339
Gabryś et al. 2011). Several examples of putative parasitengonid mites attached
to spiders were documented from Baltic amber by Wohltmann in Wunderlich
(2004); see also above.
3. Mantid flies (Neuroptera: Mantispidae) have larvae that typically feed on spider
eggs. The larva either seeks an already available egg sac or attaches itself to a
spider and waits for the eggs to be laid. Whether this constitutes parasitism sensu
stricto is debatable—it represents egg parasitoidism if the egg is presumably
killed—but Ohl (2011) reported a remarkable case from Baltic amber of a mantispid larva attached to a clubionid spider (Fig. 9.7a). Adult mantid flies are
known as far back as the Jurassic (e.g. Jepson et al. 2013).
4. At least three families of wasps (Hymenoptera) are spider parasitoids, or at least
include species which attack spiders. Pompilidae (spider wasps) are dedicated
spider hunters, depositing an ectoparasitoid egg on the paralysed victim. The
fossil record of Pompilidae is subject to debate. A Cretaceous example was
described from Burmese amber by Engel and Grimaldi (2006), but Rodriguez
et al. (2016) questioned this interpretation and restricted the group to Eocene and
younger records.
5. Ichneumon wasps (Ichneumonidae) are parasitoids which usually attack the larvae of butterflies, beetles or other hymenopterans, but occasionally also spiders.
Again, the egg is lain on (or in) the host, which may or may not be paralysed, and
the host is eventually consumed alive. Fossil ichneumonids (Labandeira and Li
2021) can be traced back to the Cretaceous (e.g. McKellar et al. 2013). An apparent case of egg parasitism of a clubionid spider by an ichneumonid was documented in Eocene Baltic amber by Poinar Jr (2004). An ichneumonid larva,
again on a clubionid spider, was described from Miocene Dominican amber by
Poinar Jr (1987).
6. Some species of digger wasp (Crabronidae) attack spiders. The family as a whole
is known from the Cretaceous, but the spider-hunting Trypoxylini lineage is first
recorded from Eocene French amber (Nel 2005).
7. Spider flies (Diptera: Acroceridae) are parasitoids whose larva enters the host’s
body and often lodges near the book lung, before consuming the hosts tissues.
The oldest fossil examples come from the Jurassic of central Asia (Gillung and
Winterton 2017; Labandeira and Li 2021) and they have also been recorded from
Burmese, Baltic and Dominican ambers. Spiders are not the only victims, and
Kerr and Winterton (2008) described a remarkable case of a larval acrocerid
attached to a whirligig mite (Anystidae) in Baltic amber.
Acknowledgements I thank Kenneth de Baets for inviting this contribution and Lidia ChitimiaDobler, Enrique Peñalver, David Penney, Michael Ohl, George Poinar Jr., Ekaterina Sidochuk,
Dieter Waloszek and Michael Zwanzig for providing images of specimens. Kenneth de Baets, an
anonymous reviewer and Ekaterina Sidochuk provided valuable comments on the typescript.
Ekaterina died in January 2019, and I would like to dedicate this work to her memory for her
extensive contributions to our understanding of fossil mites.
9 Chelicerates as Parasites
Gabryś et al. 2011). Several examples of putative parasitengonid mites attached
to spiders were documented from Baltic amber by Wohltmann in Wunderlich
(2004); see also above.
3. Mantid flies (Neuroptera: Mantispidae) have larvae that typically feed on spider
eggs. The larva either seeks an already available egg sac or attaches itself to a
spider and waits for the eggs to be laid. Whether this constitutes parasitism sensu
stricto is debatable—it represents egg parasitoidism if the egg is presumably
killed—but Ohl (2011) reported a remarkable case from Baltic amber of a mantispid larva attached to a clubionid spider (Fig. 9.7a). Adult mantid flies are
known as far back as the Jurassic (e.g. Jepson et al. 2013).
4. At least three families of wasps (Hymenoptera) are spider parasitoids, or at least
include species which attack spiders. Pompilidae (spider wasps) are dedicated
spider hunters, depositing an ectoparasitoid egg on the paralysed victim. The
fossil record of Pompilidae is subject to debate. A Cretaceous example was
described from Burmese amber by Engel and Grimaldi (2006), but Rodriguez
et al. (2016) questioned this interpretation and restricted the group to Eocene and
younger records.
5. Ichneumon wasps (Ichneumonidae) are parasitoids which usually attack the larvae of butterflies, beetles or other hymenopterans, but occasionally also spiders.
Again, the egg is lain on (or in) the host, which may or may not be paralysed, and
the host is eventually consumed alive. Fossil ichneumonids (Labandeira and Li
2021) can be traced back to the Cretaceous (e.g. McKellar et al. 2013). An apparent case of egg parasitism of a clubionid spider by an ichneumonid was documented in Eocene Baltic amber by Poinar Jr (2004). An ichneumonid larva,
again on a clubionid spider, was described from Miocene Dominican amber by
Poinar Jr (1987).
6. Some species of digger wasp (Crabronidae) attack spiders. The family as a whole
is known from the Cretaceous, but the spider-hunting Trypoxylini lineage is first
recorded from Eocene French amber (Nel 2005).
7. Spider flies (Diptera: Acroceridae) are parasitoids whose larva enters the host’s
body and often lodges near the book lung, before consuming the hosts tissues.
The oldest fossil examples come from the Jurassic of central Asia (Gillung and
Winterton 2017; Labandeira and Li 2021) and they have also been recorded from
Burmese, Baltic and Dominican ambers. Spiders are not the only victims, and
Kerr and Winterton (2008) described a remarkable case of a larval acrocerid
attached to a whirligig mite (Anystidae) in Baltic amber.
Acknowledgements I thank Kenneth de Baets for inviting this contribution and Lidia ChitimiaDobler, Enrique Peñalver, David Penney, Michael Ohl, George Poinar Jr., Ekaterina Sidochuk,
Dieter Waloszek and Michael Zwanzig for providing images of specimens. Kenneth de Baets, an
anonymous reviewer and Ekaterina Sidochuk provided valuable comments on the typescript.
Ekaterina died in January 2019, and I would like to dedicate this work to her memory for her
extensive contributions to our understanding of fossil mites.
9 Chelicerates as Parasites