388
(Greenwalt et al. 2013; Greenwalt 2021), that allows for assignment of these insect
fossils, respectively, to parasitoid and parasite life habits. The evidence is highly
variable and consists of modern and fossil molecular data, broadly construed
(Nagler and Haug 2015), but also includes morphological and interaction-related
features of the fossil organisms themselves or their taxonomic affiliations (Fig. 11.1f,
h, j–l) (De Baets and Littlewood 2015; Leung 2017). Functional features also are
important (Fig. 11.1b, e, f, h, j, l), as are damage to organisms (Fig. 11.1m, n), gut
contents (Fig. 11.1h), coprolites (Fig. 11.1a, c), and predator avoidance traits
(Fig. 11.1d, g) (Labandeira 2002). Evidence of parasitism is seen in Fig. 11.1e, h, k,
l; evidence of parasitoidism is seen in Fig. 11.1f, n.
11.3.1 Biomolecular Data
Within the deep-time context of the insect fossil record, study of biomolecules has
two basic applications for understanding the occurrence of predation and especially
parasitism and parasitoidism. The first approach involves understanding of the feeding habits and other host relationships of the fossil parasite containing the ingested
biomolecule such as blood (Greenwalt 2021; Greenwalt et al. 2013; Yao et al. 2014).
In a complementary sense, identification of the ingested biomolecule itself is necessary for specification of the host taxon (De Baets and Littlewood 2015). Identification
of both parasite and host is still is in their infancy (De Baets and Littlewood 2015);
but characterization of certain biomolecules, such as fossil keratin, shows considerable promise (Wappler et al. 2004; Briggs and Summons 2014).
The other major use of biomolecular data relevant for inferring predation, parasitism and parasitoidism in the fossil record is the production of phylogenies (De
Baets and Littlewood 2015; Nagler and Haug 2015; Warnock and Engelstädter
2021). The influence of molecular phylogenies of insects—particularly those of
fossil-calibrated cladistic studies of predator, parasite and parasitoid groups—has
provided considerable assistance in inferring the predatory, parasitic or parasitoidic
life habits of many insect lineages (Pohl and Beutel 2005; Winterton et al. 2007;
Bologna et al. 2008; Heraty et al. 2013; Winkler et al. 2015; Peters et al. 2017;
Gillung et al. 2018). In addition, such studies are a principal basis for understanding
the inter-relationships among predator and especially parasite and parasitoid lineages and their relationships to sister clades (Grimaldi and Engel 2005a).
11.3.2 Taxonomic Affiliation
The insect fossil record consists of evidence for predatory, parasitic and parasitoidic
interactions based on taxonomic similarity to their recent relatives (Boucot and
Poinar Jr 2010; Leung 2017). For example, bee cleptoparasitism was found in the
fossil record of a late Paleocene (60 Ma) site from Menat, France (Martins et al.
C. C. Labandeira and L. Li
(Greenwalt et al. 2013; Greenwalt 2021), that allows for assignment of these insect
fossils, respectively, to parasitoid and parasite life habits. The evidence is highly
variable and consists of modern and fossil molecular data, broadly construed
(Nagler and Haug 2015), but also includes morphological and interaction-related
features of the fossil organisms themselves or their taxonomic affiliations (Fig. 11.1f,
h, j–l) (De Baets and Littlewood 2015; Leung 2017). Functional features also are
important (Fig. 11.1b, e, f, h, j, l), as are damage to organisms (Fig. 11.1m, n), gut
contents (Fig. 11.1h), coprolites (Fig. 11.1a, c), and predator avoidance traits
(Fig. 11.1d, g) (Labandeira 2002). Evidence of parasitism is seen in Fig. 11.1e, h, k,
l; evidence of parasitoidism is seen in Fig. 11.1f, n.
11.3.1 Biomolecular Data
Within the deep-time context of the insect fossil record, study of biomolecules has
two basic applications for understanding the occurrence of predation and especially
parasitism and parasitoidism. The first approach involves understanding of the feeding habits and other host relationships of the fossil parasite containing the ingested
biomolecule such as blood (Greenwalt 2021; Greenwalt et al. 2013; Yao et al. 2014).
In a complementary sense, identification of the ingested biomolecule itself is necessary for specification of the host taxon (De Baets and Littlewood 2015). Identification
of both parasite and host is still is in their infancy (De Baets and Littlewood 2015);
but characterization of certain biomolecules, such as fossil keratin, shows considerable promise (Wappler et al. 2004; Briggs and Summons 2014).
The other major use of biomolecular data relevant for inferring predation, parasitism and parasitoidism in the fossil record is the production of phylogenies (De
Baets and Littlewood 2015; Nagler and Haug 2015; Warnock and Engelstädter
2021). The influence of molecular phylogenies of insects—particularly those of
fossil-calibrated cladistic studies of predator, parasite and parasitoid groups—has
provided considerable assistance in inferring the predatory, parasitic or parasitoidic
life habits of many insect lineages (Pohl and Beutel 2005; Winterton et al. 2007;
Bologna et al. 2008; Heraty et al. 2013; Winkler et al. 2015; Peters et al. 2017;
Gillung et al. 2018). In addition, such studies are a principal basis for understanding
the inter-relationships among predator and especially parasite and parasitoid lineages and their relationships to sister clades (Grimaldi and Engel 2005a).
11.3.2 Taxonomic Affiliation
The insect fossil record consists of evidence for predatory, parasitic and parasitoidic
interactions based on taxonomic similarity to their recent relatives (Boucot and
Poinar Jr 2010; Leung 2017). For example, bee cleptoparasitism was found in the
fossil record of a late Paleocene (60 Ma) site from Menat, France (Martins et al.
C. C. Labandeira and L. Li