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HYMENOPTERA, Pteromalidae (Walker 1835) -
(Chalcidoidea) -- <Images> & <Juveniles> Please refer also
to the following links for details on this group: Pteromalidae. -- The PteromaIids
are a large group of about 400 North American species. They are tiny black or metallic-green or
bronze wasps . They are parasitic and attack a wide variety of hosts with
many being valuable in the biological control of crop pests. The adults of
many species feed on the body fluids of the host, which exude from punctures
made by the parasite's ovipositor (see Photo #3 to the right below). In Habrocytus cerealellae (Ashmead) that
attacks larvae of the angoumois grain moth and where the host larvae are in
the seed and out of reach of the adult pteromalid, a viscous fluid is
secreted from the ovipositor that is then formed into a feeding
tube extending down to the host larva. The adult sucks up the body fluids
of the host through this tube. Pteromalidae now also includes the former separate families,
Cleonymidae, Miscogasteridae and Spalangiidae, which have been designated
subfamilies Cleoneminae, Miscogasterinae and Spalangiinae, respectively. For the present, discussions of the various subfamilies will be separate because
of considerable distinctness among them.
The families Asaphinae, Austroterobiinae, Austrosystasinae,
Brachyscelidiphaginae, Ceinae, Cerocephalinae, Chromeurytominae, Cleonyminae,
Coelocybinae, Colotrechinae, Cratominae, Diparinae, Ditropinotellinae,
Eunotinae, Erotolepsiinae, Eunotinae, Eutrichosomatinae, Herbertinae,
Keiraninae, Leptofoeninae, Louriciinae, Macromesinae, Miscogasterinae,
Neodiparinae, Nefoeninae, Ormocerinae, Panstenoninae, Parasaphodinae, Pireninae,
Pteromalinae, Spalantiinae and
Storeyinae are treated in the respective files, <ASAPHINA.TXT>,
<AUSTROTE.TXT>, <AUSTROSY.TXT>, <BRACHYSC.TXT>,
<CEINAE.TXT>, <CEROCEPH.TXT>, <CHROMEUR.TXT>,
<CLEONYMI.TXT>, <COELOCY.TXT>, <COLOTREC.TXT>,
<CRATOMIN.TXT>, DIPARINA.TXT>, <DITROPIN.TXT>,
<ELATOIDI.TXT>, <EROTOLEP.TXT>, <EUNOTINA.TXT>,
<EUTRICHO.TXT>, <HERBERT.TXT>, <KEIRANI.TXT>,
<LEPTOFOE.TXT>, <LOURICI.TXT>, <MACROMES.TXT>,
<MISCOGAS.TXT>, <NEODIPAR.TXT>, <NEFOEN.TXT>, <ORMOCER.TXT>,
<PANSTENO.TXT>, <PARASAPH.TXT>, <PIRENIN.TXT>,
<PTERONAE.TXT>,
<SPALANGI.TXT> and <STOREYIN.TXT> [see Taxnames for more
detail]. Among Chalcidoidea, the Pteromalidae are one of the most common
families containing many genera and species of frequent encounter as
parasitoids or hyperparasitoids of various insect pests. Dominant genera include Pteromalus, Habrocytus, Dibrachys
and Pachyneuron. The family Spalangiidae is frequently
included under Pteromalidae and is represented by many species of Spalangia (see Bou
ek
1963). Most species are external
gregarious parasitoids of larvae and pupae of Lepidoptera and Coleoptera, but
some also attack pupae of Diptera and larvae of Hymenoptera. Genera such as Spintherus, Enargopelte,
and Peridesmia are egg
predators. External and internal
parasitism within a genus is found in Dibrachys,
Pteromalus and Stenomalus. Ophelosia crawfordi Riley occurs as a predator on the eggs of Pulvinaria, Pseudococcus and Icerya
(Smith & Compere 1931), and as a hyperparasitoid of these genera and
sometimes of larvae of Coccinellidae.
