EPICNEMIAL CARINA

Adeliinae
All of the species so far known
are solitary endoparasitoids of
leaf-mining moths, especially
(perhaps exclusively) of the
family Nepticulidae (Whitfield
and Wagner, 1991), and they
emerge 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).

Agathidinae
Members of the Agathidinae are solitary
koinobiont endoparasitoids of concealed
lepidopterous larvae except for members
of the tribe Disophrini which attack freeliving 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.

Alysiinae
All alysiines are koinobiont
endoparasitoids of cyclorrhaphous
Diptera. They oviposit in host larvae
or eggs and emerge from the
puparium. Most species are solitary,
but several of the species of
Aphaereta Foerster are gregarious.
The Alysiini use a wide variety of
cyclorrhaphous hosts, often in moist
habitats and decaying, ephemeral
substrates.

Aphidiinae
Aphidiines have been reared
extensively; all are solitary,
koinobiont endoparasitoids of adult
and immature aphids.

Apozyginae
The biology of A. penyai is
unknown but due to its
basal phylogenetic position
within the Braconidae and
its general similarity to
some members of the
Doryctinae (Sharkey 1993)
it may be an idiobiont
ectoparasitoid of
xylophagous coleopterous
larvae.

Betylobraconinae
Biology unknown.Probably
enodparasitic on lepidopterous
larvae

Blacinae
The biology of all but a few species
is unknown. Blacus Nees are
parasitoids of larval Coleoptera.
Records of species being parasitic
on dipterous larvae are almost
certainly wrong. Some species of
Blacus have male mating swarms.
The two described species of
Dyscoletes Haliday are parasites of
larval Boreidae of the genus Boreus
(Mecoptera).

Braconinae
As far as is known, all New World
braconines are idiobiont ectoparasitoids
of concealed holometabolous insect
larvae, especially of the Lepidoptera and
Coleoptera, though a few species,
mostly in the genus Bracon, attack
concealed dipterous or sawfly larvae.
Braconines are synovigenic and their
large 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. Both
solitary and gregarious parasitism occur,
sometimes with closely related species
displaying different strategies.

Cardiochilinae
All 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 carry
polydnaviruses (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, Heliothis
virescens.

Cenocoeliinae
As far as known, all species are
solitary koinobiont
endoparasitoids of coleopteran
larvae with an endophytic way
of life. Most of the host records
for Nearctic species are from
Cerambycidae, but some
Neotropical species have been
reared from seed-eating
Curculionidae.

Capitonius

Cheloninae
Chelonines are solitary koinobiont egglarval endoparasitoids of Lepidoptera
that feed largely in concealed situations,
especially Tortricoidea and Pyraloidea.
Many host larvae are borers in stems,
buds, or fruit, while others create a
cryptic feeding retreat by rolling, folding,
or tying leaves with silk. In some cases,
host location and oviposition may be
simplified by attacking eggs which are
numerous and exposed. Other species,
with long ovipositors, may attack eggs in
more concealed situations. Although
egg-larval parasitoids occur in other
braconid subfamilies (e.g. Alysiinae,
Helconinae, Ichneutinae), chelonines are
the most common braconids that employ
this strategy.

Dirrhopinae
The only host records are from
leaf-mining moth larvae of the
family Nepticulidae.

Doryctinae
Very 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 is
usually formed in the host tunnel or mine.
One described and several undescribed
species of the genus Allorhogas are
phytophagous in seeds. Species of
Psenobolus have been reared from figs in
Costa Rica where they display extreme
sexual dimorphism in the males, similar to
that found in chalcid fig wasps.

Euphorinae
Euphorines are solitary or (more
rarely) gregarious koinobiont
endoparasitoids of adult Coleoptera,
adult Hymenoptera, or adult
Neuroptera, and of nymphal and
adult Heteroptera and Psocoptera. As
a group, the subfamily Euphorinae
has a host range that is substantially
broader than other braconid
subfamilies, presumably because of
diversification following the
evolution of parasitism of adult
insects.

Leiophron

Gnamptodontinae
Gnamptodontines have been reared
exclusively from leaf-mining
Lepidoptera of the family
Nepticulidae. Although apparently
koinobiont parasitoids (Shaw and
Huddleston, 1991), it is not yet
known whether gnamptodontines are
endo- or ectoparasitic. Detailed
biological studies are lacking.

Helconinae
Members of the Helconini are
solitary endoparasitoids of
cerambycid beetle larvae and
possibly other wood-boring
beetles. The biology of the
members of the Diospilini is
largely unknown. Where known,
species are solitary
endoparasitoids of phytophagous
beetles such as Curculionidae
and Nitidulidae. Members of the
Brachistini are known to be egglarval endoparasitoids of
Coleoptera, usually in the
families Curculionidae or
Bruchidae.

