Emery's rule

Emery's rule is the trend of social parasites to be parasites to species or genera they are closely related to.

History

In 1909, the Italian entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.^[1]^[2]

Over time, this pattern has been recognized in many additional cases, and generalized to what is now known as Emery's rule.^[1]

In nature

The pattern is best known for various taxa of Hymenoptera. For example, the social wasp Dolichovespula adulterina parasitizes other members of its genus such as Dolichovespula norwegica and Dolichovespula arenaria.^[3]^[4] Emery's rule is also applicable to members of other kingdoms such as fungi, red algae, and mistletoe. The significance and general relevance of this pattern are still a matter of some debate, as a great many exceptions exist, though a common explanation for the phenomenon when it occurs is that the parasites may have started as facultative parasites within the host species itself (such forms of intraspecific parasitism are well-known, even in some species of bees),^[5] but later became reproductively isolated and split off from the ancestral species, a form of sympatric speciation.

When a parasitic species is a sister taxon to its host in a phylogenetic sense, the relationship is considered to be in "strict" adherence to Emery's rule. When the parasite is a close relative of the host but not its sister species, the relationship is in "loose" adherence to the rule.^[6]

References

^ ^a ^b Richard Deslippe (2010). "Social Parasitism in Ants". Nature Education Knowledge. Retrieved 2010-10-29. In 1909, the taxonomist Carlo Emery made an important generalization, now known as Emery's rule, which states that social parasites and their hosts share common ancestry and hence are closely related to each other (Emery 1909).
^ Emery, C. "Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen". Biologisches Centralblatt 29, 352–362 (1909)
^ Carpenter, J. M., & Perera, E. P. (2006). Phylogenetic relationships among yellowjackets and the evolution of social parasitism (Hymenoptera: Vespidae, Vespinae). American Museum Novitates, 1–19.
^ Dvořák, L. (2007). Parasitism of Dolichovespula norwegica by D. adulterina (Hymenoptera: Vespidae). Archived 2016-03-04 at the Wayback Machine Silva Gabreta, 13(1) 65–67.
^ Wenseleers, Tom (2011). "Intraspecific queen parasitism in highly eusocial bee" (PDF). Biology Letters. 7 (2). Royal Society Publishing: 173–176. doi:10.1098/rsbl.2010.0819. PMC 3061179. PMID 20961883. Retrieved 9 July 2011.
^ Hines, H. M., & Cameron, S. A. (2010). The phylogenetic position of the bumble bee inquiline Bombus inexspectatus and implications for the evolution of social parasitism. Insectes Sociaux, 57(4), 379–383.

[RD-1] Richard Deslippe (2010). "Social Parasitism in Ants". Nature Education Knowledge. Retrieved 2010-10-29. In 1909, the taxonomist Carlo Emery made an important generalization, now known as Emery's rule, which states that social parasites and their hosts share common ancestry and hence are closely related to each other (Emery 1909).

[Emery_1909-2] Emery, C. "Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen". Biologisches Centralblatt 29, 352–362 (1909)

[3] Carpenter, J. M., & Perera, E. P. (2006). Phylogenetic relationships among yellowjackets and the evolution of social parasitism (Hymenoptera: Vespidae, Vespinae). American Museum Novitates, 1–19.

[4] Dvořák, L. (2007). Parasitism of Dolichovespula norwegica by D. adulterina (Hymenoptera: Vespidae). Archived 2016-03-04 at the Wayback Machine Silva Gabreta, 13(1) 65–67.

[5] Wenseleers, Tom (2011). "Intraspecific queen parasitism in highly eusocial bee" (PDF). Biology Letters. 7 (2). Royal Society Publishing: 173–176. doi:10.1098/rsbl.2010.0819. PMC 3061179. PMID 20961883. Retrieved 9 July 2011.

[hines-6] Hines, H. M., & Cameron, S. A. (2010). The phylogenetic position of the bumble bee inquiline Bombus inexspectatus and implications for the evolution of social parasitism. Insectes Sociaux, 57(4), 379–383.

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v t e Biological rules
Rules	Allen's rule Shorter appendages in colder climates Bateson's rule Extra limbs mirror their neighbours Bergmann's rule Larger bodies in colder climates Cope's rule Bodies get larger over time Deep-sea gigantism Larger bodies in deep-sea animals Dollo's law Loss of complex traits is irreversible Eichler's rule Parasites co-vary with their hosts Emery's rule Insect social parasites are often in same genus as their hosts Fahrenholz's rule Host and parasite phylogenies become congruent Foster's rule (Insular gigantism, Insular dwarfism) Small species get larger, large species smaller, after colonizing islands Gause's law Complete competitors cannot coexist Gloger's rule Lighter coloration in colder, drier climates Haldane's rule Hybrid sexes that are absent, rare, or sterile, are heterogamic Harrison's rule Parasites co-vary in size with their hosts Hamilton's rule Genes increase in frequency when relatedness of recipient to actor times benefit to recipient exceeds reproductive cost to actor Kleiber's law An animals metabolic rate decreases with its size Hennig's progression rule In cladistics, the most primitive species are found in earliest, central, part of group's area Jarman–Bell principle The correlation between the size of an animal and its diet quality; larger animals can consume lower quality diet Jordan's rule Inverse relationship between water temperature and no. of fin rays, vertebrae Lack's principle Birds lay only as many eggs as they can provide food for Rapoport's rule Latitudinal range increases with latitude Rensch's rule Sexual size dimorphism increases with size when males are larger, decreases with size when females are larger Rosa's rule Groups evolve from character variation in primitive species to a fixed character state in advanced ones Schmalhausen's law A population at limit of tolerance in one aspect is vulnerable to small differences in any other aspect Thayer's law The top of an animals coloration is darker than the bottom Thorson's rule No. of eggs of benthic marine invertebrates decreases with latitude Van Valen's law Probability of extinction of a group is constant over time von Baer's laws Embryos start from a common form and develop into increasingly specialised forms Williston's law Parts in an organism become reduced in number and specialized in function
Related	Countergradient variation Where genetics opposes environment as a factor Gigantothermy Large ectothermic animals more easily maintain constant body temperature