It has been reared from Icerya
females where hyperparasitism seemed possible through Cryptochaetum. Some
species of Asaphes are
hyperparasitoids of Aphididae through various braconids, aphelinids and
encyrtids that behave as primary parasitoids (Griswold 1929). Dibrachys
cavus Wlk. (= boucheanus Ratz.) attacks a wide range of hosts with over 45 host
species comprising 2 Coleoptera, 2 Diptera, 27 Hymenoptera and 14 Lepidoptera
(Faure & Zolstorewsky 1925). Most Pteromalidae are primary parasitoids, but hyperparasitic
species are common. Most species are
ectoparasitic, but endoparasitic species are common also. Solitary and gregarious species and races
are common. Generally, this family
has a wide host range. Most species
are gregarious ectoparasitoids of larvae and pupae of Lepidoptera and
Coleoptera, but a number of species attack larvae and pupae of Diptera as
well. Some are predaceous on eggs of
Coccidae. There are no phytophagous
species. Considerable importance has
been placed on pteromalids for biological control of Lepidoptera, Coleoptera
and synanthropic Diptera. A few species
have also been used for the biological control of Coccidae. Pteromalus puparum L., a gregarious internal parasitoid of cabbage butterfly
pupae and Ascia rapae L. in New Zealand have been
credited with marked reductions in host population densities in biological
control efforts. Species of Muscidifurax, Urolepis and Pachycrepoideus
have been deployed successfully against synanthropic Diptera (Legner et al.
1976, Rueda & Axtell 1985). Biology
& Behavior
Extensive studies have been made on species in the genera Dibrachys, Dibrachoides, Pteromalus,
Eupteromalus, Habrocytus, Muscidifurax,
and Nasonia (Mormoniella). Variation within a genus and species is aptly demonstrated in the
genus Muscidifurax, which is a
group of closely related species (superspecies) attacking puparia of
synanthropic Diptera. The five described
species occur in geographic isolation in the Nearctic and Neotropical
regions, except two species which are sympatric in the western Nearctic
(Legner 1969, 1983, Kogan & Legner 1970, Kawooya 1983). The suspected ancestor of this apparent
clade, M. raptor Girault & Sanders, is widely distributed in Europe,
Africa, North America, and portions of the Pacific area (Legner & Olton
1988, 1971; Legner et al. 1976, Rutz & Axtell 1980, Legner 1987, Smith et
al. 1987), but it has not been found in the Neotropics. There are no known clinal patterns. The genus has not been reported from Asia
and is poorly represented in equatorial regions (Legner & Olton 1968,
Legner & Greathead 1969, Legner et al. 1976, Legner 1983). Muscidifurax are most
prevalent in or near accumulated decaying organic material deposited by
humans and livestock, where they parasitize host Diptera that also breed
selectively in this habitat.
Therefore, they fit the endophilous synanthropic category (Povolny
1971, Legner et al. 1974), and their existence depends largely on
herdsmen. This has led to the
suggestion that the four species currently confined wholly to the Americas
could have evolved within the recent time period of European settlement, or during
the past 400 yr (Kogan & Legner 1970, van den Assem & Povel 1973). The only known South American member of the genus, M. raptorellus
Kogan & Legner, occurs as three or more separate populations. One population from coastal Peru is
solitary, whereas the others show various degrees of gregarious oviposition
and development (two or more eggs laid at one insertion) (Legner 1967, 1988,
Kogan & Legner 1970, Kawooya 1983).
