393
11.3.7 Coprolites
Arthropod (Fig. 11.1a) and vertebrate (Fig. 11.1c) coprolites often contain identifiable material assignable to particular insect predators and the hosts of parasites and
parasitoids, but are rare as fossils (Labandeira 2002; Robin et al. 2016; Chin 2021).
Coprolites of hosts are usually difficult to assign to a producer; nevertheless, they
potentially are a significant archive of fossilized trophic relationships that include
evidence for parasitism (De Baets and Littlewood 2015; Qvarnström et al. 2016).
Such coprolites occasionally reveal prey items consumed by predators such as swallowed insect parasites combed from hair and feathers by vertebrate grooming
behavior (Qvarnström et al. 2016). More remotely, parasitoid remains can occur in
the coprolites of small terrestrial vertebrates. However, for insects during their 412
million-year-long existence, by far the greatest populations of insect coprolites
involve plant cuticle, trichomes and vascular tissue remnants from detritivores and
herbivores, and pollen from pollinivores (Labandeira and Phillips 1996; Labandeira
1998). Fluid-feeding parasites and parasitoids would rarely produce a detectible
coprolite record.
11.3.8 Sedimentary Ichnological Evidence
Sedimentary ichnological evidence provides support for recognizing inquilinism,
cleptoparasitism, parasitoidism and scavenging in the fossil record, particularly in
the trace fossil record of Pleistocene and Holocene deposits that involve beetle,
wasp and bee hosts (Ellis and Ellis-Adam 1993; Bown et al. 1997; Mikulás and
Genise 2003; Genise and Cladera 2004). Cleptoparasitism, for example, occurs in
the sedimentary ichnological record as fossil traces of Tombownichnus pepei that
consist of pits in the walls or infillings of Coprinosphaera, the brood balls of beetles
(Sánchez and Genise 2009). The Tombownichnus–Coprinosphaera parasitoid and
host relationship has modern equivalents (Halffter and Edmonds 1982; Halffter and
Matthews 1999), although the material documented from the middle Eocene–lower
Miocene Saramiento Formation of Patagonia, Argentina (Sánchez and Genise 2009)
indicates an ancient association. The cleptoparasite culprits remain unknown,
although the Tombownichnus pepei pit type likely represents pupation chambers
excavated by last-instar larvae.
Another example is T. plenus from Semnan Province, Iran, that involves curvilinear, narrow burrows up to 8 mm long that invade bee colony cells (Bagheri et al.
2013). These bee nest structures in Celliforma nests are attributed to one of the three
main groups of bee parasitoids: Meloidae (blister beetles), Bombyliidae (bee flies)
or Mutillidae (velvet ants). Similarly, possible parasitoid attack of bee cells are present in ichnofossils of Celliforma, attributed to modern Anthophoridae (carpenter
bees), from Fuerteventura and Lanzarote of the Canary Islands, Spain. The outer
sedimentary linings of individual unopened cells exhibit numerous perforations that
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
11.3.7 Coprolites
Arthropod (Fig. 11.1a) and vertebrate (Fig. 11.1c) coprolites often contain identifiable material assignable to particular insect predators and the hosts of parasites and
parasitoids, but are rare as fossils (Labandeira 2002; Robin et al. 2016; Chin 2021).
Coprolites of hosts are usually difficult to assign to a producer; nevertheless, they
potentially are a significant archive of fossilized trophic relationships that include
evidence for parasitism (De Baets and Littlewood 2015; Qvarnström et al. 2016).
Such coprolites occasionally reveal prey items consumed by predators such as swallowed insect parasites combed from hair and feathers by vertebrate grooming
behavior (Qvarnström et al. 2016). More remotely, parasitoid remains can occur in
the coprolites of small terrestrial vertebrates. However, for insects during their 412
million-year-long existence, by far the greatest populations of insect coprolites
involve plant cuticle, trichomes and vascular tissue remnants from detritivores and
herbivores, and pollen from pollinivores (Labandeira and Phillips 1996; Labandeira
1998). Fluid-feeding parasites and parasitoids would rarely produce a detectible
coprolite record.
11.3.8 Sedimentary Ichnological Evidence
Sedimentary ichnological evidence provides support for recognizing inquilinism,
cleptoparasitism, parasitoidism and scavenging in the fossil record, particularly in
the trace fossil record of Pleistocene and Holocene deposits that involve beetle,
wasp and bee hosts (Ellis and Ellis-Adam 1993; Bown et al. 1997; Mikulás and
Genise 2003; Genise and Cladera 2004). Cleptoparasitism, for example, occurs in
the sedimentary ichnological record as fossil traces of Tombownichnus pepei that
consist of pits in the walls or infillings of Coprinosphaera, the brood balls of beetles
(Sánchez and Genise 2009). The Tombownichnus–Coprinosphaera parasitoid and
host relationship has modern equivalents (Halffter and Edmonds 1982; Halffter and
Matthews 1999), although the material documented from the middle Eocene–lower
Miocene Saramiento Formation of Patagonia, Argentina (Sánchez and Genise 2009)
indicates an ancient association. The cleptoparasite culprits remain unknown,
although the Tombownichnus pepei pit type likely represents pupation chambers
excavated by last-instar larvae.
Another example is T. plenus from Semnan Province, Iran, that involves curvilinear, narrow burrows up to 8 mm long that invade bee colony cells (Bagheri et al.
2013). These bee nest structures in Celliforma nests are attributed to one of the three
main groups of bee parasitoids: Meloidae (blister beetles), Bombyliidae (bee flies)
or Mutillidae (velvet ants). Similarly, possible parasitoid attack of bee cells are present in ichnofossils of Celliforma, attributed to modern Anthophoridae (carpenter
bees), from Fuerteventura and Lanzarote of the Canary Islands, Spain. The outer
sedimentary linings of individual unopened cells exhibit numerous perforations that
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution