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EPICNEMIAL CARINA
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Adeliinae
All of the species so far knownare solitary endoparasitoids of
leaf-mining moths, especially
(perhaps exclusively) of the
family Nepticulidae (Whitfield
and Wagner, 1991), and theyemerge from the host cocoon.
At least some of the species
resemble ants when they run
about on plants, and one species
is known to produce a formic
acid-like odor when handled
(Whitfield, 1989).
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Agathidinae
Members of the Agathidinae are solitary
koinobiont endoparasitoids of concealed
lepidopterous larvae except for members
of the tribe Disophrini which attack free-
living larvae. Members of the tribes
Agathidini, Earinini, and Eumicrodini
attack the first or second instar larvae of
their hosts; those of the Disophrini attack
later instars, and members of Cremnoptini
appear to be able to parasitize any larval
stage. In all cases the adult parasitoid
emerges after the final instar of the host
has spun its cocoon (Nickels et al., 1950;Dondale, 1954; Odebiyi and Oatman,
1972, 1977). Most species are diurnal but
many members of the Disophrini are
nocturnal with typical pale coloration and
enlarged ocelli.
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Alysiinae
All alysiines are koinobiont
endoparasitoids of cyclorrhaphousDiptera. They oviposit in host larvae
or eggs and emerge from the
puparium. Most species are solitary,
but several of the species of
AphaeretaFoerster are gregarious.
The Alysiini use a wide variety of
cyclorrhaphous hosts, often in moist
habitats and decaying, ephemeral
substrates.
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Aphidiinae
Aphidiines have been reared
extensively; all are solitary,koinobiont endoparasitoids of adult
and immature aphids.
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Apozyginae
The biology ofA. penyaiis
unknown but due to its
basal phylogenetic position
within the Braconidae and
its general similarity tosome members of the
Doryctinae (Sharkey 1993)
it may be an idiobiont
ectoparasitoid ofxylophagous coleopterous
larvae.
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Betylobraconinae
Biology unknown.Probably
enodparasitic on lepidopterous
larvae
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BraconinaeAs far as is known, all New World
braconines are idiobiont ectoparasitoidsof concealed holometabolous insect
larvae, especially of the Lepidoptera and
Coleoptera, though a few species,
mostly in the genusBracon, attack
concealed dipterous or sawfly larvae.
Braconines are synovigenic and theirlarge eggs are usually laid on the host
which was previously paralyzed by
injection of venom. Host feeding
appears to be quite common among the
smaller species and sometimes involves
the construction of feeding tubes. Bothsolitary and gregarious parasitism occur,
sometimes with closely related species
displaying different strategies.
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CardiochilinaeAll of the species for which host data are
available parasitize the larvae of Lepidoptera.
As far as is known, all of the species are
solitary, internal parasitoids, attacking the early
instar larvae and emerging from the late instar
larvae or prepupae. The known hosts are most
commonly from the Pyralidae and Noctuidae,
but a number of other host groups have been
recorded. All species appear to carrypolydnaviruses (Stoltz et al., 1984) with which
they compromise the immune system and
physiology of their host insects. The most fully
studied species is Toxoneuron(usually referred
to as Cardiochiles)nigriceps(Viereck), a
parasitoid of the tobacco budworm,Heliothisvirescens.
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Cenocoeliinae
As far as known, all species aresolitary koinobiont
endoparasitoids of coleopteran
larvae with an endophytic way
of life. Most of the host records
for Nearctic species are fromCerambycidae, but some
Neotropical species have been
reared from seed-eating
Curculionidae.
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Capitonius
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Dirrhopinae
The only host records are fromleaf-mining moth larvae of the
family Nepticulidae.
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DoryctinaeVery little is known about the habits of this
subfamily relative to the number of
described species. Most species appear to be
idiobiont ectoparasitoids of wood-boring
beetle larvae but a few attack stem boring
lepidopterous and sawfly larvae. The host
larva is paralyzed before the egg is
deposited and the parasitoid cocoon isusually formed in the host tunnel or mine.
One described and several undescribed
species of the genusAllorhogasare
phytophagous in seeds. Species of
Psenobolushave been reared from figs in
Costa Rica where they display extremesexual dimorphism in the males, similar to
that found in chalcid fig wasps.
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Gnamptodontinae
Gnamptodontines have been rearedexclusively from leaf-mining
Lepidoptera of the family
Nepticulidae. Although apparently
koinobiont parasitoids (Shaw and
Huddleston, 1991), it is not yetknown whether gnamptodontines are
endo- or ectoparasitic. Detailed
biological studies are lacking.
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HisteromerinaeObservations on the Eurasian species
H. mystacinusWesmael indicate that
Histeromerinae are gregarious
idiobiont ectoparasitoids of wood-
inhabiting beetle larvae, prepupae
and pupae. Hosts are paralyzed and
the female appears to remain with asingle host after oviposition. Shaw
(1995) suggested that the ovipositor
may not be used for oviposition but
rather that the egg may emerge
directly from the genital orifice at the
ovipositor base.