One of the gregarious populations from central Chile oviposits more
than one egg in >60% of the hosts it attacks, with subsequent successful
gregarious development. The Chilean
population compensates for a lower host-searching capacity with the
gregarious behavior, which results in a greater number of progeny per host
(Legner 1967, 1987). Adult Habits.--The time between emergence of adults and
oviposition is short. Females of Nasonia vitripennis Wlk. (= Nasonia
brevicornis Ashm., Mormoniella vitripennis Wlk.) deposit eggs within three hours of emergence,
and Habrocytus cerealellae Ashm. does so the following day. Two to three days are required by Dibrachoides dynastes Foerst. and Homoporus
braconidis Ferr. for gestation, but
Pirene graminea Hal. lays its first eggs 7-8 days after adult emergence
(Clausen 1940/1962). As is typical for Hymenoptera, adults feed extensively on
honeydew, and plant secretions. Such
food is sufficient to maintain life, but in many species it is not sufficient
to meet the optimal nutritional requirements for egg production. It was demonstrated that a protein diet is
essential for some species before normal oögenesis can proceed. The body fluids of the host provide a
suitable food of this type, and the females of Pteromalidae more generally
than of any other family, have developed the habit of feeding on fluids that
exude from punctures made with the ovipositor. Roubaud (1917) seems to be the first to realize the
significance of host-feeding, as his work with Nasonia vitripennis
showed that such feeding was essential before normal oviposition could take
place. Such feeding may be associated
with and immediately precede oviposition, or the two acts may be entirely
independent and upon different host individuals (Clausen 1940/1962, Legner
& Gerling 1967). Feeding on
exposed hosts presents minimal difficulties for the parasitoid, but where
hosts are sequestered in a burrow, cocoon, cell or puparium, direct feeding
is not possible. Therefore, it is
necessary to provide some means by which the body fluids of the host can be
brought within reach of the parasitoid.
This is accomplished by the construction of a feeding tube which
extends from the oviposition puncture in the host's body to the outside wall
of the cell, etc. Fluids rise to the
top of the tube by capillary action and are there lapped up by the parasitoid. In some species, withdrawal of the body
fluids is so great as to suggest the deployment of actual suction to bring it
to the surface. This adaptation for feeding was first discovered by Lichtenstein
(1921) while studying Habrocytus cionicida Licht. and external parasitoid
of larvae and pupae of the weevil, Cionus
thapsi F., in their cocoons. The habit has since been found in many
other genera of Pteromalidae and occasionally in other Chalcidoidea as
well. It also occurs among the
Braconidae, particularly in the genus Microbracon. A detailed account of the manner of tube
formation was given for Habrocytus cerealellae, parasitic on the
angoumois grain moth Sitotroga cerealella Oliv. (Fulton 1933). This parasitoid normally attacks larval
stages in cells in the seed, although parasitization and development are also
possible on exposed larvae. A feeding
tube is constructed only when the host is deeply embedded in the seed,
however. After stinging, the
ovipositor is withdrawn until only its tip penetrates through the hole in the
cell wall. A clear, viscid fluid
begins to ooze from it, most abundantly from near the tip. This material is spread by a twisting and
vertical movement of the ovipositor, which serves as a spatula. It is worked downward gradually, and fresh
material is added continuously until the body of the host is finally
reached. The ovipositor tip is then
slowly moved about until the original puncture in the skin is found. It is then reinserted in the puncture and
held in that position for several minutes, during which time the tube is
completed. The ovipositor is
withdrawn very slowly in order not to damage the delicate tube. In the meantime, the stylets move
alternately up and down. A small
extension of the tube appears above the surface of the opening in the cell
wall. The female then turns about and
begins feeding on the fluids from the tube, which may continue without
interruption for almost an hour. At
the completion of feeding, the female reinserts the ovipositor in the tube,
seemingly for the purpose of breaking it.
Some researchers have noted this reinsertion of the ovipositor by
different species and considered it to be for the purpose of inducing a
renewal of body fluids flow and that breaking the tube was only accidental. Faure & Zolstorewsky described an identical manner of tube
formation in Dibrachys cavus. H. D. Smith (1930) observed in Dibrachoides dynastes
Foerst., that the chalky fluid flows down the full length of the ovipositor
while its tip is still inserted in the wound, with hardening taking place
quickly after which the ovipositor was cautiously withdrawn. Flanders (1935b) noted tube formation
among egg predators. Female Spintherus sp. punctured one of a
cluster of eggs of the alfalfa weevil, Hypera
variabilis Hbst., that were
embedded in a plant stem. A tube was
formed and the entire egg contents was sucked out. Colleterial glands are thought to be the source of the tube
substance. Feeding is not always limited to host fluids but to tissues as
well. Noble (1932) reported that when
the ovipositor of Habrocytus cerealellae was withdrawn from the
wound the barbs of the sheath draw up strands of solid matter and then the
parasitoid chews vigorously upon them.
