Biology:Foster's rule

Short description: Ecogeographical rule in evolutionary biology

Garganornis ballmanni, a very large fossil goose from the Gargano and Scontrone islands of the Late Miocene

Foster's rule, also known as the island rule or the island effect, is an ecogeographical rule in evolutionary biology stating that members of a species get smaller or bigger depending on the resources available in the environment. For example, it is known that pygmy mammoths evolved from normal mammoths on small islands. Similar evolutionary paths have been observed in elephants, hippopotamuses, boas, sloths, deer (such as Key deer) and humans.^[1]^[2] It is part of the more general phenomenon of island syndrome which describes the differences in morphology, ecology, physiology and behaviour of insular species compared to their continental counterparts.

The rule was first formulated by van Valen in 1973^[3]^[4] based on the study by mammalogist J. Bristol Foster in 1964.^[5]^[6] In it, Foster compared 116 island species to their mainland varieties. Foster proposed that certain island creatures evolved larger body size (insular gigantism) while others became smaller (insular dwarfism). Foster proposed the simple explanation that smaller creatures get larger when predation pressure is relaxed because of the absence of some of the predators of the mainland, and larger creatures become smaller when food resources are limited because of land area constraints.^[7]

The idea was expanded upon in The Theory of Island Biogeography, by Robert MacArthur and Edward O. Wilson. In 1978, Ted J. Case published a longer paper on the topic in the journal Ecology.^[8]

Recent literature has also applied the island rule to plants.^[9]

There are some cases that do not neatly fit the rule; for example, artiodactyls have on several islands evolved into both dwarf and giant forms.^[10]^[11]

The Island Rule is a contested topic in evolutionary biology. Some argue that, since body size is a trait that is affected by multiple factors, and not just by organisms moving to an island, genetic variations across all populations could also cause the body mass differences between mainland and island populations.^[12]

References

↑ Juan Luis Arsuaga, Andy Klatt, The Neanderthal's Necklace: In Search of the First Thinkers, Thunder's Mouth Press, 2004, ISBN:1-56858-303-6, ISBN:978-1-56858-303-7, p. 199.
↑ Jean-Baptiste de Panafieu, Patrick Gries, Evolution, Seven Stories Press, 2007, ISBN:1-58322-784-9, ISBN:978-1-58322-784-8, p 42.
↑ Van Valen, L. (1973). "Body Size and Numbers of Plants and Animals". Evolution 27 (1): 27–35. doi:10.2307/2407116. PMID 28563673.
↑ Lokatis, S.; Jeschke, J.M. (2018). "The island rule: An assessment of biases and research trends". Journal of Biogeography 45 (2): 289–303. doi:10.1111/jbi.13160.
↑ Foster, J.B. (1964). "The evolution of mammals on islands". Nature 202 (4929): 234–235. doi:10.1038/202234a0. Bibcode: 1964Natur.202..234F.
↑ Foster, J. B. (1965) The evolution of the mammals of the Queen Charlotte Islands, British Columbia. Occasional Papers of the British Columbia Provincial Museum, 14, 1–130.
↑ Whittaker, R.J. (1998). Island biogeography: ecology, evolution, and conservation. Oxford University Press, UK. pp. 73–75. ISBN 978-0-19-850020-9.
↑ Case, T.J. (1978). "A general explanation for insular body size trends in terrestrial vertebrates". Ecology 59 (1): 1–18. doi:10.2307/1936628.
↑ Biddick, M.; Hendriks, A.; Burns, K.C. (2019-08-19). "Plants obey (and disobey) the island rule". Proceedings of the National Academy of Sciences 116 (36): 17632–34. doi:10.1073/pnas.1907424116. PMID 31427521. Bibcode: 2019PNAS..11617632B.
↑ Mazza, P.P.A.; Rossi, M.A.; Agostini, S. (2015). "Hoplitomerycidæ (Late Miocene, Italy), an Example of Giantism in Insular Ruminants". Journal of Mammalian Evolution 22 (2): 271–77. doi:10.1007/s10914-014-9277-2.
↑ van der Geer, A.A.E. (2018). "Uniformity in variety: Antler morphology and evolution in a predator-free environment". Palaeontologia Electronica (21.1.9A): 1–31. doi:10.26879/834.
↑ Lokatis, Sophie; Jeschke, Jonathan M. (2018). "The island rule: An assessment of biases and research trends". Journal of Biogeography 45 (2): 289–303. doi:10.1111/jbi.13160.

External links

"Why Do Islands Breed Giants (And Sometimes Dwarfs)?", episode of PBS's Nova

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[1] Juan Luis Arsuaga, Andy Klatt, The Neanderthal's Necklace: In Search of the First Thinkers, Thunder's Mouth Press, 2004, ISBN:1-56858-303-6, ISBN:978-1-56858-303-7, p. 199.

[2] Jean-Baptiste de Panafieu, Patrick Gries, Evolution, Seven Stories Press, 2007, ISBN:1-58322-784-9, ISBN:978-1-58322-784-8, p 42.

[Valen1973-3] Van Valen, L. (1973). "Body Size and Numbers of Plants and Animals". Evolution 27 (1): 27–35. doi:10.2307/2407116. PMID 28563673.

[Lokatis2018-4] Lokatis, S.; Jeschke, J.M. (2018). "The island rule: An assessment of biases and research trends". Journal of Biogeography 45 (2): 289–303. doi:10.1111/jbi.13160.

[5] Foster, J.B. (1964). "The evolution of mammals on islands". Nature 202 (4929): 234–235. doi:10.1038/202234a0. Bibcode: 1964Natur.202..234F.

