Rapoport's rule

Rapoport's rule is an ecogeographical rule that states that latitudinal ranges of plants and animals are generally smaller at lower latitudes than at higher latitudes.

Background

Stevens (1989)^[1] named the rule after Eduardo H. Rapoport, who had earlier provided evidence for the phenomenon for subspecies of mammals (Rapoport 1975,^[2] 1982^[3]). Stevens used the rule to "explain" greater species diversity in the tropics in the sense that latitudinal gradients in species diversity and the rule have identical exceptional data and so must have the same underlying cause. Narrower ranges in the tropics would facilitate more species to coexist. He later extended the rule to altitudinal gradients, claiming that altitudinal ranges are greatest at greater latitudes (Stevens 1992^[4]), and to depth gradients in the oceans (Stevens 1996^[5]). The rule has been the focus of intense discussion and given much impetus to exploring distributional patterns of plants and animals. Stevens' original paper has been cited about 330 times in the scientific literature.

Generality

Support for the generality of the rule is at best equivocal.^[6] For example, marine teleost fishes have the greatest latitudinal ranges at low latitudes.^[7]^[8] In contrast, freshwater fishes do show the trend, although only above a latitude of about 40 degrees North.^[8] Some subsequent papers have found support for the rule; others, probably even more numerous, have found exceptions to it.^[6]^[9] For most groups that have been shown to follow the rule, it is restricted to or at least most distinct above latitudes of about 40–50 degrees. Rohde therefore concluded that the rule describes a local phenomenon.^[10] Computer simulations using the Chowdhury Ecosystem Model did not find support for the rule.^[11]

Explanations

Rohde (1996)^[10] explained the fact that the rule is restricted to very high latitudes by effects of glaciations which have wiped out species with narrow ranges, a view also expressed by Brown (1995).^[12] Another explanation of Rapoport's rule is the "climatic variability" or "seasonal variability hypothesis".^[5]^[13] According to this hypothesis, seasonal variability selects for greater climatic tolerances and therefore wider latitudinal ranges (see also Fernandez and Vrba 2005^[14]).

Methods used to demonstrate the rule

The methods used to demonstrate the rule have been subject to some controversy. Most commonly, authors plot means of latitudinal ranges in a particular 5° latitudinal band against latitude, although modal or median ranges have been used by some.^[15] In the original paper by Stevens, all species occurring in each band were counted, i.e., a species with a range of 50 degrees occurs in 10 or 11 bands. However, this may lead to an artificial inflation of latitudinal ranges of species occurring at high latitudes, because even a few tropical species with wide ranges will affect the means of ranges at high latitudes, whereas the opposite effect due to high latitude species extending into the tropics is negligible: species diversity is much smaller at high than low latitudes. As an alternative method the "midpoint method" has been proposed, which avoids this problem. It counts only those species with the midpoint of their ranges in a particular latitudinal band.^[8] An additional complication in assessing Rapoport's rule for data based on field sampling is the possibility of a spurious pattern driven by a sample-size artifact. Equal sampling effort at species-rich and species-poor localities tends to underestimate range size at the richer localities relative to the poorer, when in fact range sizes might not differ among localities.^[16]

Biotic and abiotic factors which act against the rule

Marine benthic invertebrates and some parasites have been shown to have smaller dispersal abilities in cold seas (Thorson's rule), which would counteract Rapoport's rule. The tropics have far more uniform temperatures over a far wider latitudinal range (about 45 degrees) than high latitude species. As temperature is one of the most important (if not the most important) factor determining geographical distribution, wider latitudinal ranges in the tropics might therefore be expected.

Evolutionary age

The inconsistent results concerning Rapoport's rule suggest that certain characteristics of species may be responsible for their different latitudinal ranges. These characteristics may include, for example, their evolutionary age: species that have evolved recently in the tropics may have small latitudinal ranges because they have not had the time to spread far from their origin, whereas older species have extended their ranges.^[17]