The same habit was found in Dibrachys
clisiocampae Fitch, attacking
mature larvae of the bee moth in its cocoon (Graham 1918). Detailed studies of the relation of the feeding habit of Peridesmia and Spintherus to egg development were made by Flanders. Sufficient protein was not stored up during
the larval period to provide for egg production, which must be accounted for
later. The preference for host fluids
is shown only during oögenesis and maturation. This reaction first appears about six days before egg
deposition. If environmental conditions
were adverse, the ovarian follicles were absorbed, feeding hon host fluids
ceased, and a long period of reproductive inactivity followed. The degeneration of the ovaries under such
conditions was considered to be a form of phasic castration. The induced inactivity corresponded in
time to periods of low numerical status of the host stage that was subject to
attack. It may have a direct effect
on the ability of a parasitoid species to survive where the host undergoes
long periods of aestivation. The female is able to develop and deposit eggs for a certain
period as a result of feeding on protein substances, but carbohydrates are
necessary to provide the energy required for extended activity. Females of Pteromalus puparum
deposited eggs for a period of three weeks when confined with fresh host
pupae, to which they were limited for food, while with the addition of honey
they continued oviposition for two months (Doten 1911). Most species that attack larvae of Lepidoptera and Coleoptera
permanently paralyze the host prior to oviposition. This is especially true in Habrocytus,
Dibrachys and Dibrachoides and some other less common genera. In H.
cerealellae the parasitoid pumps
several droplets of a paralyzing fluid into the host body, which can be readily
seen as it flows down the ovipositor.
This organ is thrust deeply into the host to ensure thorough
distribution of the fluid. The
stinging act requires up to 10 min., and the host is completely paralyzed
before the ovipositor is withdrawn.
In Drabrachoides dynastes the host may be stung 3-100
times before complete immobility, which may take up to 8hrs (Clausen
1940/1962). The larvae of the bee
moth, Galleria mellonella L., die from the effects of the sting of Dibrachys clisiocampae, and the eggs of the latter are not deposited until
death occurs. Doten reported that
larvae of the codling moth and pupae of the cabbage butterfly were usually
killed by thrusts of the Meraporus
sp. ovipositor. In a large number of
species hosts are consistently killed by the sting, and decay ensues very
soon thereafter. In such cases the
larvae are scavengerous in habit rather than parasitic. A few species do not paralyze the host,
among which are Trichomalus fasciatus Thoms., an external
parasitoid of Ceutorrhynchus assimilis Payk. larvae. Pirene
graminea paralyzes its host larva Contarinia pisi Winn. temporarily (Clausen 1940/1962). Some species that are parasitic in dipterous, seal the puncture
in the puparial wall with a drop of fluid after oviposition. Clausen (1940) noted the behavior of Scymnophagus townsendi Ashm., an external parasitoid of Scymnus sp. pupae, a coccinellid predator on aphids in
Japan. During oviposition the female
stands on the exposed portion of the dorsum of the pupa and makes a
preliminary insertion with the ovipositor to form a feeding puncture. Several hours may be spent in lapping up
the body fluids exuding from the wound, after which the ovipositor is
reinserted, usually in the original puncture, and forced completely through
the body. The egg is laid on the
ventral surface of the abdomen or under a wing pad or leg. Feeding and oviposition may be repeated,
the single original puncture serving for this purpose. A compact cluster of 4-6 eggs is usually
found when the host is finally abandoned. Fulton (1933) described the mechanics of oviposition in H. cerealellae,
noting the presence of a remarkable adaptation for the passage of a large
object through a relatively minute tube such as the ovipositor. The small end of the egg approaches the
ovipositor first; the spicules of the chorion, in conjunction with the
backward directed spines and ridges on the walls of the vagina and stylets,