[6] Foster, J. B. (1965) The evolution of the mammals of the Queen Charlotte Islands, British Columbia. Occasional Papers of the British Columbia Provincial Museum, 14, 1–130.

[7] Whittaker, R.J. (1998). Island biogeography: ecology, evolution, and conservation. Oxford University Press, UK. pp. 73–75. ISBN 978-0-19-850020-9.

[8] Case, T.J. (1978). "A general explanation for insular body size trends in terrestrial vertebrates". Ecology 59 (1): 1–18. doi:10.2307/1936628.

[9] Biddick, M.; Hendriks, A.; Burns, K.C. (2019-08-19). "Plants obey (and disobey) the island rule". Proceedings of the National Academy of Sciences 116 (36): 17632–34. doi:10.1073/pnas.1907424116. PMID 31427521. Bibcode: 2019PNAS..11617632B.

[Mazza2015-10] Mazza, P.P.A.; Rossi, M.A.; Agostini, S. (2015). "Hoplitomerycidæ (Late Miocene, Italy), an Example of Giantism in Insular Ruminants". Journal of Mammalian Evolution 22 (2): 271–77. doi:10.1007/s10914-014-9277-2.

[Geer2018-11] van der Geer, A.A.E. (2018). "Uniformity in variety: Antler morphology and evolution in a predator-free environment". Palaeontologia Electronica (21.1.9A): 1–31. doi:10.26879/834.

[12] Lokatis, Sophie; Jeschke, Jonathan M. (2018). "The island rule: An assessment of biases and research trends". Journal of Biogeography 45 (2): 289–303. doi:10.1111/jbi.13160.

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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 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 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

v t e Ecology: Modelling ecosystems: Trophic components
General	Abiotic component Abiotic stress Behaviour Biogeochemical cycle Biomass Biotic component Biotic stress Carrying capacity Competition Ecosystem Ecosystem ecology Ecosystem model Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Resource
Producers	Autotrophs Chemosynthesis Chemotrophs Foundation species Mixotrophs Myco-heterotrophy Mycotroph Organotrophs Photoheterotrophs Photosynthesis Photosynthetic efficiency Phototrophs Primary nutritional groups Primary production
Consumers	Apex predator Bacterivore Carnivores Chemoorganotroph Foraging Generalist and specialist species Intraguild predation Herbivores Heterotroph Heterotrophic nutrition Insectivore Mesopredators Mesopredator release hypothesis Omnivores Optimal foraging theory Planktivore Predation Prey switching
Decomposers	Chemoorganoheterotrophy Decomposition Detritivores Detritus
Microorganisms	Archaea Bacteriophage Environmental microbiology Lithoautotroph Lithotrophy Microbial cooperation Microbial ecology Microbial food web Microbial intelligence Microbial loop Microbial mat Microbial metabolism Phage ecology
Food webs	Biomagnification Ecological efficiency Ecological pyramid Energy flow Food chain Trophic level
Example webs	Cold seeps Hydrothermal vents Intertidal Kelp forests Lakes North Pacific Subtropical Gyre Rivers San Francisco Estuary Soil Tide pool
Processes	Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer-resource systems Copiotrophs Dominance Ecological network Ecological succession Energy quality Energy Systems Language f-ratio Feed conversion ratio Feeding frenzy Mesotrophic soil Nutrient cycle Oligotroph Paradox of the plankton Trophic cascade Trophic mutualism Trophic state index
Defense, counter	Animal coloration Antipredator adaptations Camouflage Deimatic behaviour Herbivore adaptations to plant defense Mimicry Plant defense against herbivory Predator avoidance in schooling fish

v t e Ecology: Modelling ecosystems: Other components
Population ecology	Abundance Allee effect Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation in wild animals Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey (Lotka–Volterra) equations Recruitment Resilience Small population size Stability
Species	Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species Latitudinal gradients in species diversity Minimum viable population Neutral theory Occupancy–abundance relationship Population viability analysis Priority effect Rapoport's rule Relative abundance distribution Relative species abundance Species diversity Species homogeneity Species richness Species distribution Species-area curve Umbrella species
Species interaction	Antibiosis Biological interaction Commensalism Community ecology Ecological facilitation Interspecific competition Mutualism Parasitism Storage effect Symbiosis
Spatial ecology	Biogeography Cross-boundary subsidy Ecocline Ecotone Ecotype Disturbance Edge effects Foster's rule Habitat fragmentation Ideal free distribution Intermediate Disturbance Hypothesis Island biogeography Landscape ecology Landscape epidemiology Landscape limnology Metapopulation Patch dynamics r/K selection theory Resource selection function Source–sink dynamics
Niche	Ecological niche Ecological trap Ecosystem engineer Environmental niche modelling Guild Habitat Marine habitats Limiting similarity Niche apportionment models Niche construction Niche differentiation
Other networks	Assembly rules Bateman's principle Bioluminescence Ecological collapse Ecological debt Ecological deficit Ecological energetics Ecological indicator Ecological threshold Ecosystem diversity Emergence Extinction debt Kleiber's law Liebig's law of the minimum Marginal value theorem Thorson's rule Xerosere
Other	Allometry Alternative stable state Balance of nature Biological data visualization Ecocline Ecological economics Ecological footprint Ecological forecasting Ecological humanities Ecological stoichiometry Ecopath Ecosystem based fisheries Endolith Evolutionary ecology Functional ecology Industrial ecology Macroecology Microecosystem Natural environment Regime shift Systems ecology Urban ecology Theoretical ecology
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