386
An example is a social parasite that benefits from brood care and access to resources
that are managed by and benefits the host colony. Social parasitism occurs among a
broad variety of insects in which the hosts and the parasites often are closely related,
in some cases forming a sympatric species pair. By contrast, inquilines are animals
that exploit the living space, such as a termite or ant nest, resulting in a commensal
relationship in which one member benefits and the other member derives a reward
or is not harmed (Malyshev 1968). Inquilism often is a precursor to social parasitism, which in turn may be a precursor to cleptoparasitoidism (Askew 1971).
11.3 The Evidence: Distinguishing Predation, Parasitism
and Parasitoidism
Several types of evidence are important for detecting the presence of predation,
parasitism and parasitoidism in the fossil record (Fig. 11.1). Some of this evidence
is associated with particular techniques, such as synchrotron X-ray microtomography (van de Camp et al. 2018), or time-of flight secondary-ion mass spectrometry
Fig. 11.1 (continued) New York State (Shear et al. 1989). (c) Vertebrate coprolite containing
remains of a cockroach, including wings (w), legs (l) and ovipositor (o) from the Middle
Pennsylvanian Mazon Creek locality of Illinois (FMNH PE 54114). (d) From the same locality as
(c) is the insect Protdiamphipnoa woodwardi Brongniart (Cnemidolestodea: Cnemidolestidae),
with prominent forewing eyespots (Carpenter 1971). (e) The enigmatic flea or flea-like insect
Strashila incredibilis Rasnitsyn (?Siphonaptera: Saurophthiridae), from the Late Jurassic of
Transbaikalia, Russia, exhibiting aptery, chelate hind tarsus and piercing-and-sucking mouthparts
(Rasnitsyn 1992). (f) The parasitoid ensign wasp Leptephialtites caudatus Rasnitsyn (Hymenoptera:
Ephialtitidae) from the Late Jurassic of Karatau, Kazakhstan, with elongate ovipositor (o) and
ovipositor valves (ov) (Rasnitsyn 1975). (g) The digger wasp Angarosphex beiboziensis Hong
(Hymenoptera: Angarosphecidae) from the Jurassic–Cretaceous boundary of China, exhibiting
abdominal banding indicating a Batesian model (Hong 1984). (h) The peculiar chewing louse
Saurodectes vrsanskyi Rasnitsyn & Zherikhin (?Phthiraptera: Saurodectidae) from the Early
Cretaceous of Transbaikalia, Russia (Rasnitsyn and Zherikhin 1999) Note gut contents. (i) Gut
contents of the middle Eocene bat Palaeochiropteryx tupaiodon Revilliod (Chiroptera:
Palaeochiropterygidae) from Messel, Germany, containing butterfly scales and other insect fragments (Richter and Storch 1980). Approximate length of scale at right is 90 μm. (j) Robber fly
(Diptera: Asilidae) from the middle Eocene of Colorado, USA (USNM501477), displaying raptorial forelegs and mouthparts of a single, dagger-like stylet (arrow). (k) The egg (nit) of an undetermined sucking louse (Phthiraptera: Anoplura) on a mammalian hair shaft, from middle Eocene
Baltic amber of Germany (Voigt 1952). (l) Palaeopsylla klebsiana Dampf (Siphonaptera:
Hystrichopsyllidae), a mammal-parasitizing flea from the same provenance as (k), showing socketed antennae, head comb and maxillary lever–stylet complex typical of fleas (Dampf 1910). (m)
The seed Tectocarya rhenana (Cornaceae), with exit hole of a seed predator, from the early
Miocene of Germany (Schmidt et al. 1958). (n) A bee cell (Hymenoptera: Stenotritidae) from the
Pleistocene of South Australia, with a small exit hole of a probable parasitoid (Houston 1987). All
subfigures are redraw from original images or are camera lucida drawings of specimens from the
Field Museum of Natural History in Chicago (FMNH) or the National Museum of Natural History
in Washington, DC (USNM). Scale bars: solid, 1.0 cm; striped, 0.1 cm. (Reproduced, with permission of the Paleontological Society, from Fig. 2 of Labandeira 2002)
C. C. Labandeira and L. Li
An example is a social parasite that benefits from brood care and access to resources
that are managed by and benefits the host colony. Social parasitism occurs among a
broad variety of insects in which the hosts and the parasites often are closely related,
in some cases forming a sympatric species pair. By contrast, inquilines are animals
that exploit the living space, such as a termite or ant nest, resulting in a commensal
relationship in which one member benefits and the other member derives a reward
or is not harmed (Malyshev 1968). Inquilism often is a precursor to social parasitism, which in turn may be a precursor to cleptoparasitoidism (Askew 1971).
11.3 The Evidence: Distinguishing Predation, Parasitism
and Parasitoidism
Several types of evidence are important for detecting the presence of predation,
parasitism and parasitoidism in the fossil record (Fig. 11.1). Some of this evidence
is associated with particular techniques, such as synchrotron X-ray microtomography (van de Camp et al. 2018), or time-of flight secondary-ion mass spectrometry
Fig. 11.1 (continued) New York State (Shear et al. 1989). (c) Vertebrate coprolite containing
remains of a cockroach, including wings (w), legs (l) and ovipositor (o) from the Middle
Pennsylvanian Mazon Creek locality of Illinois (FMNH PE 54114). (d) From the same locality as
(c) is the insect Protdiamphipnoa woodwardi Brongniart (Cnemidolestodea: Cnemidolestidae),
with prominent forewing eyespots (Carpenter 1971). (e) The enigmatic flea or flea-like insect
Strashila incredibilis Rasnitsyn (?Siphonaptera: Saurophthiridae), from the Late Jurassic of
Transbaikalia, Russia, exhibiting aptery, chelate hind tarsus and piercing-and-sucking mouthparts
(Rasnitsyn 1992). (f) The parasitoid ensign wasp Leptephialtites caudatus Rasnitsyn (Hymenoptera:
Ephialtitidae) from the Late Jurassic of Karatau, Kazakhstan, with elongate ovipositor (o) and
ovipositor valves (ov) (Rasnitsyn 1975). (g) The digger wasp Angarosphex beiboziensis Hong
(Hymenoptera: Angarosphecidae) from the Jurassic–Cretaceous boundary of China, exhibiting
abdominal banding indicating a Batesian model (Hong 1984). (h) The peculiar chewing louse
Saurodectes vrsanskyi Rasnitsyn & Zherikhin (?Phthiraptera: Saurodectidae) from the Early
Cretaceous of Transbaikalia, Russia (Rasnitsyn and Zherikhin 1999) Note gut contents. (i) Gut
contents of the middle Eocene bat Palaeochiropteryx tupaiodon Revilliod (Chiroptera:
Palaeochiropterygidae) from Messel, Germany, containing butterfly scales and other insect fragments (Richter and Storch 1980). Approximate length of scale at right is 90 μm. (j) Robber fly
(Diptera: Asilidae) from the middle Eocene of Colorado, USA (USNM501477), displaying raptorial forelegs and mouthparts of a single, dagger-like stylet (arrow). (k) The egg (nit) of an undetermined sucking louse (Phthiraptera: Anoplura) on a mammalian hair shaft, from middle Eocene
Baltic amber of Germany (Voigt 1952). (l) Palaeopsylla klebsiana Dampf (Siphonaptera:
Hystrichopsyllidae), a mammal-parasitizing flea from the same provenance as (k), showing socketed antennae, head comb and maxillary lever–stylet complex typical of fleas (Dampf 1910). (m)
The seed Tectocarya rhenana (Cornaceae), with exit hole of a seed predator, from the early
Miocene of Germany (Schmidt et al. 1958). (n) A bee cell (Hymenoptera: Stenotritidae) from the
Pleistocene of South Australia, with a small exit hole of a probable parasitoid (Houston 1987). All
subfigures are redraw from original images or are camera lucida drawings of specimens from the
Field Museum of Natural History in Chicago (FMNH) or the National Museum of Natural History
in Washington, DC (USNM). Scale bars: solid, 1.0 cm; striped, 0.1 cm. (Reproduced, with permission of the Paleontological Society, from Fig. 2 of Labandeira 2002)
C. C. Labandeira and L. Li