References

^ Stevens, George C. (1989). "The Latitudinal Gradient in Geographical Range: How so Many Species Coexist in the Tropics". The American Naturalist. 133 (2): 240–256. doi:10.1086/284913.
^ Rapoport, E. H. (1975). Areografía. Estrategias Geográficas de las Especies. Fondo de Cultura Económica, México
^ Rapoport, E. H. (1982). Areography. Geographical Strategies of Species. Trad. B. Drausal, Pergamon Press, Oxford. ISBN 978-0-08-028914-4
^ Stevens, George C. (1992). "The Elevational Gradient in Altitudinal Range: An Extension of Rapoport's Latitudinal Rule to Altitude". The American Naturalist. 140 (6): 893–911. doi:10.1086/285447. PMID 19426029.
^ ^a ^b Stevens, George (1996). "Extending Rapoport's rule to Pacific marine fishes". Journal of Biogeography. 23 (2): 149–154. Bibcode:1996JBiog..23..149S. doi:10.1046/j.1365-2699.1996.00977.x.
^ ^a ^b Gaston, Kevin J.; Blackburn, Tim M.; Spicer, John I. (1998). "Rapoport's rule: Time for an epitaph?". Trends in Ecology & Evolution. 13 (2): 70–74. Bibcode:1998TEcoE..13...70G. doi:10.1016/S0169-5347(97)01236-6. PMID 21238203.
^ Rohde, Klaus (1992). "Latitudinal Gradients in Species Diversity: The Search for the Primary Cause". Oikos. 65 (3): 514–527. Bibcode:1992Oikos..65..514R. doi:10.2307/3545569. JSTOR 3545569.
^ ^a ^b ^c Rohde, Klaus; Heap, Maureen; Heap, David (1993). "Rapoport's Rule Does Not Apply to Marine Teleosts and Cannot Explain Latitudinal Gradients in Species Richness". The American Naturalist. 142 (1): 1–16. doi:10.1086/285526.
^ Rohde, Klaus (1999). "Latitudinal gradients in species diversity and Rapoport's rule revisited: a review of recent work and what can parasites teach us about the causes of the gradients?". Ecography. 22 (6): 593–613. Bibcode:1999Ecogr..22..593R. doi:10.1111/j.1600-0587.1999.tb00509.x.
^ ^a ^b Rohde, Klaus (1996). "Rapoport's Rule is a Local Phenomenon and Cannot Explain Latitudinal Gradients in Species Diversity". Biodiversity Letters. 3 (1): 10–13. doi:10.2307/2999704. JSTOR 2999704.
^ Stauffer, D.; Rohde, K. (2006). "Simulation of Rapoport's rule for latitudinal species spread". Theory in Biosciences. 125 (1): 55–65. arXiv:q-bio/0507033. Bibcode:2006ThBio.125...55S. doi:10.1016/j.thbio.2006.01.002. PMID 17046373.
^ Brown, J. H. (1995). Macroecology. Chicago: University of Chicago Press.
^ Letcher, Andy J.; Harvey, Paul H. (1994). "Variation in Geographical Range Size Among Mammals of the Palearctic". The American Naturalist. 144 (1): 30–42. doi:10.1086/285659.
^ Fernández, Manuel Hernández; Vrba, Elisabeth S. (2005). "Rapoport effect and biomic specialization in African mammals: revisiting the climatic variability hypothesis". Journal of Biogeography. 32 (5): 903–918. Bibcode:2005JBiog..32..903F. doi:10.1111/j.1365-2699.2004.01188.x.
^ Roy, K; Jablonski, D; Valentine, J W (1994). "Eastern Pacific molluscan provinces and latitudinal diversity gradient: no evidence for "Rapoport's rule"". Proceedings of the National Academy of Sciences. 91 (19): 8871–8874. Bibcode:1994PNAS...91.8871R. doi:10.1073/pnas.91.19.8871. PMC 44708. PMID 11607494.
^ Colwell, Robert K.; Hurtt, George C. (1994). "Nonbiological Gradients in Species Richness and a Spurious Rapoport Effect". The American Naturalist. 144 (4): 570–595. doi:10.1086/285695.
^ Rohde, Klaus (1998). "Latitudinal Gradients in Species Diversity. Area Matters, but How Much?". Oikos. 82 (1): 184–190. Bibcode:1998Oikos..82..184R. doi:10.2307/3546928. JSTOR 3546928.

External links

Klaus Rohde: Rapoport's rule

[1] Stevens, George C. (1989). "The Latitudinal Gradient in Geographical Range: How so Many Species Coexist in the Tropics". The American Naturalist. 133 (2): 240–256. doi:10.1086/284913.

[2] Rapoport, E. H. (1975). Areografía. Estrategias Geográficas de las Especies. Fondo de Cultura Económica, México

[3] Rapoport, E. H. (1982). Areography. Geographical Strategies of Species. Trad. B. Drausal, Pergamon Press, Oxford. ISBN 978-0-08-028914-4

[4] Stevens, George C. (1992). "The Elevational Gradient in Altitudinal Range: An Extension of Rapoport's Latitudinal Rule to Altitude". The American Naturalist. 140 (6): 893–911. doi:10.1086/285447. PMID 19426029.

