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Scarabaeidae constitute approximately 27,000 species, of which approximately
5% or less in our estimation are cleptoparasitoids associated with deprivation of
food and other resources from social insects (Scholtz and Grebennikov 2005). Five
subfamilies of Scarabaeidae contain cleptoparasitoid taxa. Aphodiinae adults have
specialized associations, evidenced by distinctive morphological adaptations, with
termites and ants (Tangelder and Krikken 1982; Howden and Storey 1992; Stebnicka
1999). Scarabaeinae larvae, some known as “kleptocoprids”, breed in dung that
consume food resources provisioned by other, larger dung beetles (Halffter and
Edmonds 1982; Halffter and Matthews 1999). Dynastinae include larvae that feed
on brood and stored honey reserves in beehives (Glaser 1976; Evans and Nel 1989).
Cetoniinae consist of Cremastocheilini adults that abscond the food stores in termites, ant and bee nests (Deloya 1988; Alpert 1994). Valginae adults and larvae are
associated with termite mounds (Krikken 1978). Some Scarabaeidae (scarab beetles) are cleptoparasitoids that consume provisions of other scarab and closely
related geotrupid beetles (Hammond 1976; Halffter and Matthews 1999; Scholtz
and Grebennikov 2005). In the most common cleptoparasitoid group, the adult lays
eggs into dung balls or analogous brood structures that hatch, followed by the firstinstar larvae immediately burrowing and eventually consuming the host egg and all
provisions in a dung ball or analogous brood structure (Halffter and Edmonds 1982;
Paulian 1988). Some taxa of the scarabaeid subfamilies Valginae and Cetoniinae
(Cremastocheilini) that are associates of social insect nests probably are cleptoparasitoids (Krikken 1978; Alpert 1994; Scholtz and Chown 1995). The earliest documented Scarabaeidae is latest Middle Jurassic in age (Bai et al. 2012), although
parasitoid forms more likely are no older than mid Cretaceous in age based on their
position within Scarabaeoidea phylogeny (Krell 2006).
Rhipiceridae (cedar beetles, cicada parasite beetles), constitute 7 genera and 100
species that are globally distributed, all of which are ectoparasitoids on cicadas
(Lawrence 2005). Female rhipicerids such as Sandalus oviposit eggs in the cracks
and interstices of elm tree bark, where cicadas also have oviposited (Elzinga 1977).
Later, triungulin-like larvae and nymphs are flushed by rain and redeposited on the
ground, where, after some time, a rhipicerid late-instar ectoparasitoid larva becomes
attached to the cicada nymph host (Evans and Steury 2012). In eastern North
America, there appears to be annual tracking by the rhipicerid parasite of its cicada
host based on fluctuating population levels. The earliest occurrence of rhipicerid
beetles in the fossil record is considerably recent, the Lutetian Stage of the middle
Eocene (Ponomarenko 1995).
Cleridae (checkered beetles) consist of approximately 3400 species, are mostly
predaceous (Kolibác 2010), but have two major parasitoid strategies. Their first is a
role as cleptoparasitoids of apid bees, depriving larvae of nest provisions in soil
cells and decaying plant material. The first instar larva is host searching or phoretic
and feeds only a short time on the host’s provisions (Eggleton and Belshaw 1992).
Clerids have a second role as parasitoids of lepidopteran caterpillars occurring in
galls that occur on plant shoots, although the host locating process is poorly understood (Eggleton and Belshaw 1992). The origin of the parasitoid life habit in
Cleridae likely was derived from predation of insects in dead wood habitats. By
C. C. Labandeira and L. Li
Scarabaeidae constitute approximately 27,000 species, of which approximately
5% or less in our estimation are cleptoparasitoids associated with deprivation of
food and other resources from social insects (Scholtz and Grebennikov 2005). Five
subfamilies of Scarabaeidae contain cleptoparasitoid taxa. Aphodiinae adults have
specialized associations, evidenced by distinctive morphological adaptations, with
termites and ants (Tangelder and Krikken 1982; Howden and Storey 1992; Stebnicka
1999). Scarabaeinae larvae, some known as “kleptocoprids”, breed in dung that
consume food resources provisioned by other, larger dung beetles (Halffter and
Edmonds 1982; Halffter and Matthews 1999). Dynastinae include larvae that feed
on brood and stored honey reserves in beehives (Glaser 1976; Evans and Nel 1989).
Cetoniinae consist of Cremastocheilini adults that abscond the food stores in termites, ant and bee nests (Deloya 1988; Alpert 1994). Valginae adults and larvae are
associated with termite mounds (Krikken 1978). Some Scarabaeidae (scarab beetles) are cleptoparasitoids that consume provisions of other scarab and closely
related geotrupid beetles (Hammond 1976; Halffter and Matthews 1999; Scholtz
and Grebennikov 2005). In the most common cleptoparasitoid group, the adult lays
eggs into dung balls or analogous brood structures that hatch, followed by the firstinstar larvae immediately burrowing and eventually consuming the host egg and all
provisions in a dung ball or analogous brood structure (Halffter and Edmonds 1982;
Paulian 1988). Some taxa of the scarabaeid subfamilies Valginae and Cetoniinae
(Cremastocheilini) that are associates of social insect nests probably are cleptoparasitoids (Krikken 1978; Alpert 1994; Scholtz and Chown 1995). The earliest documented Scarabaeidae is latest Middle Jurassic in age (Bai et al. 2012), although
parasitoid forms more likely are no older than mid Cretaceous in age based on their
position within Scarabaeoidea phylogeny (Krell 2006).
Rhipiceridae (cedar beetles, cicada parasite beetles), constitute 7 genera and 100
species that are globally distributed, all of which are ectoparasitoids on cicadas
(Lawrence 2005). Female rhipicerids such as Sandalus oviposit eggs in the cracks
and interstices of elm tree bark, where cicadas also have oviposited (Elzinga 1977).
Later, triungulin-like larvae and nymphs are flushed by rain and redeposited on the
ground, where, after some time, a rhipicerid late-instar ectoparasitoid larva becomes
attached to the cicada nymph host (Evans and Steury 2012). In eastern North
America, there appears to be annual tracking by the rhipicerid parasite of its cicada
host based on fluctuating population levels. The earliest occurrence of rhipicerid
beetles in the fossil record is considerably recent, the Lutetian Stage of the middle
Eocene (Ponomarenko 1995).
Cleridae (checkered beetles) consist of approximately 3400 species, are mostly
predaceous (Kolibác 2010), but have two major parasitoid strategies. Their first is a
role as cleptoparasitoids of apid bees, depriving larvae of nest provisions in soil
cells and decaying plant material. The first instar larva is host searching or phoretic
and feeds only a short time on the host’s provisions (Eggleton and Belshaw 1992).
Clerids have a second role as parasitoids of lepidopteran caterpillars occurring in
galls that occur on plant shoots, although the host locating process is poorly understood (Eggleton and Belshaw 1992). The origin of the parasitoid life habit in
Cleridae likely was derived from predation of insects in dead wood habitats. By
C. C. Labandeira and L. Li