prevent any backward movement once the surfaces are engaged. The greatly compressed egg tip is drawn
down through the ovipositor. The
chorion, being stretched, passes down the ovipositor by a pulling action rather
than any pressure exerted through the abdomen. Therefore, the portion of the chorion being drawn through the
tube is almost devoid of liquid content, and little lateral pressure is
exerted on the ovipositor. At this
time the stylets, moving together rather than alternately, work the tip of
the egg downward until it reaches the end of the ovipositor. Thereafter the egg contents flow down the
tube into the released portion, and by this action the remainder of the egg
is pulled through the ovipositor. The
reduction of egg diameter during its passage down the ovipositor is shown by
a comparison of egg size with ovipositor width. The egg averages 0.16 mm in width, while the outside diameter
of the ovipositor is ca. 1/5th of this, or 0.03 mm. The habit of adult swarming, which is rate in Chalcidoidea, was
recorded for Pteromalus deplanatus Nees (Scott 1919). During 1916-1919 in some localities in
England this species was present in huge swarms in buildings during late July
and August. Clausen (1940) believed this
to be a search for suitable quarters in which to pass the winter. Reproductive capacity of Pteromalidae is high. An egg deposition of 676 in 78 days has
been recorded by Fulton for H. cerealellae, and P. puparum has been
found to lay as many as 697 eggs. A
single female Nasonia vitripennis produced 557 progeny
(Cousin 1933), suggesting an egg capacity much in excess of that number. A series of Eupteromalus nidulans
Thoms. females laid an average of 251 eggs, with a maximum of 583 (Proper
1931). Aplastomorpha calandrae
How. lays ca. 250 and Dibrachoides dynastes a maximum of 122 during a
single month. Development of Larvae.--First instar larvae of species that feed
externally are usually very active, moving about readily on the body of the
host and in the cell containing it.
However, Smith (1930) found that larvae of D. dynastes which hatch
from eggs not placed directly on the host, usually die without reaching
it. In solitary species, there is a
pronounced cannibalistic tendency, and the larva that hatches first often
destroys any remaining eggs. In cases
where a number are able to hatch, the youngest, due to its greater mobility,
is usually victorious in the combat for the host. Such an elimination of surplus individuals seems necessary
because of the indiscriminate oviposition of the parent female, which is
apparently unable to recognize hosts that are already parasitized. Many individuals of Pteromalus
puparum develop within a single
host pupa, which results in the colony being too large for the available food
supply. Numerous dead larvae are
often found at the extremities of the pupa, these having died from starvation
(Hardy 1933). It is evident that the
food material at these points is exhausted more quickly than at the middle of
the body, as Faure (1926) found that development of individuals in the
extremities of the host is retarded.
But Voukassovitch (1926) concluded that the retardation in emergence
of a portion of the brood is independent of nutrition. It was found that emergence from pupae
that had been parasitized on known dates often extended over several
months. This appears to be a larval
diapause of uncertain duration and affects a varying portion of each
colony. Clausen (1940) regarded it
unusual for the individuals comprising a single colony of an internal
parasitoid to emerge so irregularly, and noted that it contrasts sharply with
the synchronous development and emergence of polyembryonic Encyrtidae, which
encounter the same adverse conditions through overcrowding, etc. However, Legner (1969) found that a spread
of emergence was characteristic of a number of parasitic Hymenoptera in
several families even though oviposition was limited to a 24h period. The spread was the result of differential
rates of development in different stages; and which stage differed varied
with the species. Some solitary ectophagous species show a considerable
adaptability with regard to the size of host individuals upon which
development can be successfully completed.
