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fungi in punky wood, although a broad spectrum of feeding habits are present, including parasitism (Newton 2005). One subfamily, Platypsyllinae, consists of 4 genera
and about 20 species that are ectoparasites of semiaquatic and aquatic mammals such
as insectivores (Talpidae), beavers (Castoridae), mountain beavers (Aplodontidae)
and river otters (Mustelidae), consuming flakes of decorticated skin, dermal exudates
and probably surficial blood (Newton 2005; Peck 2006). The highly specialized parasite Platypsyllus (beaver beetle) is the best documented platypsylline, and the adult,
flea-like in overall appearance, is a flat, wingless and blind form that has thoracic
hooks for attachment to the hair of its host. The larvae and adults are parasites of the
North American beaver, Castor canadensis, and the North American river otter,
Lontra canadensis, and leave their host only to pupate (Peck 2006).
Meloidae (blister beetles) are a diverse, cosmopolitan family of beetles occurring
primarily in warm temperate to arid regions that consist of four subfamilies and
about 120 genera (Bologna et al. 2010). The diets of adult Meloidae are variable,
with phytophagy dominant and predation subdominant. Most larvae are predaceous,
feeding on grasshopper egg pods and immatures of aculeate wasps and bees, with
development occurring in the soil or wood (Bologna et al. 2010). However, parasite
and parasitoid forms occur throughout the family. The life cycle of Meloidae is
complex, with the larva undergoing hypermetamorphosis and consisting of four
distinct larval instars prior to entering the pupal stage. The first larval instar is an
actively mobile, well sclerotized triungulin, which is the dispersal stage for seeking
parasite and parasitoid hosts. The triungulin instar is followed by a first instar grub,
a feeding stage of robust proportions; then a coarctate instar, a resting stage in diapause; and finally a second grub instar that is active but nonfeeding prior to pupation
(Selander and Mathieu 1964). Parasitic (and parasitoidic) larvae use grasshoppers,
ground beetles, aculeate wasps and bees as hosts (Engel 2005a; Bologna et al.
2010). Although meloid consumers of other animals are over-represented by parasitoids, the family does harbor several lineages of ectoparasites (Engel 2005a;
Bologna and Di Giulio 2011).
The fossil record of beetle parasites, as would be expected, is poor, particularly
for non-diverse families. There is no fossil record of the Platypsyllinae, although the
Leiodidae has a decent fossil record of 16 occurrences that extends to the Late
Jurassic (Perkovsky 1999). Rhipiceridae apparently has a fossil record extending to
the middle Eocene (Ponomarenko 1995). Bothrideridae has a known first occurrence from the middle Eocene (Ponomarenko 1995). The Passandridae has a sparse
fossil record of three Cenozoic occurrences; the earliest occurrence is from an
approximately 44 million-year-old middle Eocene (Lutetian) deposit in Argentina
(Ramírez et al. 2016). The Meloidae is a largely Cenozoic lineage that has a sparse
fossil record when compared to its extant diversity. The meloid fossil record, however, includes records of triungulin larvae (Engel 2005a; Bologna et al. 2008) that
provide evidence for phoresy and other features consistent with an ectoparasitic
mode of life (Engel 2005a). The meloid fossil record includes 12 fossil occurrences;
the oldest is from the 35 million-year-old late Eocene. The Ripiphoridae have a fossil record extending to the early Cenomanian (Cai et al. 2018; Batelka et al. 2016,
2019). It is unclear if any of these occurrences demonstrate a specific parasitic relationship with an actual, extinct host.
C. C. Labandeira and L. Li
fungi in punky wood, although a broad spectrum of feeding habits are present, including parasitism (Newton 2005). One subfamily, Platypsyllinae, consists of 4 genera
and about 20 species that are ectoparasites of semiaquatic and aquatic mammals such
as insectivores (Talpidae), beavers (Castoridae), mountain beavers (Aplodontidae)
and river otters (Mustelidae), consuming flakes of decorticated skin, dermal exudates
and probably surficial blood (Newton 2005; Peck 2006). The highly specialized parasite Platypsyllus (beaver beetle) is the best documented platypsylline, and the adult,
flea-like in overall appearance, is a flat, wingless and blind form that has thoracic
hooks for attachment to the hair of its host. The larvae and adults are parasites of the
North American beaver, Castor canadensis, and the North American river otter,
Lontra canadensis, and leave their host only to pupate (Peck 2006).
Meloidae (blister beetles) are a diverse, cosmopolitan family of beetles occurring
primarily in warm temperate to arid regions that consist of four subfamilies and
about 120 genera (Bologna et al. 2010). The diets of adult Meloidae are variable,
with phytophagy dominant and predation subdominant. Most larvae are predaceous,
feeding on grasshopper egg pods and immatures of aculeate wasps and bees, with
development occurring in the soil or wood (Bologna et al. 2010). However, parasite
and parasitoid forms occur throughout the family. The life cycle of Meloidae is
complex, with the larva undergoing hypermetamorphosis and consisting of four
distinct larval instars prior to entering the pupal stage. The first larval instar is an
actively mobile, well sclerotized triungulin, which is the dispersal stage for seeking
parasite and parasitoid hosts. The triungulin instar is followed by a first instar grub,
a feeding stage of robust proportions; then a coarctate instar, a resting stage in diapause; and finally a second grub instar that is active but nonfeeding prior to pupation
(Selander and Mathieu 1964). Parasitic (and parasitoidic) larvae use grasshoppers,
ground beetles, aculeate wasps and bees as hosts (Engel 2005a; Bologna et al.
2010). Although meloid consumers of other animals are over-represented by parasitoids, the family does harbor several lineages of ectoparasites (Engel 2005a;
Bologna and Di Giulio 2011).
The fossil record of beetle parasites, as would be expected, is poor, particularly
for non-diverse families. There is no fossil record of the Platypsyllinae, although the
Leiodidae has a decent fossil record of 16 occurrences that extends to the Late
Jurassic (Perkovsky 1999). Rhipiceridae apparently has a fossil record extending to
the middle Eocene (Ponomarenko 1995). Bothrideridae has a known first occurrence from the middle Eocene (Ponomarenko 1995). The Passandridae has a sparse
fossil record of three Cenozoic occurrences; the earliest occurrence is from an
approximately 44 million-year-old middle Eocene (Lutetian) deposit in Argentina
(Ramírez et al. 2016). The Meloidae is a largely Cenozoic lineage that has a sparse
fossil record when compared to its extant diversity. The meloid fossil record, however, includes records of triungulin larvae (Engel 2005a; Bologna et al. 2008) that
provide evidence for phoresy and other features consistent with an ectoparasitic
mode of life (Engel 2005a). The meloid fossil record includes 12 fossil occurrences;
the oldest is from the 35 million-year-old late Eocene. The Ripiphoridae have a fossil record extending to the early Cenomanian (Cai et al. 2018; Batelka et al. 2016,
2019). It is unclear if any of these occurrences demonstrate a specific parasitic relationship with an actual, extinct host.
C. C. Labandeira and L. Li