332
were found associated with other arthropods. Three main clades within Mesostigmata
can be recognised: Sejida, Trigynaspida and Monogynaspida. For a recent overview
of the mesostigmatid fossil record see Dunlop et al. (2018: Table 1). There are
reports of Cretaceous mesostigmatids in amber (Poinar Jr 1998; Peñalver et al.
2017), but none of them have been formally assigned to a family. Of the three main
clades, sejids are predators, the deutonymphs are often phoretic on beetles, but parasitism has not been recorded. Parasitic species are found within the two remaining
subgroups, albeit somewhat sporadically in five of the superfamilies listed below:
1. The superfamily Antennophoroidea in the cohort Antennophorina contains the
only parasites within the Trigynaspida clade. Members of the genus
Antennophorus (Antennophoridae) are associated with ants and induce the host
to regurgitate food for them (Franks et al. 1991). This may actually be an example of kleptoparasitism rather than parasitism per se as the host is not attacked
directly. There is no fossil record.
2. Uropodoidea in the cohort Uropodina are a monogynaspid group well known for
their phoretic behaviours (e.g. Faasch and Schaller 1966; see also Sect. 9.5.2)
and their association with other arthropods. At least one species of the genus
Macrodinychus (Macrodinychidae) is a parasitoid of ants which feeds on the
ants’ pupal stage. The wider Uropodoidea lineage dates back to at least
Palaeogene (ca. 44–49 Ma) Baltic amber, but these are phoretic fossils rather
than parasites.
3. Heterozerconidae, the only family in Heterozerconoidea and the cohort
Heterozerconina, are unusual mites associated with millipedes and occasionally
reptiles. They have a pair of sucker-like discs for attaching themselves to their
host (Gerdeman and Alberti 2007). Relationships are thought to be mostly commensal, but at least one species is a haematophagous parasite of worm-lizards
(Amphisbaena) (Flechtmann and Johnston 1990). There is no fossil record.
4. Ascoidea in the cohort Gamasina is a diverse superfamily which includes a number of taxa that feed on fungi or pollen (the family Ameroseiidae) as well as
species with a more typical predatory lifestyle (e.g. Walter 1987). The genus
Proctolaelaps (Ascidae) includes a handful of species known (or suspected) to
be parasites of their insect hosts (Egan and Hunter 1975). Ascoidea as a group
are known from Eocene Baltic amber, but again the record here appears to be
phoretic rather than parasitic.
5. Phytoseiodea is a gamasine superfamily which includes several predatory taxa,
some of which have potential as biological control agents against, e.g., spider
mites. One family (Otopheidomenidae) is exclusively parasitic on insects (e.g.
Zhang 1995) with different subgroups feeding on moths, true bugs, crickets or
termites. There is no fossil record.
The majority of the parasitic mesostigmatids occur in the final gamasine superfamily, Dermanyssoidea, which is thus considered in more detail below. The original lifestyle was probably free-living, as is still the case in some Laelapidae. The
remaining dermanyssoid mites are, or are thought to be, either blood-feeding ectoparasites, or endoparasites of the respiratory tract and parasitism probably arose on
J. A. Dunlop
were found associated with other arthropods. Three main clades within Mesostigmata
can be recognised: Sejida, Trigynaspida and Monogynaspida. For a recent overview
of the mesostigmatid fossil record see Dunlop et al. (2018: Table 1). There are
reports of Cretaceous mesostigmatids in amber (Poinar Jr 1998; Peñalver et al.
2017), but none of them have been formally assigned to a family. Of the three main
clades, sejids are predators, the deutonymphs are often phoretic on beetles, but parasitism has not been recorded. Parasitic species are found within the two remaining
subgroups, albeit somewhat sporadically in five of the superfamilies listed below:
1. The superfamily Antennophoroidea in the cohort Antennophorina contains the
only parasites within the Trigynaspida clade. Members of the genus
Antennophorus (Antennophoridae) are associated with ants and induce the host
to regurgitate food for them (Franks et al. 1991). This may actually be an example of kleptoparasitism rather than parasitism per se as the host is not attacked
directly. There is no fossil record.
2. Uropodoidea in the cohort Uropodina are a monogynaspid group well known for
their phoretic behaviours (e.g. Faasch and Schaller 1966; see also Sect. 9.5.2)
and their association with other arthropods. At least one species of the genus
Macrodinychus (Macrodinychidae) is a parasitoid of ants which feeds on the
ants’ pupal stage. The wider Uropodoidea lineage dates back to at least
Palaeogene (ca. 44–49 Ma) Baltic amber, but these are phoretic fossils rather
than parasites.
3. Heterozerconidae, the only family in Heterozerconoidea and the cohort
Heterozerconina, are unusual mites associated with millipedes and occasionally
reptiles. They have a pair of sucker-like discs for attaching themselves to their
host (Gerdeman and Alberti 2007). Relationships are thought to be mostly commensal, but at least one species is a haematophagous parasite of worm-lizards
(Amphisbaena) (Flechtmann and Johnston 1990). There is no fossil record.
4. Ascoidea in the cohort Gamasina is a diverse superfamily which includes a number of taxa that feed on fungi or pollen (the family Ameroseiidae) as well as
species with a more typical predatory lifestyle (e.g. Walter 1987). The genus
Proctolaelaps (Ascidae) includes a handful of species known (or suspected) to
be parasites of their insect hosts (Egan and Hunter 1975). Ascoidea as a group
are known from Eocene Baltic amber, but again the record here appears to be
phoretic rather than parasitic.
5. Phytoseiodea is a gamasine superfamily which includes several predatory taxa,
some of which have potential as biological control agents against, e.g., spider
mites. One family (Otopheidomenidae) is exclusively parasitic on insects (e.g.
Zhang 1995) with different subgroups feeding on moths, true bugs, crickets or
termites. There is no fossil record.
The majority of the parasitic mesostigmatids occur in the final gamasine superfamily, Dermanyssoidea, which is thus considered in more detail below. The original lifestyle was probably free-living, as is still the case in some Laelapidae. The
remaining dermanyssoid mites are, or are thought to be, either blood-feeding ectoparasites, or endoparasites of the respiratory tract and parasitism probably arose on
J. A. Dunlop