Noble found that Habrocytus cerealellae, when developing on small Sitotroga larvae, is able to pass through
the final larval stage without feeding and to attain the adult stage. In such cases the larval stage was
prolonged with resulting adults of minute size. Stenomalus micans Ol., 4th instar larvae, which
are parasitic in larvae of Chlorops
taeniopus Meig., bears a
specialized boring apparatus on its head that is used to break up the
internal organs of the host and to effect emergence through the hardened
shell of the host larva, which has died just as it was undergoing pupation
(Kearns 1931). No feeding occurs
during the 5th larval stage, which is very short in duration. Parasitism by Stenomalus results not only in appearance changes of the host
larva, but in its activity. Healthy
larvae move downward in a barley stem and, prior to pupation, turn about and
ascent to a point just below where the leaf leaves the stem. Reddish-brown puparia are then
formed. Parasitized individuals do
not make this position reversal, and partly formed puparia are colorless. Normally pupation in Pteromalidae occurs in the cell, cocoon or
other cavity in which the host resided.
However, Eupteromalus nidulans forms a naked pupa in the web
of its lepidopterous host. Nasonia vitripennis pupae retain the larval exuviae about the posterior
portion of the abdomen, and this, adhering to the meconium, attaches the pupa
to the host puparium wall. Enargopelte ovivora Ishii is one of the few Chalcidoidea showing a tendency
toward normal cocoon formation (Ishii 1928).
Mature larvae, of which there may be ca. 10 in the egg chamber of Lecanium sp., spin individual,
yellowish-brown cocoons. In some species many individuals are able to develop on a single
host. Martelli (1907) recorded 165
adults of Pteromalus puparum from a single pupa of the
cabbage butterfly, and Picard (1922) reared 212 males from the same host and
47 Tritneptis klugii Ratz. (= P. nematicidus Pack.) have been secured
from a cocoon of Lygaeonematus erichsonii Htg. (Hewett 1912). Roubaud reared 105 N. vitripennis from a
single dipterous puparium. Such figures
undoubtedly represent maximums for which food material was available. However, in most gregarious species
development to maturity is possible even if only a portion of the available
food material is utilized. The
different species of Dibrachys usually
develop in numbers of <10 on each host, and all the recorded species of Habrocytus are solitary. Muscidifurax
raptorellus K. & L. has both
solitary and gregarious races, the habit being under the control of polygenic
loci (Legner 1987d, 1988a, 1989b, 1991).
The gregarious races produce individuals of a characteristic size
(Kogan & Legner 1970). Life Cycle
Pteromalids usually have short life cycles, averaging ca. 3 weeks
from egg to adult at room temperature.
There was a minimum of 10 days recorded for Habrocytus cerealellae
and Nasonia vitripennis. The females
of many species require 1-2 days longer for development than do the
males. The incubation of the egg
requires from less than 1 day to 3 days, the larval stage 4-10 days and the
pupal stage 4-14 days. A notable
exception is E. ovivora, in which the egg, larval, and
pupal stages take 7 days, 20 days and ca. 11 months, respectively (Clausen
1940/1962). The availability of suitable host stages influences the number of
generations per year. Most species
produce generation after generation as long as hosts are available, but some
species are limited to a fixed number.
E. ovivora has only one generation per year which corresponds to the
host cycle. Pirene graminea and Stenomalus micans have two generations, as do their respective hosts. However, Aplastomorpha calandrae
(Cotton 1923) and H. cerealellae are able to produce
several generations to each one of the host. In these species there is no
need for synchronization of the cycles of parasitoid and host, for they
attack insects infesting stored grains which have all stages present
continuously. Trineptis klugii has
ca. 6 generations each year on one brood of the host. Most species that hibernate do so in the mature larval stage
within the host cocoon, puparium or cell.
But, Eupteromalus nidulans is found in the hibernation
webs of the satin and brown-tail moths.