Histeromerinae
Observations on the Eurasian species
H. mystacinus Wesmael indicate that
Histeromerinae are gregarious
idiobiont ectoparasitoids of woodinhabiting beetle larvae, prepupae
and pupae. Hosts are paralyzed and
the female appears to remain with a
single 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.

Homolobinae
Homolobines are solitary, koinobiont
endoparasitoids of Lepidoptera. Noctuidae
and Geometridae are the most commonly
recorded hosts of Homolobus. Most species
are nocturnal.

Hormiinae
All of the genera treated here are either
known to be or suspected to be
ectoparasitoids, usually attacking
concealed hosts. An important exception
is Monitoriella 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 are
concealed in some way (e.g. leaf-miners,
leaf-rollers, leaf-tiers, stem borers, seed
borers, gall formers, or borers in fungi).

Pambolus

Ichneutinae
Most members of the Ichneutinae are
koinobiont endoparasites of sawflies
of the families Argidae and
Tenthredinidae, however one lineage,
comprised of the genera
Oligoneurus, Paroligoneurus, and
Lispixys, has switched to leaf-mining
lepidopterous hosts. The few species
with known biologies suggest that all
members of the family may be egglarval parasitoids, attacking the egg
but emerging from the last larval
instar.

Macrocentrinae
The Macrocentrinae include
both solitary and gregarious
koinobiont endoparasitoids of
lepidopterous larvae. Where
known, the gregarious species
are polyembryonic. Most
macrocentrines are pale colored
and crepuscular or nocturnal.
The species of Macrocentrus
have long ovipositors, often
laying their eggs in early instars
of hosts concealed in leaf
whorls, stems, tubers, or rolled
leaves.

Masoninae

Biology unknown.

Mendesellinae
Only one species of the
subfamily has been reared
from its host. Epsilogaster
bicolor is a parasitoid of a
momphid (Lepidoptera)
stem galler on Cephalanthus
occidentalis (buttonbush).

Meteoridinae
Members are known to be larvalpupal endoparasitoids of
lepidopterous larva. Eggs are laid
into the host larva and the adult
parasitoid emerges from the pupal
stage of the host. Nocturnal.

Meteorinae
The Meteorinae are solitary or gregarious
koinobiont endoparasitoids of larval
Coleoptera or Lepidoptera, and many species
of Meteorus have 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 weakly
concealed (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 genus Meteorus
gained its name.

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 of
the 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 their
cocoons either singly or in masses, and the
structure of these cocoon masses can be
diagnostic.

Microtypinae
As far as is known, all species are
solitary 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.

Miracinae
Many of the species have been
reared, 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.

Neoneurinae
There is little detailed information on the
biology and behavior of neoneurines, but
they are thought to be endoparasitoids.
Although oviposition by neoneurines
into the abdomens of adult worker ants
has been observed by several authors,
the details of larval development are not
known. The process of egg deposition is
extremely rapid, taking only a fraction
of a second. Wasmann (1897) reported
rearing an adult of Elasmosoma from a
cocoon found attached to the abdomen
of a dead worker ant (Formica sp.).
Neoneurines fly close to the ground and
stay very near nests and trails of
Formica ants.

Opiinae
Opiines oviposit in either the egg or
larval stage of their host and emerge
from the host puparium. Solitary
endoparasitoids, with the vast majority
of the rearings from either
Agromyzidae or Tephritidae. Although
at least 13 other families of Diptera
have been recorded as hosts of the
Opiinae (Fischer, 1971), many of these,
particularly from the earlier literature,
need confirmation.

Most Opiinae

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 to
Coleophoridae, Gelechiidae,
Tortricidae, Pyralidae and
Oecophoridae.

Rogadinae
Rogadinae are koinobiont
endoparasitoids, mostly of exposedfeeding macrolepidopteran larvae
(M. Shaw, 1983). Of the many
braconids attacking Lepidoptera, the
Rogadinae are unique in their habit
of mummifying the remains of the
host caterpillar, thus this group is
easy to recognize when reared from
host lepidopterans. The vast majority
of rogadine species are solitary
parasitoids, but a few gregarious
species are known (e.g. Aleiodes
stigmator). Pupation is internal,
within the shrunken and mummified
remains of the host caterpillar.

Sigalphinae
Members of the Sigalphinae are
koinobiont 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 a
carapace-like metasoma appears
somewhat contradicted by this
evidence.

Ypsistocerinae
All known species in two of the three
genera, viz. Ypsistocerus manni, Y.
vestigialis and Termitobracon
emersoni, have been recorded from
nests of Nasutitermes species.
Whether ypsistocerines are actually
parasitoids of the termites themselves
rather than of some group of
inquilines, is unknown.