394
suggest considerable pre-imaginal mortality, attributed to predation or fungal
attacks (Ellis and Ellis-Adam 1993), but equally could represent parasitoid attack.
In another example, direct evidence of a parasitoid entry or an exit hole is provided
by a bee cell of Stenotritidae, from the Pleistocene of South Australia (Houston 1987).
Perforations of thickened bee cell walls and small cocoons within the bee cells
also have been described from the Paleocene–Eocene Claron Formation of southwestern Utah (Bown et al. 1997). Perforations of these cell walls are rounded,
incomplete or complete holes that suggest parasitoid presence, but of unknown
taxonomic affiliation. A second instance is the association of Lazaichnus fistulosus
with beetle pupal chambers, consisting of holes circular to broadly ellipsoidal in
outline that penetrate the cell wall and are connected to a single cavity of Monesichnus
ameghinoi (Mikulás and Genise 2003). This probable parasitoid trace fossil is from
the Late Cretaceous to Early Paleogene Ascencio Formation of Uruguay and represents the earliest occurrence of probable parasitoidism in the sedimentary ichnological record (Martin and Varricchio 2011).
11.4 Evolutionary and Ecological Biology
Parasitism, in the broadest sense of the term to include parasitoidism, is a very
ancient relationship extending into Paleozoic (Leung 2017) and likely even
Precambrian (De Baets and Littlewood 2015) time. The varieties of parasitism exist
as a continuum (Robin 2021) that ranges from symbiosis where two interacting
organisms mutually benefit, to commensalism where one organism benefits and the
other has a neutral interaction, and then to predation, parasitism and eventually
parasitoidism where one interacting organism benefits and the other is disadvantaged, including death (Araújo et al. 2003). For parasitism, one conservative estimate of the animal kingdom indicates that this life habit has originated at least 223
times in 15 phyla, with the greatest representation occurring overwhelmingly in
Arthropoda (Weinstein and Kuris 2016). Within arthropods, it is not necessarily the
most diverse clades that exhibit the greatest incidence of parasitism. Clearly, at the
family level, originations of parasitism in insects were overwhelmingly concentrated among the holometabolous insects where development included larval and
pupal instars (Poulin and Morand 2000). These originations consisted of 84 separate
events for holometabolous insects, compared with five originations for hemimetabolous insects, where larval and pupal instars are absent. Hemimetabolous insect
occurrences are Dermaptera (one occurrence), Psocoptera (one occurrence),
Phthiraptera (one occurrence, included in Psocodea) and Hemiptera (one occurrence) (Fig. 11.2). At the family level of analysis, the distribution of the incidence
of originations of parasitism is unrelated to underlying total species diversity
(Fig. 11.2). Coleoptera (beetles) are the most diverse clade, yet exhibit only ten
occurrences of parasitism. Diptera displays 60 separate occurrences of parasitism
(Feener and Brown 1997; Weinstein and Kuris 2016), whereas Hymenoptera (sawflies, wasps, ants and bees), which have a species diversity of 75% that of the Diptera
C. C. Labandeira and L. Li
suggest considerable pre-imaginal mortality, attributed to predation or fungal
attacks (Ellis and Ellis-Adam 1993), but equally could represent parasitoid attack.
In another example, direct evidence of a parasitoid entry or an exit hole is provided
by a bee cell of Stenotritidae, from the Pleistocene of South Australia (Houston 1987).
Perforations of thickened bee cell walls and small cocoons within the bee cells
also have been described from the Paleocene–Eocene Claron Formation of southwestern Utah (Bown et al. 1997). Perforations of these cell walls are rounded,
incomplete or complete holes that suggest parasitoid presence, but of unknown
taxonomic affiliation. A second instance is the association of Lazaichnus fistulosus
with beetle pupal chambers, consisting of holes circular to broadly ellipsoidal in
outline that penetrate the cell wall and are connected to a single cavity of Monesichnus
ameghinoi (Mikulás and Genise 2003). This probable parasitoid trace fossil is from
the Late Cretaceous to Early Paleogene Ascencio Formation of Uruguay and represents the earliest occurrence of probable parasitoidism in the sedimentary ichnological record (Martin and Varricchio 2011).
11.4 Evolutionary and Ecological Biology
Parasitism, in the broadest sense of the term to include parasitoidism, is a very
ancient relationship extending into Paleozoic (Leung 2017) and likely even
Precambrian (De Baets and Littlewood 2015) time. The varieties of parasitism exist
as a continuum (Robin 2021) that ranges from symbiosis where two interacting
organisms mutually benefit, to commensalism where one organism benefits and the
other has a neutral interaction, and then to predation, parasitism and eventually
parasitoidism where one interacting organism benefits and the other is disadvantaged, including death (Araújo et al. 2003). For parasitism, one conservative estimate of the animal kingdom indicates that this life habit has originated at least 223
times in 15 phyla, with the greatest representation occurring overwhelmingly in
Arthropoda (Weinstein and Kuris 2016). Within arthropods, it is not necessarily the
most diverse clades that exhibit the greatest incidence of parasitism. Clearly, at the
family level, originations of parasitism in insects were overwhelmingly concentrated among the holometabolous insects where development included larval and
pupal instars (Poulin and Morand 2000). These originations consisted of 84 separate
events for holometabolous insects, compared with five originations for hemimetabolous insects, where larval and pupal instars are absent. Hemimetabolous insect
occurrences are Dermaptera (one occurrence), Psocoptera (one occurrence),
Phthiraptera (one occurrence, included in Psocodea) and Hemiptera (one occurrence) (Fig. 11.2). At the family level of analysis, the distribution of the incidence
of originations of parasitism is unrelated to underlying total species diversity
(Fig. 11.2). Coleoptera (beetles) are the most diverse clade, yet exhibit only ten
occurrences of parasitism. Diptera displays 60 separate occurrences of parasitism
(Feener and Brown 1997; Weinstein and Kuris 2016), whereas Hymenoptera (sawflies, wasps, ants and bees), which have a species diversity of 75% that of the Diptera
C. C. Labandeira and L. Li