E. ovivora, Rhopalicus suspensus Ratz., and Merisus febriculosus Gir. pass the winter in the pupal stage, while Dibrachoides dynastes and Pseudocatolaccus
asphondyiiae Masi persist through
the winter as adults. Other may pass
winter as either mature larvae or adults. A number of Pteromalidae are able to undergo long periods of
inactivity as either larvae or adults when conditions are unfavorable. The relation of food to reproduction in Spintherus and Peridesmia was already noted, and it was shown that phasic
castration in females may continue for a long time. This is one way of maintaining a species during periods of
adverse conditions; another is larval diapause, such as is found in H. medicaginis
Gahan and Nasonia vitripennis. In the former individuals have been observed to pass almost two
years in the larval stage, as compared to the normal two weeks. Nasonia
vitripennis may even pass several
years in dipterous puparia when conditions are unfavorable (Clausen
1940/1962). Adults live much longer than many other chalcidoid
parasitoids. Species which normally pass
the winter as adults are exceptionally hardy, and D. dynastes has been
kept alive for >8 months at temperatures of 5-13°C. Species without diapause usually live 6-8
weeks. Sex
Ratio & Parthenogenesis
Females predominate in ratios up to 30:1 in Habrocytus medicaginis. Clausen (1940) noted that ratios for N. vitripennis
vary from 1:1 to 10:1. The normal
ratio for P. puparum is ca. 2:1; but George (1927) noted a seasonal variation,
the ratio being 2.8:1 in springtime and 1.1:1 in autumn. In H.
cerealellae, the field sex ratio is
ca. 3:2. Experimental determinations
give an increasing proportion of male progeny toward the end of the life of
females (Clausen 1940/1962; Legner & Gerling 1967), even in thelytokous populations
(Legner 1987c). Griswold (1929)
recorded an excess of males in the ratio of ca. 3:1 in some rearings of Asaphes americana from aphids collected in glasshouses. Extended studies in thelytoky have been done on Muscidifurax uniraptor Kogan & Legner, showing the presence of extranuclear
influences (Legner 1985a,b; 1987b,c; 1988c).
Certain bacteria have been found in the male and female reproductive
tracts, which are capable of inducing endomitosis in, unfertilized eggs,
thereby causing them to be diploid and female (E. F. Legner unpub. data, R.
Stouthamer unpub. data). For detailed descriptions of immature stages of Pteromalidae,
please see Clausen (1940/1962). References: Please refer to
<biology.ref.htm>,
[Additional references may be found at: MELVYL Library] Askew, R. R.
1961. Trans. Roy. Ent. Soc.
London 113: 155-73. Bou…ek, Z.
1958.
Eine Cleonyminen-Studie:
Bestimmungstabelle der Gattungen mit Beschreibungen und Notizen,
eingeschlossen einige Eupelmidae. Sb. Ent. Odd. Nár. Mus.
Praze 33: 384-486. Bou…ek, Z.
1959. On Chalcedectus sinaiticus
(Masi) from the Near East and Ch. quarantiticus (Strand), from Paraguay,
and a new synonymy. Sb. Ent. Odd.
Nár. Mus. Praze 33: 429-602. Bou…ek, Z.
1963. A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Acta. Ent. Mus. Nat. Prague 35: 429-512. Bou…ek, Z.
1974. The pteromalid subfamily
Eutrichosomatinae (Hymenoptera: Chalcidoidea). J. Ent. (B) 43(2):
129-38. Bou…ek, Z.
1978. A generic key to
Perilampinae (Hymenoptera, Chalcidoidea), with a revision of Krombeinius n. gen. and Euperilampuus Walker. Ent. Scand. 9(4): 199-307. Bou…ek, Z.
1988. Australasian
Chalcidoidea, a biosystematic revision of genera of fourteen families, with a
reclassification of species. CAB
Internatl., Wallingford, UK. 832 p. Burks, B. D.
1958. A North American Colotrechnus (Zanonia) (Hymenoptera: Pteromalidae). Fla. Ent. 41: 13-16. Burks, B. D.
1965. The North American
species of Metacolus (Hymenoptera,
Pteromalidae). Proc. Ent. Soc. Wash.
67(2): 116-19. Burks, B. D.
1969. The North American
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