These are tiny wasps (1 mm or less) all of which are egg
parasites. One genus, Megaphragma, which parasitizes thrips eggs,
has species 180 microns (0.18 mm) in length. These are among the smallest
known insects (Doutt and Viggiani 1968). The genus Trichogramma has
probably had more papers written about it than any other chalcidoid, which
is especially regrettable when considering that few, if any, of these papers
have used a correct species name and therefore are of dubious value. A notable
exception is more recent work by Pinto, Platner, and Oatman (1978). George
Salt (1935, 1937, 1940) published a series of papers on the behavior of
Trichogramma and other parasites. More recently, Pinto, et al. (1989)
and Southamer, et al. (1990) have investigated important biological characteristics
such as phenotypic plasticity and thelytokous forms (respectively). The
genera of Trichogrammatidae were monographed at a world level by Doutt and
Viggiani (1968) who also provided lists of all known species. In 1971 Viggiani
presented a key to 27 (of 70) genera based on male genitalia. Yousuf and
Shaffee (1986) gave a checklist and bibliography of world species. Pinto
(in Gibson, et al. 1997) keyed all the genera for North America.
Of special interest to students of chalcidoid taxonomy is the great difference
between the genus Trichogramma and the genus Aphytis (Aphelinidae).
Both have been studied fairly well, from a biosystematic standpoint, with
completely different results. For example, Aphytis species are believed
to be very host specific, with about a 1 host, 1 parasite ratio (Rosen and
DeBach 1979). Trichogramma, however, tends to show "habitat
specificity" in some species, while others seem to show little specificity
of any kind (Nagarkatti and Nagaraja 1977). Salt (1935) reviewed records
of 1 species of Trichogramma ovipositing and emerging from eggs of
53 species of Lepidoptera, 6 Diptera, 3 Coleoptera, and Hymenoptera, Hemiptera,
and Neuroptera. In fact, Salt showed that a species of Trichogramma
attempted oviposition into any object of a certain configuration including
sand, glass beads, lumps of flour, seeds, and mercury globules. Given a
choice between large sand particles or seeds and a true host egg, Trichogramma
actually preferred the false host to the true one! Salt concluded that the
Trichogramma female was more impressed with "larger size"
than with actual host suitability and that "size is the principal criterion
used by Trichogramma in its choice of hosts." A second difference
is that the taxonomy of Aphytis and most other chalcidoids is based
on female specimens and the male genitalia have had little taxonomic significance.
Trichogramma taxonomy, however, is based largely on male genitalia.
From the 2 examples cited above it would be safe to say that each family
and probably each genus of chalcidoids needs to be studied on its own merits
as well as in comparison to our overall knowledge of Chalcidoidea.
Several authors (Doutt and Viggiani 1968, Nagarkatti and Nagaraja 1977)
stated that trichogrammatids are related to eulophids, and in fact, it seems
likely that one of the major groups of eulophids (possibly euderines or
tetrastichines) gave rise to this group. However they are distinct morphologically
and are not readily confused with eulophids. In addition, recent molecular
work could change our ideas of relationships. Viggiani (1971) divided the
family into 2 subfamilies based upon male genitalia (Trichogrammatinae,
Oligosetinae). Because of the difficulty in seeing adult trichogrammatids
(let alone their genitalia), their relative uncommonness (at least in general
collecting), and the degree of specialization required to work with them,
we will treat these parasites only at the family level.
STATISTICS: Number of world species: about 650 (50
Nearctic); number of world genera: 76 (35 Nearctic).
BIOLOGY: These wasps are strictly egg parasites and have
been reared from the holometabolus orders as well as Homoptera, Heteroptera,
Orthoptera, and Thysanoptera. This family also contains some of the few
truly aquatic species of Hymenoptera known. Several species in several genera
parasitize eggs of Odonata, Dytiscidae, and other aquatic Heteroptera. Propulsion
underwater may be by swimming with the legs or with the wings depending
upon the species. As many as 70 individuals may emerge from a single dytiscid
egg.
As was mentioned previously, Salt wrote several papers on the ethology of
Trichogramma (Clausen 1940 and Askew 1971 list most of these papers).
In addition to the example cited above for host selection, Salt has reported
on at least 2 other aspects of Trichogramma biology of interest to
the study of both taxonomy and biocontrol. These aspects concern the effect
that a host has upon its parasite. The first finding was that hosts of T.
semblidis caused male sexual dimorphism (much as in Melittobia-Eulophidae).
Males reared from Sialis (Neuroptera) eggs were apterous and had
different numbers of antennal segments and enlarged legs compared to winged
males reared from Lepidoptera. Females were the same from both hosts (Salt
1937). Of particular interest is that males did not differ in size and that
the differences in morphology were truly dimorphic, there being no intermediates.
Sialis was the "normal" host, but Salt suggested that a
"variety of hosts" might be attacked in the summer. The second
finding of interest (Salt 1940) was the fact that progeny from large hosts
generally were larger in size than from small hosts and that this could
effect both morphology and behavior. Morphologically it could result in
presence or absence of wings and the relative numbers of setae on wings
and antennae being correlated to length. Biologically he found that progeny
reared from large hosts behaved differently than progeny from small hosts.
For example, large progeny parasitized hosts nearly twice as fast as small
progeny, they possibly lived longer (though there was little data to make
this statement), and they had a higher reproductive potential. Also large
progeny could attack large hosts (which they preferred) or small hosts (reluctantly)
but small progeny could successfully attack only small hosts even though
they also preferred large ones.
DISTINGUISHING CHARACTERS: This family is distinct from
all other chalcidoids by having 3-segmented tarsi. Specimens are nearly
all less than 1 mm and are very fragile so that they shrivel up when dead
(similar to eulophids and some aphelinids in that respect). The antenna
contains 2 or less funicle segments (i.e. antenna appearing 5-segmented
or less with the club counted as one). Quite often the wing setae are arranged
in rows.
COLLECTING: Usually overlooked when sweeping because of
small size, these wasps are fairly common, but in small numbers. Sweeping
meadows and herbaceous vegetation is often productive, however, collecting
and rearing eggs, especially Lepidoptera, will usually produce the best
results.
DISTRIBUTION: Trichogrammatids are distributed worldwide
and are part of the "usual" fauna of small, remote islands by
virtue of their easy dispersal by wind (Doutt and Viggiani 1968).