[Stevens1996-5] Stevens, George (1996). "Extending Rapoport's rule to Pacific marine fishes". Journal of Biogeography. 23 (2): 149–154. Bibcode:1996JBiog..23..149S. doi:10.1046/j.1365-2699.1996.00977.x.

[Rapoport;s_rule:_time_1996)-6] Gaston, Kevin J.; Blackburn, Tim M.; Spicer, John I. (1998). "Rapoport's rule: Time for an epitaph?". Trends in Ecology & Evolution. 13 (2): 70–74. Bibcode:1998TEcoE..13...70G. doi:10.1016/S0169-5347(97)01236-6. PMID 21238203.

[Rohde1992-7] Rohde, Klaus (1992). "Latitudinal Gradients in Species Diversity: The Search for the Primary Cause". Oikos. 65 (3): 514–527. Bibcode:1992Oikos..65..514R. doi:10.2307/3545569. JSTOR 3545569.

[Rohde1993-8] Rohde, Klaus; Heap, Maureen; Heap, David (1993). "Rapoport's Rule Does Not Apply to Marine Teleosts and Cannot Explain Latitudinal Gradients in Species Richness". The American Naturalist. 142 (1): 1–16. doi:10.1086/285526.

[9] Rohde, Klaus (1999). "Latitudinal gradients in species diversity and Rapoport's rule revisited: a review of recent work and what can parasites teach us about the causes of the gradients?". Ecography. 22 (6): 593–613. Bibcode:1999Ecogr..22..593R. doi:10.1111/j.1600-0587.1999.tb00509.x.

[Rohde1996-10] Rohde, Klaus (1996). "Rapoport's Rule is a Local Phenomenon and Cannot Explain Latitudinal Gradients in Species Diversity". Biodiversity Letters. 3 (1): 10–13. doi:10.2307/2999704. JSTOR 2999704.

[11] Stauffer, D.; Rohde, K. (2006). "Simulation of Rapoport's rule for latitudinal species spread". Theory in Biosciences. 125 (1): 55–65. arXiv:q-bio/0507033. Bibcode:2006ThBio.125...55S. doi:10.1016/j.thbio.2006.01.002. PMID 17046373.

[12] Brown, J. H. (1995). Macroecology. Chicago: University of Chicago Press.

[13] Letcher, Andy J.; Harvey, Paul H. (1994). "Variation in Geographical Range Size Among Mammals of the Palearctic". The American Naturalist. 144 (1): 30–42. doi:10.1086/285659.

[14] Fernández, Manuel Hernández; Vrba, Elisabeth S. (2005). "Rapoport effect and biomic specialization in African mammals: revisiting the climatic variability hypothesis". Journal of Biogeography. 32 (5): 903–918. Bibcode:2005JBiog..32..903F. doi:10.1111/j.1365-2699.2004.01188.x.

[15] Roy, K; Jablonski, D; Valentine, J W (1994). "Eastern Pacific molluscan provinces and latitudinal diversity gradient: no evidence for "Rapoport's rule"". Proceedings of the National Academy of Sciences. 91 (19): 8871–8874. Bibcode:1994PNAS...91.8871R. doi:10.1073/pnas.91.19.8871. PMC 44708. PMID 11607494.

[ColwellHurtt_1994-16] Colwell, Robert K.; Hurtt, George C. (1994). "Nonbiological Gradients in Species Richness and a Spurious Rapoport Effect". The American Naturalist. 144 (4): 570–595. doi:10.1086/285695.

[17] Rohde, Klaus (1998). "Latitudinal Gradients in Species Diversity. Area Matters, but How Much?". Oikos. 82 (1): 184–190. Bibcode:1998Oikos..82..184R. doi:10.2307/3546928. JSTOR 3546928.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

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

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 Green world hypothesis Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Resource Restoration
Producers	Autotrophs Chemosynthesis Chemotrophs Foundation species Kinetotrophs 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 Lithoautotroph Lithotrophy Marine 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	Lakes Rivers Soil Tritrophic interactions in plant defense Marine food webs cold seeps hydrothermal vents intertidal kelp forests North Pacific Gyre San Francisco Estuary tide pool
Processes	Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer–resource interactions 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 Anti-predator 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 Consumer-resource model Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey (Lotka–Volterra) equations Recruitment Small population size Stability Resilience Resistance Random generalized Lotka–Volterra model
Species	Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species / Native 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 Insular biogeography Land change modeling 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 Ontogenetic niche shift
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 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 Sexecology Systems ecology Urban ecology Theoretical ecology
Outline of ecology