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Homolobinae
Homolobines are solitary, koinobiont
endoparasitoids of Lepidoptera. Noctuidae
and Geometridae are the most commonly
recorded hosts ofHomolobus. Most species
are nocturnal.
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HormiinaeAll of the genera treated here are either
known to be or suspected to be
ectoparasitoids, usually attacking
concealed hosts. An important exception
isMonitoriella Hedqvist, which is
phytophagous. Many or most species
usually placed in the Rhyssalini,Hormiini, Pambolini and probably also
Hydrangeocolini are gregarious. Most of
the Hormiinae are also idiobionts and
develop rapidly on late-instar host larvae.
As far as is known, the vast majority of
the species parasitize hosts that areconcealed in some way (e.g. leaf-miners,
leaf-rollers, leaf-tiers, stem borers, seed
borers, gall formers, or borers in fungi).
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Pambolus
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Ichneutinae
Most members of the Ichneutinae are
koinobiont endoparasites of sawfliesof the families Argidae and
Tenthredinidae, however one lineage,
comprised of the genera
Oligoneurus,Paroligoneurus, and
Lispixys, has switched to leaf-mininglepidopterous hosts. The few species
with known biologies suggest that all
members of the family may be egg-
larval parasitoids, attacking the egg
but emerging from the last larvalinstar.
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Masoninae
Biology unknown.
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Mendesellinae
Only one species of thesubfamily has been reared
from its host. Epsilogaster
bicoloris a parasitoid of a
momphid (Lepidoptera)stem galler on Cephalanthus
occidentalis(buttonbush).
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M i
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Meteorinae
The Meteorinae are solitary or gregarious
koinobiont endoparasitoids of larval
Coleoptera or Lepidoptera, and many species
ofMeteorushave broad host ranges. The vast
majority of meteorines are solitary parasitoids
attacking exophytic (exposed-feeding)
lepidopteran larvae, and many are nocturnal.
Others utilize hosts that are only weaklyconcealed (e.g. in leaf rolls or under
webbing). The solitary parasitoids of arboreal
Lepidoptera typically emerge from the host
larva and pupate away from the host remains
in a cocoon that is often suspended by a long
slender thread, and it is from this
characteristic cocoon that the genusMeteorus
gained its name.
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Mi t i
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Microgastrinae
Microgastrine species attack virtually the
entire taxonomic and biological spectrum of
Lepidoptera, with the possible exception of
Hepialidae and a few other primitive
lineages of Lepidoptera. All species are
koinobiont, endoparasitoids of larvae, and
exit the host to pupate. A few species spin
their cocoons within the cocoon or shelter ofthe prepupal host. The majority of the
species are solitary, but a very large number
of species are gregarious, and from larger
hosts such as Sphingidae and Saturniidae,
hundreds of larvae may emerge to spin theircocoons either singly or in masses, and the
structure of these cocoon masses can be
diagnostic.
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Microtypinae
As far as is known, all species aresolitary koinobiont endoparasitoids of
microlepidopteran larvae with an
endophytic way of life. The known
hosts of Microtypinae belong to the
lepidopteran families Pyralidae,Tortricidae, Gelechiidae, and
Yponomeutidae.
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Miracinae
Many of the species have beenreared, always from leaf-miners,
usually from Nepticulidae or
Heliozelidae but also from some
Gracillariidae and Tischeriidae.The larvae are endoparasitoids;
the adults emerge from the host
cocoon.
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Opiinae
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Opiinae
Opiines oviposit in either the egg or
larval stage of their host and emergefrom the host puparium. Solitary
endoparasitoids, with the vast majority
of the rearings from either
Agromyzidae or Tephritidae. Although
at least 13 other families of Dipterahave been recorded as hosts of the
Opiinae (Fischer, 1971), many of these,
particularly from the earlier literature,
need confirmation.
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Most Opiinae
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Orgilinae
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Orgilinae
As far as known, all species are
solitary koinobiont endoparasites
of lepidopteran larvae. The
largest genus, Orgilus, contains
species attacking
microlepidopteran larvae with a
partly endophytic way of life,especially those with leaf-mining
or tunnelling early instars, whose
activities are detectable by
extruded frass. The known hosts
of Orgilinae belong toColeophoridae, Gelechiidae,
Tortricidae, Pyralidae and
Oecophoridae.
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Sigalphinae
Members of the Sigalphinae arekoinobiont endoparasites of larval
Noctuidae (Lepidoptera).
Cushman (1913) observed first
instar larvae being attacked by
Sigalphus bicolor(Cresson)though no adults were successfully
reared. Van Achterberg and
Austin's (1992) hypothesis that
sigalphines may be egg-larval
parasites because they have acarapace-like metasoma appears
somewhat contradicted by this
evidence.
Ypsistocerinae
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Ypsistocerinae
All known species in two of the three
genera, viz. Ypsistocerus manni,Y.
vestigialisand Termitobracon
emersoni, have been recorded from
nests ofNasutitermesspecies.
Whether ypsistocerines are actually
parasitoids of the termites themselves
rather than of some group of
inquilines, is unknown.