Nuptial feeding encompasses any form of nutrient transfer from
the male to the female during or directly after
courtship and/or copulation. In insects, nuptial gifts may take the
form of food captured or collected by the
male, parts, or even the whole of the male's body, or glandular
products of the male such as salivary
secretions, external glandular secretions, the spermatophore and
substances in the ejaculate. Over the past
decade, there has been considerable debate over the current function
of nuptial feeding in insects. This
debate has centred on the issue of whether nuptial gifts function as
paternal investment (i.e. function to
increase the fitness and/or number of the gift-giving male's
own offspring) or as mating effort (i.e. function
to attract females, facilitate coupling, and/or to maximize
ejaculate transfer), although the two hypotheses
are not mutually exclusive. In the present article, evidence for the
potential of nuptial gifts to function as
either paternal investment, mating effort, or both is reviewed for
each form of nuptial feeding in each insect
taxon for which sufficient data are available. Empirical evidence suggests
that many diverse forms of nuptial
feeding in different insect taxa function, at least in part, as
mating effort. For example, nuptial prey and
salivary masses in the Mecoptera, regurgitated food in Drosophila
(Diptera),
hind-wing feeding in Cyphoderris
(Orthoptera) and the secretion of the male's cephalic gland in
Neopyrochroa (Coleoptera) and Zorotypus
(Zoraptera) appear to function to entice females to copulate and/or
to
facilitate coupling. Nuptial prey and
salivary masses in the Mecoptera also appear to function to maximize
ejaculate transfer (which is also a form
of mating effort), as do nuptial prey in Empis (Diptera),
external glandular secretions in Oecanthus and
Allonemobius (Orthoptera) and the spermatophylax in gryllids and
tettigoniids (Orthoptera). Large
spermatophores in, for example, the Lepidoptera and Coleoptera, also
appear to be maintained by selection
on the male to maximize ejaculate transfer and thereby counter the effects
of sperm competition. In contrast
to the large amount of evidence in support of the mating effort hypothesis,
there is a relative lack of good
evidence to support the paternal investment hypothesis. Certain studies
have demonstrated an increase in
the weight and/or number of eggs laid as a result of the receipt of
larger gifts, or a greater number of gifts,
in tettigoniids, gryllids, acridids, mantids, bruchid beetles, drosophilids
and lepidopterans. However,
virtually all of these studies (with the possible exception of studies
of the spermatophylax in tettigoniids) have
failed to control adequately for hormonal substances in the ejaculate
that are known to affect female
reproductive output. Furthermore, in at least four tettigoniids (but
not in the case of two species), three
lepidopterans, a drosophilid and probably also bruchid beetles and
bittacids, evidence suggests that the male
has a low probability of fertilising the eggs that stand to benefit
from his nuptial gift nutrients. Therefore,
the hypothesis that paternal investment might account for the function
of nuptial gifts in general is not supported.