447
Tribe Calpini of the Subfamily Calpinae (Zaspel et al. 2012). The clade of Calyptra
consist of ten species of hematophagous vampire moths in which evidently only
males imbibe blood from large mammals that include cattle, zebu, tapirs and occasionally elephants and humans, centered in Southeast Asia but with a geographic
range extending to India and southeastern Russia (Bänziger 1971; Zaspel et al.
2007). Related species and genera within Calpini frequent the more delicate membranes of the eyes, rectum and genitalia for feeding on tears, sebum, sweat and other
dermal secretions (Bänziger 1968, 1996). A more distantly related clade of fruit
piercing and surface scarifying moths are known from the tropics of Africa and
South America (Zenker et al. 2010). The mouthpart siphons of these fruit feeding,
and blood- and lymph feeding species, are rigid, lack internal stylets, and contain
external, spine-like structures at the proboscis tip that are hydraulically stiffened for
penetration into skin. The siphonal stylet is powered by a cibarial food pump for
ingestion of fluid host tissue (Bänziger 1996; Labandeira 2019). The clade that
bears these distinctive mouthparts lacks a fossil record.
11.5.1.10 Hymenoptera (Wasps)
Social parasitism frequently is found in Hymenoptera and is particularly prevalent
among bees. A recurring feature of social parasitism is embodied in Emery’s Rule,
which is narrowly defined as social parasites are the closest relatives of their hosts,
frequently at the sister-species level (Wilson 1971). A broader version of this rule
states that a clade or related group of social parasites is solely related to their hosts
(Smith et al. 2007). An illustration of this phenomenon occurs in allodapine carpenter bees (Apidae: Xylocopinae) (Smith et al. 2007), which exhibit a looser version
of relationships between a host clade and its sister social parasite clade (Fig. 11.11),
providing evidence for sympatric speciation. As seen in the lack of rigorous correlation of sympatry in Fig. 11.11, there may be inherent flaws in the notion of Emery’s
Rule, at least in some instances. Confounding processes such as extinction or speciation within one or both of the clades, host-switching, and incomplete sampling
(Smith et al. 2007) could derail the observed patterns. Notably, older host–social
parasite relationships tend to support sympatric speciation whereas younger such
relationships do not. This discontinuity likely is because extinctions may result in a
pruning of the older clades such that they appear retrospectively to have originated
sympatrically (Smith et al. 2007).
A second type of common parasitic relationship among bees is cleptoparasitism.
Cleptoparasitism, the theft of resources by one animal from another animal, is rarely
found in the fossil record. However, based on a distinctive forewing shape and other
morphometrically-gleaned features, it was possible to assign a cleptoparasite identity to a wing of a bee from the middle Paleocene, at the 60.5 million-year-old
Menat locality in central France (Dehon et al. 2017). The presence of cleptoparasite
taxa is significant, as it establishes that before the midpoint of the geochronologic
history of bees, cleptoparasites were present.
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
Tribe Calpini of the Subfamily Calpinae (Zaspel et al. 2012). The clade of Calyptra
consist of ten species of hematophagous vampire moths in which evidently only
males imbibe blood from large mammals that include cattle, zebu, tapirs and occasionally elephants and humans, centered in Southeast Asia but with a geographic
range extending to India and southeastern Russia (Bänziger 1971; Zaspel et al.
2007). Related species and genera within Calpini frequent the more delicate membranes of the eyes, rectum and genitalia for feeding on tears, sebum, sweat and other
dermal secretions (Bänziger 1968, 1996). A more distantly related clade of fruit
piercing and surface scarifying moths are known from the tropics of Africa and
South America (Zenker et al. 2010). The mouthpart siphons of these fruit feeding,
and blood- and lymph feeding species, are rigid, lack internal stylets, and contain
external, spine-like structures at the proboscis tip that are hydraulically stiffened for
penetration into skin. The siphonal stylet is powered by a cibarial food pump for
ingestion of fluid host tissue (Bänziger 1996; Labandeira 2019). The clade that
bears these distinctive mouthparts lacks a fossil record.
11.5.1.10 Hymenoptera (Wasps)
Social parasitism frequently is found in Hymenoptera and is particularly prevalent
among bees. A recurring feature of social parasitism is embodied in Emery’s Rule,
which is narrowly defined as social parasites are the closest relatives of their hosts,
frequently at the sister-species level (Wilson 1971). A broader version of this rule
states that a clade or related group of social parasites is solely related to their hosts
(Smith et al. 2007). An illustration of this phenomenon occurs in allodapine carpenter bees (Apidae: Xylocopinae) (Smith et al. 2007), which exhibit a looser version
of relationships between a host clade and its sister social parasite clade (Fig. 11.11),
providing evidence for sympatric speciation. As seen in the lack of rigorous correlation of sympatry in Fig. 11.11, there may be inherent flaws in the notion of Emery’s
Rule, at least in some instances. Confounding processes such as extinction or speciation within one or both of the clades, host-switching, and incomplete sampling
(Smith et al. 2007) could derail the observed patterns. Notably, older host–social
parasite relationships tend to support sympatric speciation whereas younger such
relationships do not. This discontinuity likely is because extinctions may result in a
pruning of the older clades such that they appear retrospectively to have originated
sympatrically (Smith et al. 2007).
A second type of common parasitic relationship among bees is cleptoparasitism.
Cleptoparasitism, the theft of resources by one animal from another animal, is rarely
found in the fossil record. However, based on a distinctive forewing shape and other
morphometrically-gleaned features, it was possible to assign a cleptoparasite identity to a wing of a bee from the middle Paleocene, at the 60.5 million-year-old
Menat locality in central France (Dehon et al. 2017). The presence of cleptoparasite
taxa is significant, as it establishes that before the midpoint of the geochronologic
history of bees, cleptoparasites were present.
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution